25 downloads
533 Views
37MB Size
Report
This content was uploaded by our users and we assume good faith they have the permission to share this book. If you own the copyright to this book and it is wrongfully on our website, we offer a simple DMCA procedure to remove your content from our site. Start by pressing the button below!
Report copyright / DMCA form
7+( (&2/2*< 2) &<$12%$&7(5,$
7KH (FRORJ\ RI &\DQREDFWHULD 7KHLU 'LYHUVLW\LQ7LPHDQG6SDFH
(GLWHGE\
%5,$1 $ :+,7721 University of Durham, Durham, England DQG
0$/&2/0 32776 Virginia Tech., Blackburg, U.S.A.
./8:(5$&$'(0,&38%/,6+(56 1(:<25.%26721'25'5(&+7/21'21026&2:
H%RRN ,6%1 3ULQW,6%1
0-306-46855-7 0-7923-4735-8
.OXZHU$FDGHPLF3XEOLVKHUV 1HZ
KWWSZZZNOXZHURQOLQHFRP KWWSZZZHERRNVNOXZHURQOLQHFRP
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
YL
DAVID M. WARD 'HSDUWPHQW RI /DQG5HVRXUFHV DQG (QYLURQPHQWDO 6FLHQFH 0RQWDQD 6WDWH 8QLYHUVLW\ %R]HPDQ 86$ XPEGZ#PRQWDQDHGX BRIAN A. WHITTON 'HSDUWPHQW RI %LRORJLFDO 6FLHQFHV8QLYHUVLW\ RI 'XUKDP 'XUKDP '+ /( 8. EDZKLWWRQ#GXUKDPDFXN DAVIDD.WYNN-WILLIAMS %ULWLVK $QWDUFWLF 6XUYH\ +LJK &URVV 0DGLQJOH\ 5RDG &DPEULGJH&%2(78. GGZZ#SFPDLOQHUFEDVDFXN
YLL
&RQWHQWV /LVW RI $XWKRUV 3UHIDFH &RORXU 3ODWHV
Y [YLL [L[
,QWURGXFWLRQ WR WKH &\DQREDFWHULD %ULDQ $ :KLWWRQ DQG 0DOFROP 3RWWV 6XPPDU\ , :KDW DUH &\DQREDFWHULD" ,, (FRORJLFDO'LYHUVLW\LQWKH3DVWDQG3UHVHQW ,OO 0RUSKRORJLFDO 'LYHUVLW\ ,9 7D[RQRP\ 9 0ROHFXODU(FRORJ\ 9, ,QWHUDFWLRQV ZLWK 2WKHU 2UJDQLVPV 9,, %ORRPV DQG 7R[LQV 9,,, &\DQREDFWHULDDV+HDOWK)RRG ,; 8VHRI&XOWXUHVDQG&XOWXUH0HGLD ; 7KH)XWXUH 5HIHUHQFHV
7KH )RVVLO 5HFRUG 7UDFLQJ WKH 5RRWV RI WKH &\DQREDFWHULDO /LQHDJH - :LOOLDP 6FKRSI 6XPPDU\ , 7UDFLQJWKH5RRWVRI&\DQREDFWHULD3URJUHVVDQG3UREOHPV ,, $QFLHQW&\DQREDFWHULD ,OO +RZ 2OG LV WKH &\DQREDFWHULDO /LQHDJH" ,9 3DOHRELRORJ\)RVVLOV*HRORJ\DQG*HRFKHPLVWU\ 9 3DOHRELRORJ\)LQDO$UELWHURI&RPSHWLQJ7KHRULHV $FNQRZOHGJHPHQWV 5HIHUHQFHV
&\DQREDFWHULD LQ *HRWKHUPDO +DELWDWV 'DYLG 0 :DUG 5LFKDUG : &DVWHQKRO] 6XPPDU\ , ,QWURGXFWLRQ ,, 'LVWULEXWLRQRI7KHUPRSKLOLF&\DQREDFWHULD%DVHGRQ 0RUSKRORJ\DQG(QULFKPHQW&XOWXUH ,OO 'LVWULEXWLRQ RI &\DQREDFWHULD %DVHG RQ 0ROHFXODU$QDO\VLV ,9 3K\VLRORJLFDODQG%HKDYLRUDO(FRORJ\RI&\DQREDFWHULD RI*HRWKHUPDO+DELWDWV 9 &RQFOXVLRQ $FNQRZOHGJHPHQWV 5HIHUHQFHV
L[
-
&\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV
/XFDV - 6WDO 6XPPDU\ , ,QWURGXFWLRQ ,, 0LFURELDO 0DWV 6WURPDWROLWHV DQG WKHLU (QYLURQPHQWV ,OO 7KH2UJDQLVPV&\DQREDFWHULDWKDW%XLOG0LFURELDO0DWV ,9 0RWLOLW\&KHPR DQG3KRWRWD[LVRI&\DQREDFWHULDLQ0LFURELDO0DWV 9 &DUERQ 0HWDEROLVP 9, &DOFLILFDWLRQLQ0DWVDQG6WURPDWROLWHV 9,, 1LWURJHQ 0HWDEROLVP DQG 1LWURJHQ )L[DWLRQ 9,,, &\DQREDFWHULDDQGWKH6XOIXU&\FOHLQ0LFURELDO0DWV ,; ,QWHUDFWLRQVRI&\DQREDFWHULDZLWK,URQ ; 3KRVSKRUXVLQ0LFURELDO0DWV ;, &RQFOXVLRQV $FNQRZOHGJHPHQWV 5HIHUHQFHV
0DULQH 3ODQNWRQ
+DQV : 3DHUO 6XPPDU\ , ,QWURGXFWLRQ ,, .H\ )XQFWLRQDO *URXSV RI 0DULQH 3ODQNWRQLF &\DQREDFWHULD ,OO 3ODQNWRQLF &\DQREDFWHULDO +DELWDWV 3K\VLFDO &KHPLFDO DQG %LRWLF &RQVLGHUDWLRQV ,9 6\QWKHVLV 7KH (FRV\VWHP 3HUVSHFWLYH $FNQRZOHGJHPHQWV 5HIHUHQFHV
)UHVKZDWHU %ORRPV
5RGHULFN / 2OLYHUDQG *HRUJH * *DQI , ,QWURGXFWLRQ ,, %ORRP )RUPLQJ&\DQREDFWHULD ,OO 'LVWULEXWLRQ ,9 *DV9DFXROHV*DV9HVLFOHV%XR\DQF\DQGLWV5HJXODWLRQ 9 0L[LQJ 5HJLPHV DQG &\DQREDFWHULD 9, 3K\VLFDO &RQWURO RI&\DQREDFWHULD 9,, &HOO 6L]H *URZWK 5DWHDQG 7HPSHUDWXUH 9,,, /LJKW&DSWXUH ,; 1XWULHQWV ; 3KRVSKRUXV ;, 1LWURJHQ ;,, 5HVSRQVHVRI&\DQREDFWHULDWR1DQG3 ;,,, ,QRUJDQLF&DUERQ ;,9 *UD]LQJ ;9 &RQFOXGLQJ 5HPDUNV 5HIHUHQFHV
[
3LFRSODQNWRQ DQG 2WKHU 1RQ%ORRP )RUPLQJ &\DQREDFWHULD LQ /DNHV -RKQ * 6WRFNQHU&ULVWLDQD &DOOLHUL DQG *HUWUXG &URQEHUJ 6XPPDU\ , ,QWURGXFWLRQ ,, 6DPSOLQJ3UHVHUYDWLRQDQG(QXPHUDWLRQ ,OO 7KH1RQ%ORRP)RUPHUV:KDWDUH7KH\" ,9 $&RPPRQ(FRORJ\" 9 &RQFOXVLRQV $FNQRZOHGJHPHQWV 5HIHUHQFHV
6RLOVDQG 5LFH)LHOGV %ULDQ $ :KLWWRQ
6XPPDU\ , ,QWURGXFWLRQ ,, 6RLOV ,OO 6XEDHULDO +DELWDWV ,9 5LFHILHOGV 9 3UDFWLFDO0HWKRGV 9, &RQFOXGLQJ &RPPHQWV 5HIHUHQFHV
/LPHVWRQHV $OODQ3HQWHFRVWDQG %ULDQ $ :KLWWRQ 6XPPDU\ , ,QWURGXFWLRQ ,, 3K\VLFDODQG&KHPLFDO)HDWXUHV ,OO (QYLURQPHQWV DQG WKHLU &KDUDFWHULVWLF &RPPXQLWLHV ,9 7KH2UJDQLVPV 9 'HSRVLWLRQ 9, &RORQL]DWLRQ6XFFHVVLRQDQG:HDWKHULQJLQ7HUUHVWULDO DQG)UHVKZDWHU(QYLURQPHQWV 9,, ,QIOXHQFHRI7HPSRUDO&KDQJHVLQ1XWULHQWV 9,,, &RQFOXGLQJ&RPPHQWV $FNQRZOHGJHPHQWV 5HIHUHQFHV
[L
6DOWV DQG %ULQHV $KDURQ 2UHQ
6XPPDU\ , ,QWURGXFWLRQ ,, +\SHUVDOLQH(QYLURQPHQWVDQGWKHLU&\DQREDFWHULDO&RPPXQLWLHV ,OO 3K\VLRORJLFDO3URSHUWLHVRIWKH0DMRU+DORSKLOLF&\DQREDFWHULD ,9 $QR[\JHQLF3KRWRV\QWKHVLVE\&\DQREDFWHULDLQ +\SHUVDOLQH(QYLURQPHQWV 9 2VPRWLF$GDSWDWLRQRI&\DQREDFWHULD/LYLQJDW +LJK6DOW&RQFHQWUDWLRQV 9, ,QWHUDFWLRQV%HWZHHQ&\DQREDFWHULDDQGRWKHU0LFURRUJDQLVPV LQ+\SHUVDOLQH(QYLURQPHQWV 9,, %LRWHFKQRORJLFDO$VSHFWVRI+DORSKLOLF&\DQREDFWHULD 9,,, &RQFOXVLRQV $FNQRZOHGJHPHQWV 5HIHUHQFHV
2LO 3ROOXWLRQ DQG &\DQREDFWHULD 6DPLU6 5DGZDQDQG5HGKD + $O+DVDQ 6XPPDU\ , ,QWURGXFWLRQ ,, &RPSRVLWLRQRI&UXGH2LO ,OO %LRGHJUDGDELOLW\RI+\GURFDUERQV ,9 &\DQREDFWHULDLQWKH%LRJHQHVLVRI2LO 9 +\GURFDUERQ'HJUDGDWLRQ3RWHQWLDORI&\DQREDFWHULD 9, &RQFOXGLQJ5HPDUNV $FNQRZOHGJHPHQWV 5HIHUHQFHV
&\DQREDFWHULDO 'RPLQDQFH LQ WKH 3RODU 5HJLRQV :DUZLFN ) 9LQFHQW 6XPPDU\ , ,QWURGXFWLRQ ,, +DELWDWVDQG&RPPXQLWLHV ,,, %LRGLYHUVLW\DQG(QGHPLVP ,9 (FRSK\VLRORJ\RI3RODU&\DQREDFWHULD 9 :K\'R&\DQREDFWHULD'RPLQDWH2U1RW " $FNQRZOHGJHPHQWV 5HIHUHQFHV
[LL
&\DQREDFWHULDLQ'HVHUWV /LIHDWWKH /LPLW" '':\QQ:LOOLDPV 6XPPDU\ , $+LVWRULFDO3HUVSHFWLYH ,, 'LYHUVLW\RI'HVHUW(FRV\VWHPV ,OO 7KH2UJDQLVPV %LRGLYHUVLW\ ,9 'HVHUW1LFKHVDQG,QWHUDFWLRQV 9 6WUHVV)DFWRUVLQ$ULG5HJLRQV 9, %LRZHDWKHULQJDQG1XWULHQW$YDLODELOLW\ 9,, 7KUHVKROGVDQG([WLQFWLRQ $FNQRZOHGJHPHQWV 5HIHUHQFHV
'HWHFWLQJWKH (QYLURQPHQW 1LFKRODV+ 0DQQ
6XPPDU\ , ,QWURGXFWLRQ ,, 3URWHLQ3KRVSKRU\ODWLRQ ,OO /LJKW'DUNDQG5HGR[6HQVLQJ ,9 /RZ0ROHFXODU:HLJKW6LJQDOOLQJ0ROHFXOHV 9 %HKDYLRXUDO5HVSRQVHV 9, $OWHUDWLRQVLQ7UDQVFULSWLRQDO6SHFLILFLW\ 9,, 2WKHU6LJQDOOLQJ3URFHVVHV 9OOO &URVV7DONDQG,QWHJUDWLRQ ,; &RQFOXVLRQV $FNQRZOHGJHPHQWV 5HIHUHQFHV
0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO6WUHVV 'HYDNL %KD\D5DNHIHW6FKZDU]DQG$UWKXU5*URVVPDQ 6XPPDU\ , ,QWURGXFWLRQ ,, $FFOLPDWLRQWR/LJKW ,OO 5HVSRQVHVWR1XWULHQW/LPLWDWLRQ ,9 &RQFOXGLQJ5HPDUNV $FNQRZOHGJHPHQWV 5HIHUHQFHV
0HWDO 0HWDEROLVP DQG7R[LFLW\ 5HSHWLWLYH'1$ 1LJHO-5RELQVRQ -XOLDQ&5XWKHUIRUG0DWKHZ5 3RFRFN DQG-HQQLIHU6 &DYHW 6XPPDU\ , 0HWDO0HWDEROLVPDQG7R[LFLW\ ,, 5HSHWLWLYH'1$LQ&\DQREDFWHULD ,OO &RQFOXGLQJ5HPDUNV $FNQRZOHGJHPHQWV 5HIHUHQFHV
[LLL
1RVWRF
0DOFROP 3RWWV 6XPPDU\ , ,QWURGXFWLRQ ,, 1DWXUDO&RPPXQLWLHV ,,, (YROXWLRQ ,9 3K\VLRORJ\DQG*URZWK 9 3DWHQWVDQG$SSOLFDWLRQV 9, $&KURQRORJ\RI1RVWRF 9,, (SLORJXH $FNQRZOHGJHPHQWV 5HIHUHQFHV
$UWKURVSLUD 6SLUXOLQD 6\VWHPDWLFV DQG (FRSK\VLRORJ\ $YLJDG 9RQVKDN DQG/XLVD7RPDVHOOL 6XPPDU\ , ,QWURGXFWLRQ ,, 0RUSKRORJ\ ,OO 6\VWHPDWLFV ,9 2FFXUUHQFHDQG'LVWULEXWLRQ 9 3K\VLRORJ\RI$UWKURVSLUD 9, &RQFOXGLQJ5HPDUNV 5HIHUHQFHV
6\PELRWLF,QWHUDFWLRQV 'DYLG*$GDPV 6XPPDU\ , ,QWURGXFWLRQ ,, 7KH6\PELRVHVDQGWKHLU(QYLURQPHQWDO,PSDFW Ill. 7KH6\PELRQWV ,9 +RVW&\DQREDFWHULD,QWHUDFWLRQV3ULRUWR,QIHFWLRQ 9 +RVW6WUXFWXUHVDQGWKHLU,QIHFWLRQ 9, +RVW&\DQRELRQW,QWHUDFWLRQV3RVW,QIHFWLRQ 9,, 5HFRQVWLWXWLRQRIWKH6\PELRVHV 9,,, &RQFOXGLQJ5HPDUNV $FNQRZOHGJHPHQWV 5HIHUHQFHV
[LY
&\DQRSKDJHV DQG 7KHLU 5ROH LQ WKH (FRORJ\ RI &\DQREDFWHULD &XUWLV$ 6XWWOH 6XPPDU\ , ,QWURGXFWLRQ ,, 7D[RQRP\0RUSKRORJ\DQG(YROXWLRQRI&\DQRSKDJHV ,OO 'LYHUVLW\ ,9 'LVWULEXWLRQ$EXQGDQFHDQG6HDVRQDO'\QDPLFV 9 )DWHRI&\DQRSKDJHVLQWKH1DWXUDO(QYLURQPHQW 9, (IIHFWRI&\DQRSKDJHVRQ&\DQREDFWHULDO3RSXODWLRQV DQG &RPPXQLWLHV 9,, (QYLURQPHQWDODQG3K\VLRORJLFDO(IIHFWVRQ&\DQRSKDJHV 9,,, /\VRJHQ\ ,; &RQFOXVLRQ $FNQRZOHGJHPHQWV 5HIHUHQFHV
&\DQREDFWHULDO 5HVSRQVHV WR 895DGLDWLRQ 5LFKDUG : &DVWHQKRO] DQG)HUUDQ *DUFLD3LFKHO 6XPPDU\ , ,QWURGXFWLRQ ,, 6WUDWHJLHVRI8957ROHUDQFHE\&\DQREDFWHULD ,OO (IIHFWVRI895LQ1DWXUHDQG:KROH&RPPXQLW\5HVSRQVHV ,9 &RQFOXVLRQV $FNQRZOHGJHPHQWV 5HIHUHQFHV
&\DQRWR[LQV &UDZIRUG 6 'RZ DQG 8WKD\D . 6ZRERGD
6XPPDU\ , ,QWURGXFWLRQ ,, 3RLVRQLQJ,QFLGHQWV ,OO :KLFK7D[DDUH7R[LF" ,9 7\SHVRI&\DQREDFWHULDO7R[LQV 9 7R[LQ$QDO\VLV 9, (FRORJLFDO,PSOLFDWLRQV 9,,&RQFOXGLQJ&RPPHQWV 5HIHUHQFHV
2UJDQLVP LQGH[
*HQH DQG *HQH 3URGXFW ,QGH[
6XEMHFW ,QGH[
XV
3UHIDFH
7KH F\DQREDFWHULD KDYH ILJXUHG SURPLQHQWO\ LQ GLVFXVVLRQV EHWZHHQ WKH HGLWRUV RYHU WKH \HDUV LQ GLIIHUHQW FRXQWULHV DQG GLYHUVH VHWWLQJV VRPH PRUH H[RWLF WKDQ RWKHUV :H KDYH OHDUQHG WKDW 1DWXUH JLYHV XS KHU VHFUHWV ZLWK FRQVLGHUDEOH UHOXFWDQFH 0DQ\ GHWDLOV DQG DVSHFWV RI F\DQREDFWHULDO HFRORJ\ UHPDLQ HOXVLYH 2XU LQWHQWLRQ LQ DVVHPEOLQJ WKLV ERRN ZDV WR KHOS DOO WKRVH WU\LQJ WR XQUDYHO WKHLU VHFUHWV DQG WR UHYHDO ZKDW LV NQRZQ WR D ZLGH DXGLHQFH 7KH SXEOLFDWLRQ LQ RI The Molecular Biology of Cyanobacteria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he comparative physiology and biochemistry of the bluegreen algae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
[YLL
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ractical Uses and Problems of Cyanobacteria. $WWKHWLPHRIZULWLQJWKLVWKHVHTXHQFLQJRIRQH F\DQREDFWHULDOJHQRPHLVFRPSOHWHGDQGWKUHHRWKHUV DUH LQSURJUHVV 2QH WKDWRI Nostoc, LV WKH ODUJHVW
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p 7RXPRLV ZDV LQFKDUJHRISURGXFWLRQ %5,$1 :+,7721 0$/&2/032776
[YLLL
&RORXU 3ODWHV
Plate 5 5 a, b, c +XQWHU V +RW 6SULQJV 2UHJRQ a 2QH RI !& VRXUFHV VKRZLQJ WKH ³9´ RI FRROLQJ WR a & ZLWK JUHHQ Synechococcus VSS IROORZHG DW a & E\ UHGGLVKEURZQ Oscillatoria FI terebriformis PDW LQ VRPH DUHDV WKLV PDW KDV UHWUDFWHG H[SRVLQJ XQGHUPDW RI Chloroflexus; E (GJH RI SRRO ZLWK WKLFN PDW RI Oscillatoria ZKHUH LW UHSODFHV JUHHQ Synechococcus PDW DW a&RVWUDFRGV Potamocypris VS ZKLWH REMHFWV WUXH VL]H aPP ORQJ KDYH JUD]HG Oscillatoria EHORZ a & c 2VWUDFRGV JUD]LQJ RQ JUD]HUUHVLVWDQW JUD]HUGHSHQGHQW Pleurocapsa/Calothrix FRPPXQLW\ JOLVWHQLQJ VSRW LQ HDFK DQLPDO LV WKH H\H d, e, f
10 &\DQREDFWHULD LQ WKH VHD a, b Trichodesmium thiebautii IURPEORRPVLQWKHZHVWHUQ$WODQWLF2FHDQ a UDGLDO
SXII DJJUHJDWH b IXVLIRUPWXIWDJJUHJDWHVIURPVLWHNPRIIFRDVWRI 1RUWK &DUROLQD c &RQWRXU SORWRIVSDWLR WHPSRUDOGLVWULEXWLRQRISODQNWRQEORRPVDORQJIUHVKZDWHUNP WR HVWXDULQH NP VDOLQLW\JUDGLHQW LQ 1HXVH 5LYHU (VWXDU\ 1 &DUROLQD 'DWD EDVHG RQ ELZHHNO\ VDPSOLQJDWVWDWLRQV FDURWHQRLGSLJPHQWV TXDQWLILHGE\ +3/& d 0DULQH6\QHFKRFRFFXVWDJJHG ZLWK IOXRUHVFHQWO\ODEHOOHGYLUXVHV Synechococcus VWUDLQ%%&KDVEHHQDGGHGWR D QDWXUDO PDULQH PLFURELDO FRPPXQLW\DQGWDJJHGZLWKF\DQRSKDJHVODEHOOHGZLWKWKHEOXHIOXRUHVFHQWVWUDLQ3232 7KH\DSSHDUSLQNVXUURXQGHGE\D EOXHKDOR 8QWDJJHG Synechococcus VSSIOXRUHVFHUHGRU \HOORZZKLOHD KHWHURWULRSKLFEDFWHULXPKDV EHHQ WDJJHGZLWKDYLUXVWKDWKV EHHQ ODEHOOHG ZLWKDJUHHQIOXRUHVFHQWVWDLQ<2<2 3KRWRV a,b,c+:3DHUOd, &$ 6XWWOH
Plate 10
Plate 21 &\DQREDFWHULDRI%HDFRQ 6DQGVWRQH LQ WKH$QWDUFWLF a 9HUWLFDO IUDFWXUHSURILOHRIFU\SWRHQGROLWKLFFRPPXQLW\ VKRZLQJ DGLVWLQFWF\DQREDFWHULDO]RQH a 6 PPIURPWKHVXUIDFH RIWKHURFN EODFN ]RQHQHDUHUWKH VXUIDFH FRQWDLQV SLJPHQWHG OLFKHQ ZKLOH WKH ZKLWH ]RQH FRPSULVHVPDLQO\ K\DOLQH IXQJL ZLWK VLRPH PLFURDOJDH b $UWLILFLDOO\ H[SRVHG F\DQREDFWHULD JUHHQEURZQ FRORXUDWLRQ K\DOLQH IXQJL ZKLWH EODFNSLJPHQWHG OLFKHQ QHDUHVW WKH VXUIDFH RI DQ H[IROLDWLQJ FU\SWRHQGROLWKLF FRPPXQLW\DW 7LPEHU 3HDN 3ULHVWOH\*ODFLHU FHQWUDO 9LFWRULD /DQG ZKLWH ODEHO LV FP ZLGH 3KRWRV '' :\QQ:LOOLDPV
Plate 22 22 &\DQREDFWHULD LQ KRW GHVHUWV a 5RFN\ GHVHUW 1HJHY b 'HVHUW SDYHPHQW1HJHY c 'HVHUW SDYHPHQWFORVHXS d 'HVHUWSDYHPHQWZLWKRQHVWRQH RYHUWXUQHG WRUHYHDOK\SROLWKLF F\DQREDFWHULD RI VWRQHVRLOLQWHUIDFH e 6XUIDFH RI1XELDQVDQGVWRQHZLWK SDUWV FKLSSHG RII WR UHYHDO FU\SWRHQGROLWKLF F\DQREDFWHULDOFRORQL]DWLRQXQGHUWKHVXUIDFH 1HJHY f &KDVPRHQGROLWKLF FRORQL]DWLRQ LQ JUDQLWH VKRZQ LQWKLQ VHFWLRQ RIURFN 6RQRUDQ'HVHUW 0H[LFR 3KRWRV(,)ULHGPDQQ
Plate 23 23 2OG OHDG ]LQF PLQLQJ DUHDV KLJKO\ FRQWDPLQDWHGZLWK=Q&G DQG 3E a-e 0LVVRXUL 86$ f (LQHVEHUJHU =HQWUDOH +DU] *HUPDQ\ a (OYLQV7DLOLQJV ODJRRQV PJ / =Q ZLWKSODQNWRQLFSynechococcus. b 'UDLQDJH VWUHDPPJ / =Q DWVDPHVLWHVKRZLQJEURZQFRORXUGXHWRPlectonema VKHDWKV c &ORVHXS ZLWKSDUWRIPDW VHFWLRQHG FRPPXQLW\ LV VLPLODU WR WKDW DW PDQ\ RWKHU FRQWDPLQDWHG VLWHV PL[WXUH RIQDUURZ Plectonema DQG SURWRQHPDO VWDJH RIWKHPRVV Dicranella. d Schizothrix FRPPXQLW\ ZLWK UHGGLVK VKHDWK W\SLFDO RIFRQWDPLQDWHG DOWHUQDWHO\ ZHW GU\DUHDV e )ORDWLQJPDWVRI PL[HGSRSXODWLRQVRIQDUURZVKHDWKHG2VFLOODWRULDFHDHW\SLFDO RI DUHDVZLWKVWDQGLQJ ZDWHU f 3DUWRIROGVSRLOKHDSRQO\ SDUWLDOO\FRORQL]HGE\KLJKHUSODQWVDQGVKRZLQJ GDUN FRYHURIPL[HG Plectonema DQGPRVVSURWRQHPDZKLFKLVDSSDUHQWO\ WKHW\SLFDOWHUUHVWULDOSLRQHHUFRPPXQLW\ RI VXFK FRQWDPLQDWHG DUHDV ZRUOGZLGH 3KRWRV %$ :KLWWRQ
Plate 24 24 Nostoc. a N. pruniforme. b 6ROXWLRQ KROORZ RQ OLPHVWRQH ZLWK PRLVW Nostoc commune FRORQLHV DQG DVVRFLDWHG RUJDQLFGHEULVSHUPLWWLQJ JURZWKRIKLJKHU SODQW (Portulaca) $OGDEUD$WROO VLPLODUVROXWLRQKROORZV ZLWK N. commune KDYH EHHQ UHSRUWHGIURPPDQ\FRXQWULHV c Nostoc FRORQLHVSHUVLVWLQJDWERWWRPRIHPSW\ SRRO GXULQJWKHGU\VHDVRQ RQ $OGDEUD $WROOWKLV FUXPEO\ IRUP WDNHVXSDQGORVHVZDWHUPRUHVORZO\WKDQWKHW\SLFDO IRUP d 6SKHULFDOFRORQLHVSHDUOV RIN. commune JURZLQJ RQ DJDU VXSSOHPHQWHG ZLWK FDOFLXPDQGPDJQHVLXP FDUERQDWHV e 1RPDUVNLLQWHUIHUHQFH PLFURVFRS\ RIWKLFN VHFWLRQRI N. commune WKDOOXV VKRZLQJ EURZQ DQG \HOORZ OD\HUV GXH WR VF\WRQHPLQLQGLYLGXDO ILODPHQWV YLVLEOH WKURXJKRXW WKH H[WUDFHOOXODU JO\FDQ 3KRWRVD%$ :KLWWRQ 5:&DVWHQKRO] b,c %$:KLWWRQ d,e 03RWWV
Plate 25 25 Scytonema a &RORQ\ RQ XSSHU SDUW RI PDQJURYH SQHXPDWRSKRUH QHDU (LODW ,VUDHO b, F 0DWV LQ XSSHU LQWHUWLGDO LQODQG UHJLRQV RI$QGURV ,V %DKDPDV b SRO\JRQV FRQVLVW RI ODPLQDWHG VWUXFWXUH WUDSSLQJ SDUWLFOHV c PDWV GHYHORSLQJ RQ VXUIDFH LQ VKHOWHUHG DUHD d 0DWV LQ \RXQJ WUDQVSODQWHG ULFHILHOG 0DQLNJDQM %DQJODGHVK e +HPL VSKHULFDO FRORQLHV RI S. myochrous, D FKDUDFWHULVWLF VSHFLHV RI PRLVW WUDYHUWLQHGHSRVLWLQJ UHJLRQV UHSRUWHG IURP PDQ\ FRXQWULHV VHHDOVR3ODWH K 3KRWRV a 0 3RWWV b,c %$ :KLWWRQ d,e ' /LYLQJVWRQH
Plate 26 26 5LYXODULDFHDHDUHFKDUDFWHULVWLFRI VLWHVVKRZLQJPDUNHGWHPSRUDORUVSDWLDOYDULDWLRQLQDPELHQWSKRVSKDWH a 6WUHDP 8SSHU 7HHVGDOH (QJODQGZLQWHU GUDLQLQJUHJLRQZLWK OLPHVWRQH DQGSHDW DQGZLWK H[WHQVLYH Rivularia FRYHU WKURXJKRXW WKH\HDU b Gloeotrichia natans FRORQLHV%DQJODGHVK c 2QFRLGVW\SLFDOO\ FP IRUPHG E\VPDOO Rivularia FRORQLHV LQ FDOFDUHRXVIOXVK6XQELJJLQ(QJODQG d 6HFWLRQ RI RQFRLGIURPWKLV VLWH e, f 0DULQH LQWHUWLGDO Rivularia (R. atra): e, VHFWLRQ RI W\SLFDO FRORQ\ VKRZLQJ RXWHU VF\WRQHPLQULFK OD\HU DQG LQWUDFHOOXODU ORFDWLRQ RI KHWHURF\VWV f KRUPRJRQLDIRUPDWLRQIROORZLQJ H[SRVXUHWRSXOVH RI HOHYDWHGSKRVSKDWH 3KRWRV a,b,c,e,f %$ :KLWWRQ d *$/ -RKQVRQ
Plate 27 27 6SLUXOLQD Arthrospira a, b 2XWGRRUSRQGV DW(DUWKULVH)DUPV &DOLIRUQLD c 3URFHVVLQJ 6SLUXOLQD DW (DUWKULVH )DUPV d 1DWXUDO SRSXODWLRQ RI Arthrospira IURP $IULFDQ QDWURQ ZDWHU EDU PDUNHU P e 7ULFKRPH VKRZLQJ JDV YDFXROHV 3KRWRVa,b,c (DUWKULVH)DUPVd & 6LOL e 0 0LLKOLQJ
Plate 28 28 6\PELRWLF DVVRFLDWLRQV PDFURVFRSLFYLHZV a Gunnera chilensis, LQFOXGLQJDUHDDW EDVH RI OHDYHV FRQWDLQLQJ VWHPJODQGVLQIHFWHGZLWKNostoc. b &\FDGVDVDQ XQGHUVWRUH\ LQ D QHZO\EXUQWEucalyptus IRUHVW LQ $XVWUDOLD c 6SRURSK\WH RI WKHIHUQAzolla pinnata YLHZHG IURP DERYHWKHNostoc V\PELRQWLVFRQWDLQHGZLWKLQVZROOHQGRUVDO OREHV RI WKHOHDYHVUDLVHG DERYHWKHZDWHUVXUIDFH d 7KH OLYHUZRUW Blasia pusilla VKRZLQJGDUNNostoc FRORQLHV VFDWWHUHGDURXQGWKHSURPLQHQWPLGULE e 7KH WULSDUWLWHOLFKHQPeltigera aphthosa FRPSRVHG RI DIXQJXVVHHQDV ZKLWH DUHDV DWWKH FXUOHGHGJHV DJUHHQ DOJD FRORXU RIPDLQWKDOOXV DQG Nostoc FRQWDLQHG LQ VPDOO GDUNVSRWV FHSKDORGLD VFDWWHUHG RYHU DWKDOOXVW\SLFDOO\ FP LQ GLDPHWHU f 7KHIXQJXV Geosiphon pyriforme LVFRORXUHG E\LWVHQGRV\PELRWLF Nostoc. 3KRWRV a,c,d '*$GDPV b,e % %HUJPDQ f '0ROOHQKDXHU
Plate 29 29 6\PELRWLF DVVRFLDWLRQV PLFURVFRS\ a c )OXRUHVFHQFH VWXGLHV ZLWK FDOFRIOXRU ZKLWH VWDLQLQJ a 9HQWUDO VXUIDFH RI Pkaeoceros, VKRZLQJ WKUHH VWRPDWDOLNH HQWUDQFHV WKURXJK ZKLFK LQIHFWLQJ KRUPRJRQLD JDLQ HQWU\ WR VOLPH FDYLWLHV ZLWKLQ WKH WKDOOXV b 9HQWUDO VXUIDFH RI Blasia, VKRZLQJ D VLQJOH DXULFOH DV DQ DOPRVW VSKHULFDO VWUXFWXUH RQ VXUIDFH RI WKDOOXV D VOLPH SDSLOOD DOPRVW ILOOV WKH DXULFOH H[FHSW IRU D QDUURZ VSDFH WKDW DSSHDUV DV D GDUNHU XQVWDLQHG UHJLRQ WKLV VSDFH EHFRPHV LQIHFWHG E\ F\DQREDFWHULD DQG WKH DXULFOH HQODUJHV DV WKH V\PELRWLF VWUXFWXUH GHYHORSV c Blasia, VKRZLQJ WKH UHG DXWRIOXRUHVFHQFH RI Nostoc ZLWKLQ DQ HQODUJHG DXULFOH d Blasia DXULFOH VKRZLQJ QHZO\ GHYHORSHG Nostoc FRORQ\ VXUURXQGLQJ WKH FHQWUDO VOLPH SDSLOOD e Blasia LQIHFWHG LQ ODERUDWRU\ ZLWK WZR GLIIHUHQW Nostoc VWUDLQV RQH EOXHJUHHQGXH WR FKORURSK\OO DQG SK\FRF\DQLQ DQG WKH RWKHU EURZQ GXH WR WKH DGGLWLRQDO SK\FRHU\WKULQ WKH FRORXUV RI WKH V\PELRWLF FRORQLHV FDQ EH VHHQ WKURXJK WKH DXULFOH ZDOOV f Nostoc LVRODWHG IURP Peltigera apktkosa; KHWHURF\VWRXV WULFKRPHV RFFXU LQ SDFNDJHV VXUURXQGHG E\ D SURQRXQFHG PXFLODJLQRXV VKHDWK 3KRWRV a,b,c,d, f 6 %DELF e % %HUJPDQ
Plate 30 30 a 6F\WRQHPLQULFKPDWRICalothrix LQDWHSLGKRWVSULQJVWUHDP3RWWV%DVLQ:HVW7KXPERI
Plate 31 31 a 6F\WRQHPLQULFK PDW PDLQO\ Lyngbya FI aestuarii, LQ K\SHUVDOLQH LQWHUWLGDO IODWV /DJXQD *XHUUHUR 1HJUR %DMD &DOLIRUQLD 6 0H[LFR b 'HVHUW FUXVW LQ 0RDE 8WDK VKRZLQJ H[SHULPHQWDO DUHD ZLWK GDUN FRORUDWLRQ GXH WR WKH KLJK DPRXQWV RI VF\WRQHPLQ LQ Nostoc commune, Scytonema VSS DQG WKH F\DQROLFKHQ Collema cristata; QRWH EDUH VRLOV LQ TXDGUDWV ZKHUH FUXVWV KDYH EHHQ UHPRYHG DQG LQ WKH ZDONLQJ SDWKV F d, e 7KH F\DQROLFKHQ Collema FI coccophorum $UFKHV 1DWLRQDO 3DUN 8WDK FRQWDLQV KLJK DPRXQWV RI VF\WRQHPLQ V\QWKHVL]HG E\ Nostoc LQ LWV XSSHU OREH VR WKLV UHJLRQ GRHV QRW IOXRUHVFH GXH WR WKH 89$ VKLHOGLQJ E\ VF\WRQHPLQ OLFKHQ OREH LV DERXW PP LQ GLDPHWHU c /REH VKRZQ ZLWK QDWXUDO FRORUDWLRQ d $XWRIOXRUHVHFHQFH ZKHQ H[FLWHG ZLWK JUHHQ OLJKW e $XWRIOXRUHVFHQFH ZKHQ H[FLWHG ZLWK 89$ UDGLDWLRQ 3KRWRV a 5:&DVWHQKRO] b d ) *DUFLD3LFKHO
&KDSWHU ,QWURGXFWLRQ WR WKH &\DQREDFWHULD %ULDQ $ :KLWWRQ DQG 0DOFROP 3RWWV 'HSDUWPHQW RI %LRORJLFDO 6FLHQFHV 8QLYHUVLW\ RI 'XUKDP 'XUKDP '+ /( 'HSDUWPHQW RI %LRFKHPLVWU\ 9LUJLQLD 3RO\WHFKQLF ,QVWLWXWHDQG6WDWH8QLYHUVLW\ %ODFNVEXUJ 9$ 86$
6XPPDU\ , :KDW DUH &\DQREDFWHULD" ,, (FRORJLFDO 'LYHUVLW\LQWKH3DVW DQG 3UHVHQW ,OO 0RUSKRORJLFDO 'LYHUVLW\ ,9 7D[RQRP\ 9 0ROHFXODU (FRORJ\ 9, ,QWHUDFWLRQVZLWK2WKHU2UJDQLVPV 9,, %ORRPV DQG7R[LQV 9,,, &\DQREDFWHULD DV +HDOWK )RRG ,; 8VH RI &XOWXUHV DQG&XOWXUH0HGLD ; 7KH )XWXUH
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bFRQWDLQLQJ RUJDQLVPVWKH\DUHWUHDWHG KHUH&KDSWHU DVPHPEHUVRI WKHF\DQREDFWHULDLQ WKHEURDGHVWVHQVHEXWVHHS
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rochlorothrix, ZKLFK GR QRW
,, (FRORJLFDO'LYHUVLW\LQ WKH 3DVWDQG 3UHVHQW *HRORJLVWV DQG JHRFKHPLVWV DOO DJUHH WKDW F\DQREDFWHULDKDYH KDG D ORQJ HYROXWLRQDU\ KLVWRU\ DQG PRVW WKRXJKQRWDOO DJUHHWKDWWKLVH[WHQGVWR
B. A. Whitton and M. Potts (eds), The Ecology of Cyanobacteria, 1-11. 2000 Kluwer Academic Publishers. Printed in the Netherlands.
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hormidium hendersonii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
%$:KLWWRQDQG03RWWV UHDFKHG E\ Synechococcus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hroococcidiopsis, VRPHWLPHVIRUPSDUWRI DPLFURELDOFRPPXQLW\ VHYHUDOPLOOLPHWUHVEHORZ WKH VXUIDFH &KDSWHU :\QQ:LOOLDPV 7KHVH RUJDQLVPVDUHFOHDUO\JURZLQJQHDUWKHOLPLWVIRUOLIH LQ WKH GU\ GHVHUWV RI $QWDUFWLFD DQG LW KDV EHHQ VXJJHVWHGWKDW WKHLUGRXEOLQJWLPHPD\ EH DV VORZ DV WHQ WKRXVDQG \HDUV 1LHQRZ DQG )ULHGPDQQ 7KLV GLIIHUV E\ DIDFWRU RI [ IURP WKH IDVWHVW NQRZQ JURZWKUDWHIRUDF\DQREDFWHULXPDGRXEOLQJ WLPH RI K IRU "Anacystis nidulans"
&KDSWHU ,QWURGXFWLRQWRWKH&\DQREDFWHULD (Synechococcus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
ORZHU S+ YDOXHV DUH KHWHURF\VWRXV VR SHUKDSV WKH\ DUH RQO\ DEOH WR FRPSHWH HIIHFWLYHO\ LQ DQ HQYLURQPHQW ZKHUH WKH DELOLW\ WR IL[ QLWURJHQ LV LPSRUWDQW
,,, 0RUSKRORJLFDO 'LYHUVLW\ &\DQREDFWHULD VKRZ FRQVLGHUDEOH PRUSKRORJLFDO GLYHUVLW\ 7KH\ PD\ EH XQLFHOOV HJ Chroococcus: 3ODWHV D E RU ILODPHQWV HJ Anabaena: 3ODWH I DQG WKHVH PD\ RFFXU VLQJO\ RU JURXSHG LQ FRORQLHV 3ODWH Schizothrix: 3ODWH H 8QLFHOOV PD\ GLYLGH LQ RQH WZR (Merismopedia: 3ODWH E RU WKUHH SODQHV 6RPH XQLFHOOV GLYLGLQJ LQ RQH SODQH VKRZ DV\PPHWULF GLYLVLRQ LQ Chamaesiphon WKH VPDOOHU FHOOV JOLGH GRZQ WR WKH EDVH RI WKH ODUJHU FHOO DQG UHSHDWV RI WKLV SURFHVV HYHQWXDOO\ JLYH ULVH WR D FRORQ\ RI PDQ\ FHOOV 7KH UHVXOWLQJ FRORQLHV FDQ RIWHQ EH VHHQ DV EURZQLVK VSRWV RQ VXEPHUJHG URFNV HJ & fuscus: 3ODWH OJ %UDQFKLQJ RI ILODPHQWV PD\ EH IDOVH RU WUXH )DOVH EUDQFKLQJ LV ZKHQ GLYLVLRQ LQ D ILODPHQW ZLWK D VKHDWK OHDGV WR WKH IRUPDWLRQ RIWZR VHSDUDWH VKHDWKHG ILODPHQWV DQG RQH RI WKHVH JURZV VHSDUDWHO\ IURP WKH RWKHU Tolypothrix: 3ODWH J Scytonema: 3ODWH K 7UXH EUDQFKLQJ RFFXUV ZKHQ FHOOV LQ D ILODPHQW KDYH WKH SRWHQWLDO IRU GLYLGLQJ LQ PRUH RQH SODQH HJ Hapalosiphon 0DLQWHQDQFH RI WKH FRORQLDO VWUXFWXUH LQ PDQ\ RI WKHVH IRUPV LV DLGHG E\ WKH SUHVHQFH RI H[RSRO\VDFFKDULGHV &KDSWHU 3RWWV VXFK DV PXFLODJH DQGRU D ILUP VKHDWK 7KH SUHVHQFH RU DEVHQFH RI D KHWHURF\VW LV DQ LPSRUWDQW IHDWXUH VHSDUDWLQJ JHQHUD +RZHYHU JDVYDFXROHV &KDSWHU 2OLYHU DQG *DQI VWUXFWXUHV ZKLFK DLG EXR\DQF\ DUH IRXQG LQ VSHFLHV RI PDQ\ GLIIHUHQW JHQHUD HJ Anabaena: 3ODWH I 6HYHUDO RWKHU WHUPV DUH XVHIXO IRU LQWHUSUHWLQJ WKH PRUSKRORJ\ RI F\DQREDFWHULD 7KH WULFKRPH UHIHUV WR WKH FHOOXODU SDUW RI D ILODPHQW ZLWK D VKHDWK HJ Lyngbya: 3ODWH D HYHQ WKRXJK D VLPLODU PRUSKRORJLFDO IRUP RFFXUULQJ ZLWKRXW D VKHDWK LV RIWHQ WHUPHG D ILODPHQW DV LQ Oscillatoria. ,Q VRPH IDOVHEUDQFKHG ILODPHQWRXV IRUPV WKH ILODPHQW JURZV DSLFDOO\ DQG GLHV DW WKH EDVH :KHUH WKH VKHDWK LV SHUVLVWHQW WKLV FDQ OHDG WR WKH GHYHORSPHQW RI D PDVV RI ROG VKHDWKV XQGHU WKH OLYLQJ PDWHULDO DV LQ VRPH VSHFLHV RI Scytonema. $QRWKHU ZLGHVSUHDG PRUSKRORJLFDO IRUP LV ZKHUH FHOO GLYLVLRQ LV ORFDOL]HG QHDUHU WKH EDVH RI D VKHDWKHG WULFKRPH DQG WKH DSLFDO
FHOOV EHFRPH QDUURZHG DQG GHYHORS LQWR D PXOWLFHOOXODU KDLU W\SLFDOO\ WKH FHOOV ORVH WKHLU FKORURSK\OO DQG WKH WK\ODNRLGV VSOLW DSDUW WR IRUP OLTXLGILOOHG YDFXROHV HJ Calothrix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ivularia DUH W\SLFDOO\ IRUPHG E\ WKH DJJUHJDWLRQ RI D QXPEHU RI KRUPRJRQLD ZKHUHDV RWKHUV HJ Nostoc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
%$ :KLWWRQ DQG03RWWV Chroococcus turgidus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ynechococcus DQG Synechocystis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scillatoria, ZLWK SHUKDSV PRUHWKDQD KXQGUHG FHOOV PLJKW FRQWDLQ PDQ\ KXQGUHGV RI FRSLHV RI WKH JHQRPH ,W VKRXOG WKHUHIRUH EH QR VXUSUL]HZKHQ FORQDOLVRODWHVRI DILODPHQWVRPHWLPHV JLYHULVHWR SKHQRW\SLFDOO\GLYHUVHILODPHQWVZLWKLQ D VLQJOHFXOWXUHIODVN 7KHJHQHUDODFFHSWDQFHRI WKHF\DQREDFWHULDORULJLQ RI FKORURSODVWV SOD\HG DQLPSRUWDQWUROHLQ KHOSLQJ WR HVWDEOLVK WKH ZLGHU WKHRU\ RI WKH UROH RI HQGRV\PELRVLV LQ WKHHYROXWLRQRI WKHHXNDU\RWLFFHOO &DUU ,QIRUPDWLRQ DERXW WKRVH PRGHUQ RUJDQLVPV VXFK DV Cyanophora paradoxa, ZKLFK KDYH LQWUDFHOOXODUF\DQHOOHVZLWK DVOLJKWUHVHPEODQFH WR IUHHOLYLQJF\DQREDFWHULDLV WKHUHIRUH RI SDUWLFXODU LQWHUHVW&KDSWHU $GDPV ,W LVXVXDOO\DVVXPHG WKDW F\DQHOOHVUHSUHVHQWDQHYROXWLRQDU\GHDGHQG DQG DUH QRW KDOIZD\ VWDJHV WR WKH GHYHORSPHQW RI D W\SLFDOFKORURSODVW 1HYHUWKHOHVVKDUGO\ DQ\WKLQJLV NQRZQ RI WKHHFRORJLFDOUHDVRQVIRUWKHVXFFHVV RI WKH RUJDQLVPV ZKLFK FRQWDLQF\DQHOOHV
,97D[RQRP\ ,W LV FOHDUO\ LPSRUWDQWWKDW HFRORJLFDO DFFRXQWV XVH QDPHVZKLFKFDQ EH LQWHUSUHWHGHDVLO\DQG UHOLDEO\ E\
&KDSWHU ,QWURGXFWLRQWRWKH&\DQREDFWHULD RWKHU UHVHDUFKHUV 8QIRUWXQDWHO\ WKLV LV RIWHQ IDU IURP WKH FDVH DQG WKHUH LV QR RQH ERRN RU HYHQ VHULHV RI ERRNV ZKLFK UHVROYHVDOOWKHSUREOHPV ,W LV HVVHQWLDO IRU DQ\RQHZDQWLQJWRLQWHUSUHWDQG ZULWH WKHLU RZQ DFFRXQWV RI F\DQREDFWHULDO HFRORJ\ WR NQRZ VRPHWKLQJDERXW WKH UHDVRQV IRU WKLV 0RUH GHWDLOHG EDFNJURXQG LV JLYHQ E\ &DVWHQKRO] DQG :DWHUEXU\ DQG :KLWWRQ 7KH WD[RQRPLF V\VWHP ZKLFK GHYHORSHGGXULQJWKH WK FHQWXU\ ZDV EDVHG DOPRVW HQWLUHO\ RQ PRUSKRORJ\DQGWKH,QWHUQDWLRQDO&RGHRI %RWDQLFDO 1RPHQFODWXUH WDNHVYDULRXVSDSHUVSXEOLVKHG GXULQJ WKHSHULRG DV WKHVWDUWLQJSRLQWIRUWKH YDOLG SXEOLFDWLRQ RI QDPHV RI ILODPHQWRXV IRUPV 5DWKHU VXUSUL]LQJO\ WKH VWDUWLQJ SRLQW IRU WKH PRUSKRORJLFDOO\ OHVVFRPSOH[IRUPVLV PXFK HDUOLHU HJ /LQQDHXV DOWKRXJKWKHUH ZDV RI FRXUVH OLWWOHXQGHUVWDQGLQJRI WKHLUGLYHUVLW\DWWKDWWLPH $ QXPEHU RI IORUDV VXPPDUL]LQJWKHNQRZQ VSHFLHV RI EOXHJUHHQ DOJDH WKH ERWDQLFDO QDPH LQ SDUWLFXODU UHJLRQVKDYHEHHQ SXEOLVKHGGXULQJWKHWKFHQWXU\ 6HYHUDO RI WKHVHSURYLGH D ORWRI LQIRUPDWLRQDERXW VSHFLHVRFFXUULQJHOVHZKHUHLQ WKH ZRUOG ,Q DQ\ FDVH LW LV FOHDU WKDW PDQ\ VSHFLHV KDYH D YHU\ ZLGH GLVWULEXWLRQ 7KHEHVWNQRZQ IORUDLVWKDWRI *HLWOHU ZKLFK IRFXVVHG RQ FHQWUDO(XURSHEXWLVVWLOO DQ HVVHQWLDO VRXUFH RI LQIRUPDWLRQIRUWKRVH ZDQWLQJ WR QDPH WKH RUJDQLVPV LQ DQ\ SDUW RI WKH ZRUOG )Up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
ZKDW KDG EHHQ GRQHVLQFH WKH V E\ WKHPDMRULW\RI PLFURELRORJLVWVGHDOLQJ ZLWK RWKHU JURXSV :KLWWRQ 'URXHWUHYHUWHG WR WKH XVH RI RQO\ D IHZ VLPSOH PRUSKRORJLFDO IHDWXUHVWKH PRUH WKDQ VSHFLHVRI EOXHJUHHQ DOJDH GHVFULEHG LQ WKH OLWHUDWXUH ZHUH HYHQWXDOO\ UHGXFHG WR VSHFLHV 7KH QRPHQFODWXUDO VLPSOLFLW\ RI WKHV\VWHPWHPSWHGPDQ\ HFRORJLFDOUHVHDUFKHUVLQ WKH 86$ GXULQJ WKH V DQG V WR XVH KLV QDPHV 1R H[WHQVLYHFULWLTXHRI WKHV\VWHPKDVHYHU EHHQ SXEOLVKHG EXW LW KDV TXLWH ULJKWO\ IDOOHQ LQWR
5 GLVXVH DSDUW IURP WKH IDFW WKDW WZR JHQHULF QDPHV Anacystis DQG Agmenellum, ZHUH XVHG IRU VWUDLQV ZKLFK KDYHVLQFHEHFRPHZLGHO\ XVHG LQUHVHDUFK $FFRXQWV RI QHZ VSHFLHV DQG DVHULHVRI UHYLHZV E\ - .RPiUHN ZLWK HLWKHU ( .DQQ RU . $QDJQRVWLGLV KDYH VRXJKWWREULQJWKHFODVVLFDOERWDQLFDOV\VWHPXS WR GDWH LQ PRUH FRQYHQWLRQDOZD\V 6HYHUDO RI WKH HDUOLHU UHYLHZV SURYLGH FRPSUHKHQVLYH DFFRXQWV RI LPSRUWDQW JHQHUD VXFK DV Chamaesiphon .DQQ DQG Phormidium .DQQ DQG .RPiUHN EXW ZHUH SXEOLVKHG LQ MRXUQDOV ZLWK RQO\ D OLPLWHG GLVWULEXWLRQ DQG ZULWWHQ LQ *HUPDQ DQG KHQFH KDYH RIWHQ EHHQ RYHUORRNHG 0RUHUHFHQWDFFRXQWV HJ .RPiUHN DQG $QDJQRVWLGLV KDYH EHHQ XVHG PRUH ZLGHO\ DOWKRXJK WKH\ LQWURGXFH PDMRU QRPHQFODWXUDOFKDQJHV IRU VRPH JHQHUD HVSHFLDOO\ IRUPV OLNH QRQKHWHURF\VWRXV ILODPHQWRXV Oscillatoria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
%$ :KLWWRQDQG03RWWV
Table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³IDOVH´ EUDQFKHVWULFKRPHVGRQRW IRUP KHWHURF\VWV $NLQHWHVDSSDUHQWO\ QRWUHFRUGHG 2UGHU1RVWRFDOHV %LQDU\ GLYLVLRQLQ RQHSODQHJLYLQJULVH WR VHQDWHWULFKRPHVWKRXJKVRPHWLPHVZLWK ³IDOVH´EUDQFKHVRQH RU PRUH FHOOV SHU WULFKRPHGLIIHUHQWLDWHLQWRDKHWHURF\VWDW OHDVW ZKHQ FRQFHQWUDWLRQRI FRPELQHG QLWURJHQ LV ORZ 6RPHDOVRSURGXFHDNLQHWHV 2UGHU6WLJRQHPDWDOHV %LQDU\ GLYLVLRQSHULRGLFDOO\RU FRPPRQO\ LQ PRUH WKDQRQHSODQH JLYLQJULVH WR PXOWLVHULDWHWULFKRPHVRU WULFKRPHV ZLWK WUXH EUDQFKHV RU ERWKDSSDUHQWO\DOZD\VSRVVHVVDELOLW\ WR IRUPKHWHURF\VWVVRPHDOVRIRUPDNLQHWHV
EHHQ XVHG WRHVWDEOLVKD VLPLODULW\PDWUL[IRU VWUDLQV RIAnabaena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iUHN DQG $QDJQRVWLGLV LQFOXGHV JHQHUD ZKLFK ZHUH ILUVW GHVFULEHG DFFRUGLQJ WRWKH,QWHUQDWLRQDO&RGHRI 1RPHQFODWXUH
RI %DFWHULD 7KH DFFRXQWV ODUJHO\ E\ 5: &DVWHQKRO] DQG -% :DWHUEXU\ LQ %HUJH\¶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
VLQJOHPRUSKRORJLFDO IRUPVXFKDV Trichodesmium &KDSWHU 3DHUO Aphunothece halophytica &KDSWHU 2UHQ DQG Nostoc commune &KDSWHU 3RWWV %HFDXVH VXFK IRUPV PD\ EH UHDGLO\ LGHQWLILHG EXW RFFXU LQ GLYHUVH ORFDOLWLHV WKH FRQFHSW RI D IRUP VSHFLHV LVKHOSIXOLQ WKHLUVWXG\ *DUFLD3LFKHO HW DO FRQFOXGHG WKDW Microcoleus chthonoplastes PD\ LQ IDFW EH FRQVLGHUHG D VLQJOH FRVPSROLWDQ VSHFLHV EXW WKH H[WHQW WR ZKLFK RWKHU VXFK VSHFLHV IRUPFORVHJHQHWLFJURXSLQJVUHPDLQVWREHHYDOXDWHG FULWLFDOO\ ,Q WKH FDVH RI KRW VSULQJ FRPPXQLWLHV UHFHQW VWXGLHV LQGLFDWH WKDW WKHLU GLYHUVLW\ LV PXFK PRUH FRPSOH[ WKDQ HDUOLHUWKRXJKW&KDSWHU :DUG DQG &DVWHQKRO]
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
,Q WKH .D]X]D '1$ 5HVHDUFK ,QVWLWXWH RI -DSDQ FRPPHQFHG VHTXHQFH DQDO\VLV RI WKH HQWLUH JHQRPH RI Synechocystis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rochlorococcus LQ WKH 86$ DQG Anabaena LQ -DSDQ 7KHVH PD\ EH FRPSDUHG$XJXVW ZLWK ZKDW KDV EHHQ GRQH IRU RWKHUEDFWHULD FRPSOHWH VHTXHQFHV SOXV LQ SURJUHVV 'HYHORSPHQWV RI LQFUHDVLQJO\ UHILQHG PHWKRGV WR VHTXHQFH JHQRPHV DQG DQDO\]H VHTXHQFH GDWD ZLOO SHUPLW WKH JHQHWLF DQDO\VLV DQG PDQLSXODWLRQ RI PRUSKRORJLFDOO\ PRUH FRPSOH[ F\DQREDFWHULD LQFOXGLQJ JOREDO ³IRUP´ VSHFLHV VXFK DV Trichodesmiurn, ZKLFK SOD\ VXFK SURPLQHQW UROHV LQ WKH HDUWK¶V HFRORJ\
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
%$ :KLWWRQ DQG03RWWV SK\FRHU\WKULQULFK Synechococcus FDQ EH H[WUHPHO\ DEXQGDQW LQ FRDVWDO PDULQH HQYLURQPHQWV ZKHUH WLWUHV PD\ H[FHHG P/ &KDSWHU 6XWWOH 7KH KLJK GHQVLW\ RI ERWKF\DQRSKDJH DQG Synechococcus KHUH UHVXOW LQ KLJK HQFRXQWHU IUHTXHQFLHV DQG VHOHFWLRQ IRU Synechococcus SRSXODWLRQV WKDW DUH ODUJHO\ UHVLVWDQW WR LQIHFWLRQ ZKHUHDV WKH ORZHU GHQVLW\ RI Synechococcus LQ RFHDQLF ZDWHUV DQG KHQFH PXFK ORZHUHQFRXQWHUIUHTXHQFLHVLVSUREDEO\ UHVSRQVLEOH IRU WKH ORZHU UHVLVWDQFH RI WKH Synechococcus WR LQIHFWLRQ
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
9,,, &\DQREDFWHULDDV+HDOWK )RRG 7KH F\DQREDFWHULXPRIWHQ NQRZQ DV 6SLUXOLQDEXW WD[RQRPLFDOO\ Arthrospira, LV ZLGHO\ PDUNHWHG DV D KHDOWK IRRG%HOD\HW DO 7KHQRPHQFODWXUDO SUREOHPUHVXOWV IURP*HLWOHU ZKR VXEPHUJHG ZKDW LV D YHU\ GLIIHUHQW RUJDQLVP ZLWKLQ WKH JHQXV Spirulina. 6SLUXOLQDLVJURZQ XQGHUDZLGHUDQJHRI V\VWHPV IURP FUXGH SRQGV WR ODUJHVFDOH RXWGRRU UDFHZD\V3ODWHDE DQG VRSKLVWLFDWHG ELRUHDFWRUV $V D UHVXOW D ORW LV NQRZQ DERXW WKH SK\VLRORJLFDO HFRORJ\ RI WKHRUJDQLVPLQ WKHODERUDWRU\DQG ODUJH VFDOHFRPPHUFLDOFXOWXUH&KDSWHU 9RQVKDN DQG 7RPDVHOOL WKRXJK OHVV DERXW KRZ WKHVH IHDWXUHV UHODWH WR JURZWK RI WKH RUJDQLVP LQ LWV QDWXUDO HQYLURQPHQW ,Q FRQWUDVW OLWWOH LV NQRZQ DERXW WKH SK\VLRORJLFDO HFRORJ\ RI Aphanizomenon, DQRWKHU F\DQREDFWHULXP PDUNHWHGRQDODUJHVFDOHDV D KHDOWK
&KDSWHU ,QWURGXFWLRQWRWKH&\DQREDFWHULD IRRG DQG ZKLFK LVDSSDUHQWO\DW SUHVHQW REWDLQHGIRU WKLV SXUSRVHVROHO\IURPQDWXUDO EORRPVLQ RQHODNH .ODPDWK /DNH LQ 2UHJRQ 86$ ,Q YLHZ RI LWV HFRQRPLF LPSRUWDQFH LW LV VXUSUL]LQJKRZ OLWWOHKDV EHHQ SXEOLVKHG DERXW WKLV WKRXJK 'UDSHDX DQG *XWHUPXWK KDYHSURYLGHGDVWDUW
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rthrospira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
GLUHFWHG WR DQVZHULQJ TXHVWLRQV DERXW WKH HQYLURQPHQW )XUWKHU WKH SUREOHPV RI LVRODWLQJ VWUDLQVKDYHVRPHWLPHVEHHQH[DJJHUDWHGDWOHDVWLQ WKH FDVH RI ILODPHQWRXVIRUPV ,WLVRI FRXUVHGLIILFXOW WR VLPXODWHKLJK89%UDGLDWLRQRUGLHOF\FOHVRI (K DQG 2 LQ WKHODERUDWRU\ 1HYHUWKHOHVVLQ PDQ\ FDVHV DOOWKDWLV QHHGHG WRLVRODWHDFXOWXUHRI DILODPHQWRXV VSHFLHV LV DQ XQGHUVWDQGLQJ RI WKH ELRORJ\ RI WKH RUJDQLVP VXFKDVWKHIDFWRUVOHDGLQJWRWKHIRUPDWLRQ RI KRUPRJRQLD RU DNLQHWHV 7KH JUHDW PDMRULW\ RI VWUDLQV ZLWKRXW JDVYDFXROHV VXUYLYH ZHOO LQ OLTXLG QLWURJHQ IRU SHULRGV XS WR DW OHDVW WZHQW\ \HDUV ZLWKRXW DQ\ VSHFLDO SUHFDXWLRQV RWKHU WKDQ WKH SUHVHQFH RI GLPHWK\OVXOSKR[LGH DXWKRUV XQSXEOLVKHG GDWD 2QO\WKRVHRUJDQLVPVOLNHPDQ\ VWUDLQV RI 6SLUXOLQD ZKLFK KDYH JDVYDFXROHV XQGHU DOO HQYLURQPHQWDO FRQGLWLRQV WHVWHG 0 0XKOLQJ SHUV FRPP SRVH D UHDO SUREOHP IRU ORQJWHUP PDLQWHQDQFH 6XEFXOWXUHV VKRXOG QRUPDOO\ EH PDGH IURP ROG EDWFK FXOWXUHV ZKHUH JURZWK RI WKH RUJDQLVP KDV EHHQ OLPLWHG E\ WKHVDPHIDFWRUWKDWXVXDOO\ OHDGVWR JURZWK OLPLWDWLRQ LQ WKH ILHOG %HFDXVH RI WKH ZLGHVSUHDG DELOLW\ RI VWUDLQV ZKLFK DUH OLNHO\ WR HQFRXQWHUQLWURJHQOLPLWDWLRQLQWKHILHOGWREHDEOHWR IL[QLWURJHQLW VHHPVSUREDEOH WKDW JURZWK RI PRUH VSHFLHV LQ QDWXUH ZLOO EH OLPLWHG E\ WKHDYDLODELOLW\RI SKRVSKDWH WKDQ E\ DQ\ RWKHU IDFWRU ,W LV WKHUHIRUH HVSHFLDOO\ LPSRUWDQW WR UHGXFH WKH SKRVSKDWH FRQFHQWUDWLRQ RI PDQ\ZLGHO\XVHGPHGLD
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
5HIHUHQFHV %HOD\ $ 2WD < 0L\DNDZD . DQG6KLPDPDWVX+ &XUUHQW NQRZOHGJH RQ SRWHQWLDO KHDOWK EHQHILWV RI Spirulina. - $SSO 3K\FRO %LQGHU %- DQG &KLVKROP6: 5HODWLRQVKLSEHWZHHQ'1$ F\FOH DQG JURZWK UDWH LQ Synechococcus VS VWUDLQ %DFWHULRO %LQLV]NLHZLF] ' &HVQDYLFLHQH ( DQG 6KXE '$ 6HOI VSOLFLQJ JURXS , LQWURQ LQ F\DQREDFWHULDOLQLWLDWRU PHWKLRQLQH W51$ HYLGHQFHIRUODWHUDOWUDQVIHURI LQWURQVLQEDFWHULD(0%2 - %U\DQW '$ HG 7KH0ROHFXODU%LRORJ\ RI &\DQRDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 'RUGUHFKW 7KH 1HWKHUODQGV &DPSEHOO(/ %UDKDPVKD % 0HHNV -& 0XWDWLRQ RI DQ DOWHUQDWLYH VLJPD IDFWRU LQ WKH F\DQREDFWHULXP 1RVWRF punctiforme UHVXOWV LQ LQFUHDVHGLQIHFWLRQRI LWVV\PELRWLFSODQW SDUWQHU$QWKRFHURVSXQFWDWXV - %DFWHULRO &DUU 1* )UHLEXUJ WR9LHQQD ORRNLQJ DW F\DQREDFWHULD RYHU WKHWZHQW\ILYH\HDUV ,Q 3HVFKHN *$ /|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scillatoria limnetica. %DFWHULRO 'HPRXOLQ 9 DQG -DQVVHQ 03 5HODWLRQVKLS EHWZHHQ GLDPHWHU RI WKHILODPHQWDQG FHOO VKDSH LQ EOXHJUHHQDOJDH %U 3K\FRO - 'HVLNDFKDU\ 79 &\DQRSK\WD ,QGLDQ &RXQFLO RI $JULFXOWXUDO 5HVHDUFK 1HZ 'HOKL 'UDSHDX&DQG*XWHUPXWK )% 7XUQLQJQXLVDQFHDOJDHLQWR KHDOWK IRRG WK ,QWHUQDWLRQDO &RQIHUHQFH RQ 7R[LF &\DQREDFWHULD6HSWHPEHU 2FWREHU%HDXIRUW1RUWK &DUROLQD &RPSLODWLRQ RI $EVWUDFWV S 'URXHW ) DQG 'DLO\ :$ 5HYLVLRQ RI WKH FRFFRLG 0\[RSK\FHDH %XWOHU 8QLY6WXG )RJJ*( *URZWKDQGKHWHURF\VWSURGXFWLRQ LQ Anabaena cylindrica /HPP 1HZ3K\WRORJLVW )UpP\ 3 &\DQRSK\FpHVGHV &{WHV G¶(XURSH 0WP 6RF 1DWQ 6FL 1DW 0DWK&KHUERXUJ )UWP\ 3 /HV 0\[RSK\FpHV GH O $IULTXH (TXDWRULDOH )UDQoDLVH $UFKLYHV GH %RWDQLTXH ,,, 0WPRLUH 1R SS UHSULQWHGSULYDWHO\ &DHQ *DUFLD3LFKHO ) 3UXIHUW%HERXW DQG0X\]HU* 3KHQRW\SLF DQG SK\ORJHQHWLFDQDO\VHVVKRZ Microcoleus chthonoplastes WR EH D FRVPRSROLWDQF\DQREDFWHULXP $SSO (QYLURQ 0LFURELRO *HLWOHU/ &\DQRSK\FHDH ,Q 5DEHQKRUVW V .U\SWRJDPHQ )ORUD9ROXPH $NDGHPLVFKH 9HUODJVJHVHOOVFKDIW/HLS]LJ +HUGPDQ0-DQYLHU 0 5LSSND 5DQG6WDQLHU 5< *HQRPH VL]HRIF\DQREDFWHULD - *HQ0LFURELRO +HUGPDQ0 DQG 5LSSND 5 +RUPRJRQLD DQG EDHRF\WHV 0HWKRGV LQ(Q]\PRORJ\ +LURNL 0 6KLPL]X $ /L 5 :DWDQDEH 0 DQG :DWDQDEH 00 3K\FRORJLFDO5HVHDUFK6XSSO
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
&KDSWHU ,QWURGXFWLRQWRWKH&\DQREDFWHULD RI V\PELRWLF F\DQREDFWHULD LQ VRPH OLFKHQV $SSO (QYLURQ 0LFURELRO 3RVW$) 7KHSURFKORURSK\WHV DQDOJDOHQLJPD %LRORJ\ RIFKORURSK\OODEFRQWDLQLQJSKRWRV\QWKHWLFSURNDU\RWHV ,Q 6HFNEDFK-HG (QLJPDWLF0LFURRUJDQLVPVDQG/LIHLQ([WUHPH (QYLURQPHQWV SS .OXZHU $FDGHPLF 3XEOLVKHUV 'RUGUHFKW7KH1HWKHUODQGV 5DVPXVVHQ 8 DQG 6YHQQLQJ 09 )LQJHUSULQWLQJ RI F\DQREDFWHULDEDVHGRQ3&5ZLWKSULPHUVGHULYHGIURPVKRUW DQGORQJWDQGHPO\UHSHDWHGUHSHWLWLYHVHTXHQFHV$SSO(QYLURQ 0LFURELRO 5LSSND5'HUXHOOHV-%:DWHUEXU\-%+HUGPDQ0DQG6WDQLHU 5< *HQHULFDVVLJQPHQWVVWUDLQKLVWRULHVDQGSURSHUWLHV RISXUHFXOWXUHVRIF\DQREDFWHULD -*HQ0LFURELRO 5XGL . 6NXOEHUJ 20 DQG-DNREVHQ .6 (YROXWLRQ RI F\DQREDFWHULD E\ H[FKDQJH RI JHQHWLF PDWHULDO DPRQJ SK\OHWLFDOO\UHODWHGVWUDLQV-%DFWHULRO 5XGL.6NXOEHUJ20/DUVHQ)DQG-DNREVHQ.6 6WUDLQ FKDUDFWHUL]DWLRQDQGFODVVLILFDWLRQRIR[\SKRWREDFWHULDLQFORQH FXOWXUHVRQWKHEDVLVRI6U51$VHTXHQFHVIURPWKHYDULDEOH UHJLRQV99DQG9$SSO(QYLURQ0LFURELRO 6FKHOGHPDQ3%DXUDLQ'%RXK\56FRWW00XKOLQJ0:KLWWRQ %$%HOD\$DQG:LOPRWWH$ Arthrospira µ6SLUXOLQD¶ VWUDLQVIURPIRXUFRQWLQHQWVDUHUHVROYHGLQWRRQO\WZRFOXVWHUV EDVHGRQDPSOLILHGULERVRPDO'1$UHVWULFWLRQDQDO\VLV RIWKH LQWHUQDOO\WUDQVFULEHGVSDFH )(060LFURELRO/HWW 6FKRSI JW DQG:DOWHU05 2ULJLQDQGHDUO\HYROXWLRQRI F\DQREDFWHULD 7KH JHRORJLFDOHYLGHQFH ,Q &DUU 1* DQG :KLWWRQ%$HGV 7KH%LRORJ\RI&\DQREDFWHULDSS %ODFNZHOO2[IRUGDQG8QLYHUVLW\RI&DOLIRUQLD3UHVV%HUNHOH\ 6LPRQ5'(1980) '1$FRQWHQWRIKHWHURF\VWVDQGVSRUHVRIWKH ILODPHQWRXV F\DQREDFWHULXP $QDEDHQD variabilis. )(06 0LFURELRO/HWW 6LYRQHQ.DQG-RQHV* &\DQREDFWHULDOWR[LQV ,Q&KRUXV ,DQG%DUWUDP-HGV 7R[LF&\DQREDFWHULDLQ:DWHU $*XLGH WR WKHLU 3XEOLF +HDOWK &RQVHTXHQFHV 0RQLWRULQJ DQG 0DQDJHPHQW ( )16SRQ/RQGRQ 1HZ
:DWHUEXU\ -% :DWVRQ 6: *XLOODUG 55/ DQG%UDQG/( :LGHVSUHDG RFFXUUHQFH RI D XQLFHOOXODU PDULQH SODQNWRQLF F\DQREDFWHULXP1DWXUH :KLWWRQ %$ 'LYHUVLW\ HFRORJ\ DQG WD[RQRP\ RI WKH F\DQREDFWHULD ,Q 0DQQ1+DQG &DUU1*HGV 3KRWRV\QWKHWLF 3URNDU\RWHV SS 3OHQXP 1HZ
&KDSWHU 7KH )RVVLO 5HFRUG 7UDFLQJ WKH 5RRWV RI WKH &\DQREDFWHULDO /LQHDJH - :LOOLDP 6FKRSI ,*33 &HQWHU IRU WKH 6WXG\ RI (YROXWLRQ DQG WKH 2ULJLQ RI /LIH 'HSDUWPHQWRI(DUWKDQG6SDFH6FLHQFHV DQG0ROHFXODU%LRORJ\,QVWLWXWH 8QLYHUVLW\RI&DOLIRUQLD/RV$QJHOHV &DOLIRUQLD 86$
6XPPDU\ , 7UDFLQJWKH5RRWVRI &\DQREDFWHULD 3URJUHVV DQG3UREOHPV $ 'DUZLQ V'LOHPPD7KH 0LVVLQJ 3UHFDPEULDQ )RVVLO 5HFRUG % 7KH8QLYHUVDO7UHH RI /LIH & /LPLWDWLRQV RI WKH (DUO\ )RVVLO5HFRUG ' 7LPLQJ(YROXWLRQE\WKH$PLQR$FLG &ORFN ,, $QFLHQW &\DQREDFWHULD $ )RVVLODQG0RGHUQ/RRNDOLNHV % $UH )RVVLO &\DQREDFWHULD &\DQREDFWHULD" ,OO +RZ2OG LV WKH &\DQREDFWHULDO /LQHDJH" $ (YLGHQFH )URPWKHa *DROG $SH[&KHUW ,9 3DOHRELRORJ\ )RVVLOV *HRORJ\ DQG *HRFKHPLVWU\ $ *HRORJLF(YLGHQFHRI &\DQREDFWHULDO3KRWRV\QWKHVLV % *HRFKHPLFDO (YLGHQFHRI (DUO\HYROYLQJ /LQHDJHV &DUERQ,VRWRSLF(YLGHQFH RI 3URFDU\RWLF 3KRWRDXWRWURSKV 6XOIXU,VRWRSLF(YLGHQFH RI 6XOIDWHUHGXFLQJ%DFWHULD &DUERQ,VRWRSLF(YLGHQFH RI 0HWKDQRJHQLF$UFKDHDQV 9 3DOHRELRORJ\ )LQDO $UELWHU RI &RPSHWLQJ 7KHRULHV
6XPPDU\ 6LQFH WKH PLGV IROORZLQJ D FHQWXU\ RI XQUHZDUGHG VHDUFK LPSUHVVLYHSURJUHVV KDV EHHQ PDGH WRZDUG GHFLSKHULQJ WKH 3UHFDPEULDQIRVVLOUHFRUGHYLGHQFHRIOLIHH[WDQWGXULQJWKHHDUOLHVWVHYHQHLJKWKV RI JHRORJLF WLPH +XQGUHGV RI IRVVLOLIHURXV XQLWV KDYH EHHQ GLVFRYHUHG FRQWDLQLQJ WKRXVDQGV RI PLFURELDO IRVVLOV GRPLQDQWO\EXWQRW H[FOXVLYHO\F\DQREDFWHULDO²DQGWKHGRFXPHQWHG DQWLTXLW\ RI OLIHKDVEHHQH[WHQGHGWRDQ DJH URXJKO\ WKUHHTXDUWHUV WKDW RI WKH(DUWK 0XWXDOO\UHLQIRUFLQJOLQHV RI HYLGHQFHSDOHRQWRORJLFDOJHRORJLFDO DQG LVRWRSLF JHRFKHPLFDOLQGLFDWHWKDW VWURPDWROLWLFPLFURELDO HFRV\VWHPVHYLGHQWO\LQFOXGLQJF\DQREDFWHULDDQG RWKHU PHPEHUV RI WKHEDFWHULDOGRPDLQZHUH H[WDQW a 0D DJR PHWKDQRJHQLF DUFKDHDQV E\ a 0DDJR DQG *UDPQHJDWLYH VXOIDWHUHGXFLQJEDFWHULDDWOHDVWDVHDUO\ DV a 0D DJR 7KH GLVFUHSDQF\ EHWZHHQWKHVH GDWHV DQG WKRVH VXJJHVWHGIRUHPHUJHQFH RI WKHVH JURXSV E\ D UHFHQWO\ SURSRVHG DPLQRDFLGEDVHG PROHFXODU FORFN LV WRR JUHDW DQG WRR FRQVLVWHQWWR EH LJQRUHG 7KH FKDOOHQJH LV WR XQLI\ WKH PROHFXODUGDWD ZLWK WKH LQFUHDVLQJO\ZHOOHVWDEOLVKHG SDOHRELRORJLFUHFRUG
B. A. Whitton and M. Potts (eds), The Ecology of Cyanobacteria, © Kluwer Academic Publishers. Printed in the Netherlands.
-:6FKRSI
, 7UDFLQJ WKH 5RRWV RI &\DQREDFWHULD 3URJUHVV DQG3UREOHPV $ 'DUZLQ V'LOHPPD7KH0LVVLQJ 3UHFDPEULDQ )RVVLO 5HFRUG ,Q ZKHQ &KDUOHV 'DUZLQ XQYHLOHG KLV PRQXPHQWDO PRQRJUDSK The Origin of Species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terra incognita. %XW'DUZLQGLG NQRZ WKDW LI DQVZHUVZHUHQRWIRUWKFRPLQJKLVWKHRU\ ZDVLQMHRSDUG\ "If the theory [of evolution] be true it is indisputable that before the lowest Cambrian stratum was deposited ... the world swarmed with living creatures. [Yet] to the question why we do notfind rich fossiliferous deposits belonging to these earliest periods ... I can give no satisfactory answer. 7KH FDVHDWSUHVHQWPXVWUHPDLQLQH[SOLFDEOHDQGPD\EH WUXO\ XUJHGDV D YDOLG DUJXPHQW DJDLQVW WKH YLHZV KHUHHQWHUWDLQHG 'DUZLQ &KDSWHU; 7KRXJK'DUZLQSRVHGWKHSUREOHPLWZDVQRWXQWLO D FHQWXU\ ODWHU ² LQWKHPLGV ZLWK WKH ELUWK RI D QHZ ILHOG RI VFLHQFH 3UHFDPEULDQ SDOHRELRORJ\ ² WKDW WKLV HDUOLHVW PLVVLQJ IRVVLO UHFRUG EHJDQ WR EH XQFRYHUHG &ORXG 6FKRSI D 3URJUHVV VLQFH WKH V KDV EHHQ LPSUHVVLYH 7KH GRFXPHQWHG UHFRUG RI OLIH KDV EHHQ H[WHQGHGVWHDGLO\ DQGQRZVWUHWFKHVWRQHDUO\ ELOOLRQ\HDUV*D DJR DGDWHLQ WKHJHRORJLFSDVWDSSURDFKLQJWKHDJH *D RI WKH (DUWKLWVHOI
Abbreviations: BIF - Banded Iron Formation; Ga - billion (109) years; Ma- million (106) years
% 7KH 8QLYHUVDO7UHHRI /LIH $V WKH HDUO\ IRVVLO UHFRUG FDPH LQFUHDVLQJO\ LQWR IRFXV GXULQJ WKH V DQG V EHQFKPDUN DGYDQFHV ZHUHXQGHUZD\LQ VWXGLHVRI WKH PROHFXODU ELRORJ\ RI OLYLQJV\VWHPV %\ WKH ODWH VWKHVH KDG FRDOHVFHGLQWRSRZHUIXOPHDQVWR GHWHUPLQH WKH HYROXWLRQDU\ SK\ORJHQHWLF UHODWLRQV DPRQJ all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² WKH Eucarya RUJDQLVPV KDYLQJFHOOVLQ ZKLFK FKURPRVRPHV DUH HQFDSVXODWHG LQ D GLVWLQFW PHPEUDQHERXQGHG QXFOHXV WKH Archaea QRQ QXFOHDWHG PLFURRUJDQLVPV LQFOXGLQJ PHWKDQRJHQLF SURFDU\RWHV DQG PDQ\ H[WUHPRSKLOHV PLFUREHV WKDW
&KDSWHU 7KH )RVVLO 5HFRUG&\DQREDFWHULD, /LQHDJH WKULYH LQ H[FHSWLRQDOO\ DFLGLF KLJK WHPSHUDWXUH VHWWLQJV DQG WKH Bacteria, WKH GRPDLQWKDWLQFOXGHV WKH F\DQREDFWHULD DQG EDFWHULD RI WUDGLWLRQDO FODVVLILFDWLRQV )RXUWK WKH(XFDU\DLV PRUH FORVHO\ UHODWHG WR WKH $UFKDHD WKDQ WKH%DFWHULDDQG WKH ODVW FRPPRQ DQFHVWRU RI DOO RUJDQLVPVOLYLQJWRGD\ ² WKH URRW VWRFN RI WKH 8QLYHUVDO 7UHH RI /LIH ²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
& /LPLWDWLRQVRIWKH(DUO\)RVVLO5HFRUG $FFXUDWHGDWLQJRIWKHWLPHV RIRULJLQ RIWKH PDMRU ELRORJLF OLQHDJHV KDV EHHQ DORQJVWDQGLQJ JRDO LQ 3UHFDPEULDQSDOHRELRORJ\6FKRSI 6FKRSIHW DO 6FKRSI E 7KLV LV D \RXQJ ILHOG KRZHYHU DQG WKH HDUO\ IRVVLO UHFRUG LV WRR LQFRPSOHWHO\ NQRZQ WR SURYLGH SUHFLVH DQVZHUV 0RUHRYHUHYHQXQGHUWKHEHVWRIFLUFXPVWDQFHVIRVVLO HYLGHQFHFDQUHFRUGRQO\WKHILUVWdetected RFFXUUHQFH RIDELRORJLFOLQHDJHQRWLWVILUVWactual RFFXUUHQFH 7RJHWKHU IRVVLOV DQG DVVRFLDWHG JHRORJLFDO DQG JHRFKHPLFDOLQGLFDWRUVRIELRORJLFDFWLYLW\WKDWLV SDOHRELRORJLF HYLGHQFH FDQHVWDEOLVKDPLQLPXP DJHIRUDOLQHDJHEXWWKH\FDQQRWUHYHDOKRZPXFK HDUOLHUWKHOLQHDJHDFWXDOO\H[LVWHG
' 7LPLQJ(YROXWLRQE\WKH$PLQR$FLG &ORFN $QRWKHU ZD\ WR VROYH WKLV SUREOHP EDVHG RQ WKH DPLQRDFLGFRPSRVLWLRQ RI SURWHLQV KDVUHFHQWO\EHHQ SURSRVHG E\ 'RROLWWOH HW DO 3URWHLQVHYROYH
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older DJHV WKDQ WKRVH GRFXPHQWHG E\ IRVVLOV
-:6FKRSI
&KDSWHU7KH)RVVLO5HFRUG&\DQREDFWHULDO/LQHDJH PLVLQWHUSUHWHG DQG DUH XQUHODWHG WR SUHVHQWGD\ OLQHDJHV 3HUKDSV OLIH RULJLQDWHG EHFDPH H[WLQFW WKHQ RULJLQDWHG D VHFRQG WLPH DERXW 0D DJR 2U SHUKDSV OLNHWKHU51$WUHHVWKHDPLQRDFLGGDWD \LHOG UHOLDEOHHYLGHQFHRQO\ RI WKHEUDQFKLQJRUGHU RI HYROXWLRQ QRW WKH WLPLQJ RI WKDWEUDQFKLQJ +RZ ILUPO\ HVWDEOLVKHG is WKH HDUO\IRVVLOHYLGHQFH" :KDW DPRXQW RI GDWD ZRXOG KDYH WR EHRYHUWXUQHGIRU WKH DPLQRDFLGFORFN WR SURYHFRUUHFW"
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a 0DROG %LWWHU 6SULQJV )RUPDWLRQ RI FHQWUDO $XVWUDOLD )LJ -6 6FKRSI6FKRSIDQG %ODFLF 6LQFH WKDW WLPH VXFK VLPLODULWLHV KDYH EHHQ HQFRXQWHUHG VR IUHTXHQWO\ WKDW LW LV QRZ FRPPRQ SUDFWLFH DPRQJ 3UHFDPEULDQ SDOHRELRORJLVWVWRQDPHIRVVLOWD[DDIWHU WKHLU PRGHUQ PRUSKRORJLFDOFRXQWHUSDUWVE\ DGGLQJ DSSURSULDWH SUHIL[HV SDODHR HR RU VXIIL[HV RSVLV LWHV WR WKH QDPHV RI SUHVHQWGD\ F\DQREDFWHULDO JHQHUD 0HQGHOVRQ DQG 6FKRSI 6FKRSI D 0RUHWKDQ VXFK QDPHVDNHV IRUH[DPSOH Palaeolyngbya, Eomicrocoleus, Aphanocapsaopsis, DQG Oscillatorites) DUH LQ XVDJH ZRUOGZLGH 7KH VWULNLQJ VLPLODULW\ RI 3UHFDPEULDQPLFURIRVVLOV WR PRGHUQ F\DQREDFWHULDLV FKDUDFWHULVWLFRI D ZLGH UDQJH RI PRUSKRORJLF W\SHV DPRQJ ZKLFK DUHIRVVLOV WKDW DUH KXQGUHGV RI PLOOLRQV RI \HDUV ROGHU WKDQ WKH GDWH RI a 0DVXJJHVWHGE\ WKHDPLQRDFLGFORFN IRU WKH RULJLQDWLRQ RI WKH JURXS )RU H[DPSOH Eoentophysalis belcherensis 3ODWHE ² FRPSDUHG ZLWK VSHFLHV RI PRGHUQ Entophysalsis 3ODWHD DQG RQ WKLV EDVLV RI UHIHUUHG WR WKH OLYLQJF\DQREDFWHULDO IDPLO\ (QWRSK\VDOLGDFHDH *ROXELF DQG +RIPDQQ ² LV SUHVHUYHG LQ PLFURELDOO\ ODPLQDWHG VWURPDWROLWHV RI 1RUWKZHVW 7HUULWRULHV &DQDGD WKDW DUH a 0D ROG +RIPDQQ 7KRXJK WKH
IRVVLO WD[RQ LV GHFLGHGO\ WRR ROG WR EH F\DQREDFWHULDODFFRUGLQJWRWKHDPLQRDFLG FORFN WKH IRVVLO DQG PRGHUQ VSHFLHV DUH PRUSKRORJLFDOO\ LQGLVWLQJXLVKDEOH LQ FHOO VKDSH DQG LQ IRUP DQG DUUDQJHPHQW RI RULJLQDOO\ PXFLODJLQRXV FHOO HQFRPSDVVLQJ HQYHORSHV H[KLELW VLPLODU IUHTXHQF\ GLVWULEXWLRQV RI GLYLGLQJFHOOVDQG HVVHQWLDOO\ LGHQWLFDO SDWWHUQV RI FHOOXODUGHYHORSPHQWUHVXOWLQJIURP FHOO GLYLVLRQ LQ WKUHH SHUSHQGLFXODU SODQHV IRUP PLFURWH[WXUDOO\ VLPLODU VWURPDWROLWLF VWUXFWXUHV LQ FRPSDUDEOH LQWHUWLGDO WR VKDOORZ PDULQH HQYLURQPHQWDO VHWWLQJV XQGHUJR HVVHQWLDOO\LGHQWLFDO SRVWPRUWHP GHJUDGDWLRQ VHTXHQFHV DQG RFFXU LQ PLFURELDO FRPPXQLWLHV WKDW DUH FRPSDUDEOH LQ ERWK VSHFLHVFRPSRVLWLRQDQG ELRORJLFDOGLYHUVLW\*ROXELF DQG +RIPDQQ 0DQ\ RWKHUF\DQREDFWHULDOORRNDOLNHV YLUWXDOO\ DOO IURP LQWHUWLGDO WR VKDOORZ PDULQH VWURPDWROLWLF VHWWLQJV OLNH WKRVH LQKDELWHG E\ WKHLU OLYLQJ PRUSKRORJLFDO FRXQWHUSDUWV DUH WRR ROG E\ WKH DPLQR DFLG GDWD $PRQJ WKHVH LV Oscillatoriopsis majuscula .QROO HW DO D a 0DROG ILODPHQWRXVWD[RQPRUHWKDQ PP LQ GLDPHWHU DQG WKXV YHU\ PXFK EURDGHU WKDQ DOPRVW DOO QRQF\DQREDFWHULDO SURFDU\RWLF ILODPHQWV RI ZKLFK DUH PP LQ GLDPHWHU 6FKRSI D 1XPHURXV RWKHUVXFKH[DPSOHVFDQEHFLWHG ,QGHHG DV VXPPDUL]HG LQ )LJ IRVVLOV LQWHUSUHWHG DV F\DQREDFWHULD DQG UHIHUUHG WR WKH &KURRFRFFDFHDH (QWRSK\VDOLGDFHDH DQG 2VFLOODWRULDFHDH RFFXU LQ PDQ\ JHRORJLF XQLWV ROGHU WKDQ SHUPLWWHG E\ WKH DPLQR DFLG FORFN ² LI any RI WKHVH WKRXVDQGV RI IRVVLOV DQG WKHPRUHWKDQ F\DQREDFWHULDOVSHFLHV WR ZKLFK WKH\KDYHEHHQUHIHUUHGKDVEHHQ LGHQWLILHG FRUUHFWO\WKH SURWHLQEDVHG DPLQR DFLG FORFN LV LQ HUURU
% $UH )RVVLO &\DQREDFWHULD &\DQREDFWHULD" :KDW DERXW WKH LGHD WKDW OLIH RULJLQDWHG EHFDPH H[WLQFW WKHQ RULJLQDWHGDVHFRQGWLPHOHDGLQJWR WKH DGYHQW RI F\DQREDFWHULD a 0D DJR" 3HUKDSV 3UHFDPEULDQ SDOHRELRORJLVWV KDYH VR DFFHSWHG WKH SUHVXPHG F\DQREDFWHULDO DIILQLW\ RI WKH IRVVLOV IRXQG WKH\ KDYH QHJOHFWHG WR FRQVLGHU RWKHU SRVVLELOLWLHV HVSHFLDOO\IRUYHU\ DQFLHQWIRUPV ,VWKHUHDEUHDNLQ WKHIRVVLOUHFRUGRI F\DQREDFWHULD a 0DDJR" 2QH ZD\ WR DGGUHVVWKLVSUREOHPREMHFWLYHO\ IUHH RI WD[RQRPLF SUHGLOHFWLRQV LV WKURXJK GHWDLOHG PRUSKRPHWULF DQDO\VHV RI WKH IRVVLO UHFRUG 6XFK VWXGLHVKDYHEHHQ FDUULHG RXW RQ 55 FRPPXQLWLHV RI
-: 6FKRSI
Fig. 2. (QVKHDWKHG FRORQLHV RI FRFFRLG IRVVLO F\DQREDFWHULD IURP VWURPDWROLWLF FKHUW RI WKH 0DROG &KLFKNDQ )RUPDWLRQ RI WKH =KDQDWDV UHJLRQ VRXWKHUQ .D]DNKVWDQ (A) Terraphycus bistratosus, &KURRFRFFDFHDH 2JXUWVRYD DQG 6HUJHHY (B) Palaeopleurocapsa fusiforma, 3OHXURFDSVDFHDH 2JXUWVRYD DQG 6HUJHHY (C) Palaeopleurocapsa reniforma, 3OHXURFDSVDFHDH 2JXUWVRYDDQG6HUJHHY
&KDSWHU 7KH )RVVLO 5HFRUG&\DQREDFWHULDO /LQHDJH
Fig. 3. &RORQLDO (A, B, E and F) DQG XQLFHOOXODU (D) IRVVLO F\DQREDFWHULD DQG D FRORQLDO IRVVLO EDFWHULXP (C), IURP VLOWVRQH (D) DQG VWURPDWROLWLF FKHUWV (A-C, E DQG F) RI WKH 0DROG 3DUDGLVH &UHHN )RUPDWLRQ RI HDVWFHQWUDO 4XHHQVODQG $XVWUDOLD (A-C) DQG WKH 0DROG 6DWND )RUPDWLRQ RI VRXWKHUQ %DVKNLULD 5XVVLD (D-F). (A DQG B) &XERLGDO EucapsisOLNH FRORQ\ &KURRFRFFDFHDH /LFDUL HW DO (C) Metallogenium VS Eoastrion VS D P\FRSODVPDOLNHLURQ DQG PDQJDQHVHR[LGL]LQJ EDFWHULXP &ORXG (D) Eomarginata striata, Chroococcaceae (Jankauskas 1980). (E) Palaeopleurocapsa kelleri, Pleurocapsaceae (Krylov and Sergeev, 1987). (F)Gloeodiniopsisuralicus,&KURRFRFFDFHDH.U\ORYDQG6HUJHHY
-:6FKRSI
Fig.4. )RVVLO F\DQREDFWHULDO FRORQLHV A-D DQG J) XQLFHOOV E-G DQG I DQG HQVKHDWKHG SDLUHG XQLFHOOV + K and L) IURP VWURPDWROLWLF FKHUW RI WKH a 0DROG 6NLOORJDOHH 'RORPLWH RI FHQWUDO 6RXWK $XVWUDOLD $XVWUDOLD (A) 7DEXODU 0HULVPRSHGLDOLNH FRORQ\ &KURRFRFFDFHDH6FKRSIDQG )DLUFKLOG (B, D DQG J) 5RVHWWHOLNHFRORQLHV VLPLODUWRMyxosarcina, 3OHXURFDSVDFHDH)DLUFKLOG (C, E, F-I, K DQG L) &KURRFRFFDFHDQ PLFURIRVVLOV 6FKRSIDQG )DLUFKLOG )DLUFKLOG 6FKRSI
&KDSWHU 7KH )RVVLO 5HFRUG&\DQREDFWHULDO /LQHDJH
Fig. 5. )RVVLO F\DQREDFWHULDO WXEXODU VKHDWK (A) DQG FHOOXODU WULFKRPHV (B-F) IURP VLOWVWRQHV RI WKH 0DROG 6KWDQGLQ )RUPDWLRQ RI VRXWKHUQ%DVKNLULD 5XVVLD (A) Siphonophycuscostatus, 2VFLOODWRULDFHDH-DQNDXVNDV ( B andD) Tortunema VS 2VFLOODWRULDFHDH -DQNDXVNDV (C) Calyptothrixgeminata, 2VFLOODWRULDFHDH -DQNDXVDNDV (EandF) Calyptothrixalternata, 2VFLOODWRULDFHDH -DQNDXVNDV
-: 6FKRSI
Fig. 6. )LODPHQWRXV (A, B, H-S) DQG FRFFRLGDO (C-G) F\DQREDFWHULDO IRVVLOV IURP VWURPDWROLWLF FKHUWV RI WKH 0DROG .ZDJXQW )RUPDWLRQ RIQRWKHUQ $UL]RQD 86$(A-I) DQG WKH 0DROG %LWWHU6SULQJV )RUPDWLRQ RIVRXWKHUQ 1RUWKHUQ 7HUULWRU\ $XVWUDOLD (J-S). (A, B, H, ZDQG S) Eomycetopsis VSS WXEXODU VKHDWKV RIRVFLOODWRULDFHDQ ILODPHQWV 6FKRSI 6FKRSIHW DO . (C-G)&KURRFRFFDFHDQ XQLFHOOV 6FKRSI HW DO (J) Filiconstrictosus diminutus, 2VFLOODWRULDFHDH 6FKRSI DQG %ODFLF . (K) Palaeolyngbya minor 6FKRSI DQG %ODFLF (L) Veteronostocale amoenum 6FKRSI DQG %ODFLF (M DQG N) Caudiculophycus rivularioides, "5LYXODULDFHDH 6FKRSI (O) Caudiculophycus acuminatus, "5LYXODULDFHDH 6FKRSI DQG %ODFLF ( P ) Parritiofilum gongyloides, 2VFLOODWRULDFHDH 6FKRSI DQG %ODFLF (Q) Palaeolyngbya barghoomiana, 2VFLOODWRULDFHDH 6FKRSI (R) Filiconstrictosus majusculus, 2VFLOODWRULDFHDH 6FKRSIDQG%ODFLF
&KDSWHU 7KH )RVVLO 5HFRUFG&\DQREDFWHULDO /LQHDJH
Fig. 7. )RVVLO FHOOXODU F\DQREDFWHULDO WULFKRPHV (A) DQG RULJLQDOO\ WXEXODU ILODPHQWRXV VKHDWKV (B, C) IURP VLOWVWRQHV RI WKH 0DROG /DNKDQGD )RUPDWLRQ RI HDVWHUQ 6LEHULD 5XVVLD (A) Pulaeolyngbya helva, 2VFLOODWRULDFHDH +HUPDQQ ( B ) Leiothrichoides tenuitunicatus, 2VFLOODWRULDFHDH+HUPDQQ (C)Palaeovaucheriaclavata,"2VFLOODWRULDFHDH+HUPDQQ
FRFFRLGDORUILODPHQWRXVPLFURIRVVLOVa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a *DORQJSHULRGDQDO\]HG $Q\ PDUNHG ELRWLF GLVFRQWLQXLW\RFFXUULQJGXULQJ WKLV SHULRG ² OLNH WKDW SUHGLFWHG E\ WKH DPLQR DFLG GDWD ²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a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living genera RI FKURRFRFFDFHDQV RU RVFLOODWRULDFHDQV 0RUHRYHU RI FRFFRLGDO WD[D RI ROGHUWKDQ 0DDQG RI RI WKRVH\RXQJHUDUHLQGLVWLQJXLVKDEOHLQ PRUSKRORJ\ IURP living species RI FKURRFRFFDFHDQV RI ILODPHQWRXVWD[D RI ROGHUWKDQ 0D DQG
-:6FKRSI RI RI WKRVH \RXQJHU KDYH living species-level PRUSKRORJLFDO FRXQWHUSDUWV DPRQJ WKH 2VFLOODWRULDFHDH DQG YLUWXDOO\ DOO FRORQ\ IRUPV H[KLELWHG E\ OLYLQJ FKURRFRFFDFHDQV RFFXU DOVR DPRQJ FKURRFRFFDFHDQOLNH IRVVLOV ERWK ROGHU DQG \RXQJHU WKDQ 0D )LJV 6FKRSI G HI
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
$ (YLGHQFH)URPWKH*DROG$SH[ &KHUW 7KRXJK WKHUH LV UHDVRQ WR TXHVWLRQ VRPHUHSRUWV RI H[FHHGLQJO\ DQFLHQW PLFURIRVVLOV 6FKRSI DQG :DOWHU VXFKXQFHUWDLQW\ GRHV QRWDSSO\WR WKH PLFURELDO DVVHPEODJH UHFHQWO\ GHVFULEHG IURP WKH a 0DROG FRQJORPHUDWLF $SH[ FKHUW 6FKRSI F 7KH JHRJUDSKLF DQG VWUDWLJUDSKLFVRXUFHRI WKH IRVVLOLIHURXV KRUL]RQ LV NQRZQ ZLWK FHUWDLQW\ IRVVLO EHDULQJVDPSOHVKDYLQJEHHQ FROOHFWHGIURPRXWFURS
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scillatoriopsis VSS DQG PRGHUQ (Oscillatoria VSS 7KH SUHVHQFH RI RVFLOODWRULDFHDQF\DQREDFWHULDLQ WKH $SH[ DVVHPEODJH LV FRQVLVWHQW DOVR ZLWK IRXU RWKHU OLQHV RI HYLGHQFH )LUVW PRGHOV RI WKH HDUO\ JOREDO HFRV\VWHP DQG WUDFH HOHPHQW FHULXP DQG HXURSLXP FRQFHQWUDWLRQV LQ $SH[DJH VHGLPHQWV VKRZ WKDW DHURELF UHVSLUDWLRQ DQG 2SURGXFLQJ
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
,9 3DOHRELRORJ\ )RVVLOV *HRORJ\ and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¶V DWPRVSKHUHIRUH[DPSOH DUHWRRORQJWHUP WR IDOO ZLWKLQ WKH VFRSH RI 3KDQHUR]RLF VWXGLHV 6LPLODUO\ EHFDXVH YLUWXDOO\ DOO 3KDQHUR]RLF IRVVLOV DUH UHDGLO\ UHODWDEOH WR OLYLQJ ELRORJLF JURXSV TXHVWLRQV UHJDUGLQJ WKHLU EDVLF PHWDEROLF FKDUDFWHULVWLFV GR QRW DULVH 8QFHUWDLQW\ PD\ H[LVW ZKHWKHU RQHRUDQRWKHUJURXSRI GLQRVDXUV ZDV PRUH RUOHVVZDUPEORRGHGEXWLWLVVDIHWRDVVXPHDOOZHUH R[\JHQUHTXLULQJ REOLJDWHDHUREHVOLNHWKHLUUHSWLOLDQ
-:6FKRSI
Primaevifilum amoenum Fig. 8. &HOOXODU ILODPHQWRXV F\DQREDFWHULXPOLNH IRVVLOV Primaevifilum amoenum "2VFLOODWRULDFHDH IURP WKH a 0DROG $SH[ FKHUW RI QRUWKZHVWHUQ :HVWHUQ $XVWUDOLD $XVWUDOLD 6FKRSI F
DQG DYLDQ GHVFHQGDQWV DQG HDUO\HYROYLQJODQG SODQWV DVVXUHGO\ ZHUH R[\JHQSURGXFLQJ SKRWRV\QWKHVL]HUV HYHQ WKRXJK WKLV OLQHDJHOLNHWKH GLQRVDXUVLV ORQJ H[WLQFW 7KH 3UHFDPEULDQ (RQ KRZHYHU LV QHDUO\ DQ RUGHU RI PDJQLWXGH ORQJHU WKDQ WKH3KDQHUR]RLFDQG HYHQWV WKDW DUH PLQRU RU HYHQ LPSHUFHSWLEOH RYHU 3KDQHUR]RLF WLPHFDQKDYHFXPXODWLYHLPSDFWLI WKH\ VSDQ DSSUHFLDEOH SRUWLRQV RI HDUOLHU (DUWK KLVWRU\ )RU H[DPSOH WKH GUDPDWLF FKDQJH IURP DQ HDUO\ DQR[LF JOREDOHQYLURQPHQWWRWKHFXUUHQWR[\JHQULFK a DWPRVSKHUH LI VSDQQLQJ WKH WLPH IURP WKH $SH[ IRVVLOV WR WKH SUHVHQW ZRXOG KDYH UHTXLUHG
VXVWDLQHG DGGLWLRQ RI RQO\ 2 SHU PLOOLRQ \HDUV
&KDSWHU 7KH )RVVLO 5HFRUG&\DQREDFWHULDO /LQHDJH
&21,&$/(1'&(//6Primaevifilumconicoterminatum Fig. 9 &HOOXODU ILODPHQWRXV F\DQREDFWHULXPOLNH IRVVLOV (Archaeocillatoriopsis disciformis, DERYH Primaevifilum conicoterminatum, EHORZ "2VFLOODWRULDFHDH IURP WKH a 0DROG $SH[ FKHUW RI QRUWKZHVWHUQ :HVWHUQ $XVWUDOLD $XVWUDOLD6FKRSI F
SDOHRQWRORJ\ 'DWD IURP JHRORJ\ DQG JHRFKHPLVWU\ DUHDOVRFUXFLDO WRXQUDYHOLQJWKHHDUO\KLVWRU\ RI OLIH
$ *HRORJLF (YLGHQFH RI &\DQREDFWHULDO 3KRWRV\QWKHVLV 7KH FDSDELOLW\ WR FDUU\ RXW R[\JHQSURGXFLQJ SKRWRV\QWKHVLV LV D XQLYHUVDO FKDUDFWHULVWLF RI F\DQREDFWHULDWKDW GLVWLQJXLVKHVWKHP IURP DOO RWKHU SURFDU\RWHV ,I WKH LQIHUHQFH LV FRUUHFW WKDW F\DQREDFWHULD DUH UHSUHVHQWHG DPRQJ WKH $SH[ IRVVLOV WKH$SH[VHTXHQFH DQG FRHYDOJHRORJLFXQLWV VKRXOGFRQWDLQHYLGHQFH RI WKH UHDFWDQWV UHTXLUHG IRU
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
-: 6FKRSI
Primaevifilum laticellulosum Fig. 10. &HOOXODU ILODPHQWRXV F\DQREDFWHULXPOLNH IRVVLOV Primaevifilum laticellulosum "2VFLOODWRULDFHDH IURP WKH a 0DROG $SH[ FKHUWRIQRUWKZHVWHUQ:HVWHUQ$XVWUDOLD$XVWUDOLD6FKRSI
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a WR a E\ ZHLJKW 6WUDXVV DQG 0RRUH
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
&KDSWHU 7KH )RVVLO 5HFRUG&\DQREDFWHULDO /LQHDJH IHUUXJLQRXV EHGV DUH FRPSRVHG RI ILQH UXVWOLNH SDUWLFOHV RI KHPDWLWH )H2 DQG LQ VRPH GHSRVLWV PDJQHWLWH )H2 %,)VKDYHDFKDUDFWHULVWLFGXOOWR EULJKW UHG FRORU 7KH LURQHYLGHQWO\RZHV LWVRULJLQWR WKH K\GURWKHUPDO FLUFXODWLRQ RI VHDZDWHU WKURXJK RFHDQLF FUXVW SULPDULO\ DW GHHS VXEPDULQH ULGJH V\VWHPV ,Q D GLVVROYHG IHUURXV VWDWH LW WKHQ FLUFXODWHG XSZDUG LQWRVKDOORZHUUHDFKHVRI WKH ZDWHU FROXPQ ² RIWHQ VHDVRQDOO\ JLYLQJ ULVH WR WKH GLVWLQFWLYH PLOOLPHWULF EDQGLQJ ² ZKHUH LW ZDV R[LGL]HG WR IHUULF LURQ FKLHIO\ E\ UHDFWLRQ ZLWK GLVVROYHG PROHFXODU R[\JHQ DQG SUHFLSLWDWHG DV D ILQH UXVW\ UDLQ RI PLQXWH LQVROXEOH LURQ R[LGH SDUWLFOHV 7KH (DUWK¶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a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
EHFDXVH WKH PLQHUDOV DUH HDVLO\ GHVWUR\HG GXULQJ ZHDWKHULQJ LQ WKH SUHVHQW R[\JHQULFK DWPRVSKHUH 1HYHUWKHOHVV ODUJH VHGLPHQWDU\ RUH GHSRVLWV WKDW FRQWDLQGHWULWDOSHEEOHVRI XUDQLQLWH DQG S\ULWH RFFXU LQ JHRORJLF XQLWV ROGHU EXW QRW \RXQJHU WKDQ a 0D +ROODQG 7KH SHUVLVWHQFH RI SHEEOHV RI XUDQLQLWH DQG S\ULWH GXULQJ ZHDWKHULQJ IOXYLDO WUDQVSRUW DQG GHSRVLWLRQ LQ FRQJORPHUDWLF GHSRVLWV PRUHWKDQ 0D DJRLV FRQVLVWHQWZLWK D ORZ2 DWPRVSKHUH XS WR WKLVWLPHDVDUHRWKHUJHRORJLFGDWD PRVWQRWDEO\WKH PLQHUDORJ\ RI SDOHRVROVDQFLHQWVRLO KRUL]RQV 7DNHQWRJHWKHUWKHVHDQG UHODWHG OLQHV RI HYLGHQFHVXSSRUWWKHFRQFOXVLRQWKDW WKH 2 FRQWHQW RI WKHDWPRVSKHUHLQFUHDVHGGUDPDWLFDOO\EHWZHHQ DQG 0D DJR HYLGHQWO\IURP WR RI WKHSUHVHQWDWPRVSKHULFOHYHO+ROODQG *HRORJLF DQG SDOHRQWRORJLF HYLGHQFH WKXV DJUHH ZLWK WKH LQIHUUHG H[LVWHQFH RI R[\JHQSURGXFLQJ F\DQREDFWHULD DV HDUO\ DV a0DDJR DERXWWZR ELOOLRQ\HDUVHDUOLHUWKDQ VXJJHVWHG E\ WKH DPLQR DFLG FORFN 'DWD IURP LVRWRSLF JHRFKHPLVWU\ DUH DOVR FRQVLVWHQW ZLWK WKLVLQWHUSUHWDWLRQDQGOLNHWKHUHFRUG RI FHOOXODUO\SUHVHUYHGQRQF\DQREDFWHULDOSURFDU\RWHV )LJ HYLGHQFHWKHSUHVHQFHRI RWKHUHDUO\HYROYHG PLFURELDOOLQHDJHVDV ZHOO
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¶/HDU\ +HQFH UHODWLYHWRWKHFDUERQ LVRWRSH 3'% VWDQGDUG cyanobacteria, R[\JHQLF
SKRWRV\QWKHVL]HUV WHQG WR EH &HQULFKHG E\ DQR[\JHQLF photosynthetic bacteria, E\ methanogenic DQG FKHPRDXWRWURSKLF )LJ ,VRWRSLF archaeans, E\ VLJQDWXUHV RI VXFK DXWRWURSKVDUH SUHVHUYDEOHLQ WKH JHRORJLF UHFRUG ZKHUH LQUHODWLYHO\ \RXQJ VHGLPHQWV WKH\ FDQ EH OLQNHGWR VSHFLILFPHWDEROLF VRXUFHV $ SULPH H[DPSOH LV WKH (RFHQH 0DROG 0HVVHO 6KDOH RI *HUPDQ\ LQ ZKLFK ELRPDUNHU LQGLFDWRUV RI R[\JHQLF SKRWRV\QWKHVLV FKORURSK\OO GHULYDWLYHV DQR[\JHQLF EDFWHULDO SKRWRDXWRWURSK\ GHJUDGDWLRQ SURGXFWV RIEDFWHULRFKORURSK\OOd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individual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
&DUERQ,VRWRSLF(YLGHQFHRI3URNDU\RWLF 3KRWRDXWRWURSKV 7KH FDUERQ LVRWRSLF FRPSRVLWLRQV RI R[\JHQLF F\DQREDFWHULDO DQG DQR[\JHQLF SKRWRV\QWKHWLF EDFWHULDO SURFDU\RWLFSKRWRDXWRWURSKVDUHUDWKHUZHOO FKDUDFWHUL]HG 'DWD DUHDYDLODEOHIRUD ODUJH QXPEHU RI PRGHUQ F\DQREDFWHULXPGRPLQDWHG VWURPDWROLWLF FRPPXQLWHV 'HV0DUDLVHW DO DV ZHOO DVIRU LQGLYLGXDO VSHFLHV RI FKURRFRFFDFHDQV Agmenellum, Anacystis, Coccochloris, Synechococcus DQG Microcoleus, Oscillatoria, RVFLOODWRULDFHDQV Schizothrix FXOWXUHG XQGHU GLYHUVHFRQGLWLRQV )LJ
-:6FKRSI :LWK UHJDUG WR SKRWRV\QWKHWLFEDFWHULDFDUERQ LVRWRSLF IUDFWLRQDWLRQ RFFXUULQJ GXULQJ SKRWRDXWRWURSK\ KDV EHHQ FKDUDFWHUL]HG LQ FXOWXUHV IRU UHSUHVHQWDWLYHV RI WKH &KORURELDFHDH Chlorobium &KORURIOH[DFHDH Chloroflexus 5KRGRVSLULOODFHDH Rhodospirillum DQG &KURPDWLDFHDH Chromatiurn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² PLQLPXP IUDFWLRQDWLRQ RFFXULQJ DW ORZ &2 OHYHOV DVVRFLDWHG FRPPRQO\ ZLWKKLJKSRSXODWLRQGHQVLWLHV DQG PD[LPXP IUDFWLRQDWLRQ DW KLJK &2 FRQFHQWUDWLRQV WR WR WLPHV WKH SUHVHQW DWPRVSKHULFOHYHO OLNHWKRVH SRVWXODWHG IRU WKHHDUO\DQG PLG3UHFDPEULDQ HQYLURQPHQW.DVWLQJ ² DQDO\VHV RI NHURJHQRXV RUJDQLF PDWWHU LQ PRUH WKDQ D WKRXVDQG UHODWLYHO\ XQPHWDPRUSKRVHG 3UHFDPEULDQ VDPSOHV HVWDEOLVK WKDW WKH FDUERQ LVRWRSLF VLJQDWXUH RI SKRWRV\QWKHVLV H[WHQGV WR DW OHDVWa0DDJR 6FKLGORZVNL HW DO6WUDXVV HW DO +D\HV 6FKRSI E 7KHVH GDWD WRJHWKHU ZLWK WKH JHRORJLF DQG SDOHRQWRORJLF HYLGHQFHRXWOLQHGDERYHVXSSRUWWKHFRQFOXVLRQWKDW WKH F\DQREDFWHULDO SKRWRV\QWKHWLF EDFWHULDO DQG *UDPSRVLWLYH EDFWHULDO OLQHDJHV KDYH EHHQ H[WDQW VLQFH DW OHDVW a 0DDJRDERXWWZRELOOLRQ\HDUV HDUOLHU WKDQ VXJJHVWHGE\ WKH DPLQR DFLG FORFN)LJ
6XOIXU,VRWRSLF(YLGHQFH RI6XOIDWH UHGXFLQJ %DFWHULD *HRFKHPLFDO HYLGHQFH DOVR UHFRUGV WKH SUHVHQFH RI *UDPQHJDWLYHEDFWHULDDW D WLPH PXFK WRRROG E\ WKH DPLQR DFLG FORFN 7KLV OLQHDJH LQFOXGHV
&KDSWHU 7KH)RVVLO5HFRUG&\DQREDFWHULDO/LQHDJH
Fig. 11. &DUERQ LVRWRSLFFRPSRVLWLRQVRILQRUJDQLFFDUERQUHVHUYRLUVSURFDU\RWLFDXWRWURSKV DQG3UHFDPEULDQNHURJHQV )UDFWLRQDWLRQ 'DWDIURP$EHOVRQDQG+RHULQJ UDQJHVIRUFXOWXUHVDQGDQDHURELF PLFURELRWDV DUH UHFDOFXODWHG IRU D &2 VRXUFH ZLWK &DOGHU DQG 3DUNHU :RQJ HW DO *DPHV DQG +D\HV 3DUGXHHWDO )XH[ %DUJKRRPHWDO 6LUHYnJHWDO )XFKV HW DO 0L]XWDQLDQG:DGD %HO\DHY HW DO +D\HV +D\HV HW DO 6FKLGORZVNL HW DO +ROR DQG 6LUHYDJ 5XEH\ HW DO 'HV 0DUDLV HW DO 6WUDXVV DQG 0RRUH DQG 6WUDXVVHWDO
GLVVLPLODWRU\ VXOIDWHUHGXFLQJ EDFWHULD VXFK DV Desulfovibrio, Desulfotomaculum, DQG Desulfomonas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a UDQJH RI DQG DUH WKXV GLVWLQJXLVKDEOHIURPWKH VXOILGH PLQHUDOV LQ LJQHRXV URFNV ZKLFK IDOO JHQHUDOO\ZLWKLQD QDUURZ UDQJH RI DERXW 7KH LVRWRSLF VLJQDWXUH RI PLFURELDO VXOIDWH UHGXFWLRQ DQG WKXV RI *UDP QHJDWLYH EDFWHULDFDQEHWUDFHGEDFNWR DWOHDVW 0D DJR 6FKLGORZVNL HW DO DQG WKH OLQHDJH PD\ ZHOO KDYHEHHQH[WDQWDV HDUO\DV 0D DJR 2KPRWR HW DO .DNHJDZD HW DO QHDUO\ WZR ELOOLRQ \HDUV WRRHDUO\ E\ WKHDPLQRDFLGFORFN )LJ
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
-:6FKRSI &2 +2 &2 + +&2+ &+2+
&+&2+
&+ &2 NFDO
&+ +2 NFDO &+ &2 +2 &+ &2 +2
&+&2
7KH PHWKDQHSURGXFHGKDV WKHODUJHVWYDULDWLRQ RI WKH && UDWLR RIDQ\NQRZQ ELRJHQLF PDWHULDO ZLWK YDOXHV UDQJLQJ IURP WR )XH[ 0D[LPXP LVRWRSLF IUDFWLRQDWLRQ DWWDLQDEOH GXULQJ SKRWRV\QWKHVLVERWK REVHUYHG :RQJ HW DO 3DUGXHHW DO 6LUHYnJHW DO DQG FDOFXODWHG 6XPPRQV DQG +D\HV+D\HV LV 7KXV KLJKO\ &HQULFKHG ELRJHQLF PHWKDQH FDQ EH GLVWLQJXLVKHGUHDGLO\ IURPSURGXFWV RI SKRWRV\QWKHVLV DQG LWV LQFRUSRUDWLRQ LQWR VHGLPHQWHG RUJDQLF PDWWHU OHDYHV DQ XQPLVWDNDEOH VLJQDWXUH LQ WKH SUHVHUYHG NHURJHQ .DSODQ DQG 1LVVHQEDXP 7KH SUHVHQFH RI NHURJHQV ZLWK YDOXHV UDQJLQJ IURP WR LQ DW OHDVW JHRORJLFXQLWV a WR a 0D LQDJH6WUDXVV DQG 0RRUH KDV WKHUHIRUHEHHQ LQWHUSUHWHG DV VWURQJHYLGHQFHIRUWKHSUHVHQFHRI ELRJHQLF PHWKDQH DQG WKXV RI PHWKDQRJHQLF DUFKDHDQVDWOHDVW 0D DJR+D\HV
9 3DOHRELRORJ\ )LQDO$UELWHURI &RPSHWLQJ 7KHRULHV 7DNHQ WRJHWKHU SDOHRQWRORJ\ JHRORJ\ DQG JHRFKHPLVWU\ SURYLGH FRQYLQFLQJ HYLGHQFH RI WKH DQWLTXLW\ DQG HYROXWLRQDU\FRQWLQXLW\RI WKH EDFWHULDO DQG DUFKDHDQ GRPDLQV 7KH OLQHV RI SDOHRELRORJLF HYLGHQFH DUH GLVWLQFW \HW PXWXDOO\ UHLQIRUFLQJ WKH GDWD YROXPLQRXV 6WURPDWROLWLFPLFURELDOHFRV\VWHPV HYLGHQWO\ LQFOXGLQJ F\DQREDFWHULD SKRWRV\QWKHWLF EDFWHULD DQG RWKHU PHPEHUV RI WKHEDFWHULDOGRPDLQ ZHUHH[WDQWa0DDJRPHWKDQRJHQLF DUFKDHDQV E\ a 0D DJR DQG *UDPQHJDWLYH VXOIDWH UHGXFLQJEDFWHULDDWOHDVWDV HDUO\DVa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² WKH VFLHQFH RI SDOHRELRORJ\ ² LV WKH FRXUW RI ODVWUHVRUW WKH ILQDO DUELWHU RI FRPSHWLQJWKHRULHVDERXWWKHKLVWRU\RI OLIH
$FNQRZOHGJPHQWV 7KLV SDSHU LV D UHYLVHG DQG H[SDQGHG YHUVLRQ RI D FRQWULEXWLRQSUHSDUHGHDUOLHUIRUDGLIIHUHQWDXGLHQFH 6FKRSI ,Q 3UHVV ,Q SUHSDULQJ LW , KDYH KDG WKH EHQHILW RI WKRXJKWIXO FRUUHVSRQGHQFH IURP 5XVVHOO 'RROLWWOH ZKLFK ,JUHDWO\DSSUHFLDWH 3UHSDUDWLRQRI WKLV PDQXVFULSWKDV EHHQ VXSSRUWHGE\ 1$6$ *UDQW 1$*:
5HIHUHQFHV $EHOVRQ3+DQG+RHULQJ7 &DUERQLVRWRSHIUDFWLRQDWLRQLQ IRUPDWLRQRIDPLQRDFLGVE\SKRWRV\QWKHWLFRUJDQLVPV 3URF1DW $FDG 6FL 86$ $OWHUPDQQ : DQG 6FKRSI -: 0LFURIRVVLOV IURP WKH 1HRDUFKHDQ&DPSEHOO *URXS *ULTXDODQG:HVW 6HTXHQFH RIWKH 7UDQVYDDO 6XSHUJURXS DQG WKHLU SDOHRQYLURQPHQWDO DQG HYROXWLRQDU\ LPSOLFDWLRQV 3UHFDPEULDQ 5HV %DUJKRRP (6 .QROO $+ 'HPEULFNL + DQG 0HLQVFKHLQ :* 9DULDWLRQ LQ VWDEOH FDUERQ LVRWRSHV LQ RUJDQLF PDWWHU IURP WKH *XQIOLQW ,URQ )RUPDWLRQ *HRFKLP &RVPRFKLP $FWD ²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² &DPEULGJH8QLYHUVLW\3UHVV1HZ
+D\HV -0 7DNLJLNX 5 2FDPSR 5 &DOORW +- DQG $OEUHFKW 3 ,VRWRSLF FRPSRVLWLRQV DQG SUREDEOH RULJLQV RI RUJDQLF PROHFXOHVLQWKH(RFHQH0HVVHO6KDOH1DWXUH² +HUPDQQ 71 1LWFKDW\H PLNURRUJDQL]P\ ODNKDQGLQVNRM VYLW\UHNL0DL >)LODPHQWRXVPLFURRUJDQLVPVIURPWKH/DNKDQGD )RUPDWLRQ RQ WKH 0D\D 5LYHU@ 3DOHRQWRO =KXU ² LQ5XVVLDQ +RIPDQQ +- 3UHFDPEULDQ PLFURIORUD %HOFKHU ,VODQGV &DQDGD 6LJQLILFDQFH DQG V\VWHPDWLFV - 3DOHRQWRO ² +RIPDQQ +- DQG 6FKRSI -: (DUO\ 3URWHUR]RLF PLFURIRVVLOV ,Q 6FKRSI-: HG (DUWK V (DUOLHVW %LRVSKHUH ,WV 2ULJLQDQG(YROXWLRQSS²3ULQFHWRQ8QLYHUVLW\3UHVV 3ULQFHWRQ1- +ROODQG +' (DUO\ 3URWHUR]RLF DWPRVSKHULF FKDQJH ,Q %HQJVWVRQ 6 HG (DUO\/LIHRQ(DUWKSS ² &ROXPELD 8QLYHUVLW\3UHVV1HZ1HZDOJDHIURPWKHXSSHU5LSKHDQ RI WKH VRXWKHUQ 8UDOV DQG DGMDFHQW DUHDV@ 3DOHRQWRO =KXU ²LQ5XVVLDQ .DNHJDZD7.DZDL+DQG2KPRWR+ %LRORJLFDODFWLYLWLHV DQG K\GURWKHUPDO DFWLYLW\ UHFRUGHG LQ WKH a 0RXQW 0F5DH 6KDOH+DPHUVOH\'LVWULFW:HVWHUQ$XVWUDOLD ,, 6XOIXULVRWRSLF FRPSRVLWLRQRIS\ULWH 5HVRXUFH*HRO ² .DSODQ ,5 DQG 1LVVHQEDXP $ $QRPDORXV FDUERQ LVRWRSH UDWLRVLQQRQYRODWLOHRUJDQLFPDWHULDO6FLHQFH ² .DVWLQJ -) 3URWHUR]RLF FOLPDWHV 7KH HIIHFW RI FKDQJLQJ DWPRVSKHULF FDUERQ GLR[LGH FRQFHQWUDWLRQV ,Q 6FKRSI-: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\SS ²&DPEULGJH8QLYHUVLW\3UHVV1HZ5LSKHDQ PLFURIRVVLOV RI WKH VRXWKHUQ8UDOV QHDUWKHWRZQRI.XVD@ 3DOHRQWRO=KXU ²LQ5XVVLDQ /LFDUL *5 &ORXG 3( -U DQG 6PLWK :' $ QHZ FKURRFRFFDFHDQ DOJD IURP WKH 3URWHUR]RLF RI4XHHQVODQG 3URF 1DW$FDG6FL86$² 0HQGHOVRQ &9 DQG 6FKRSI -: 3URWHUR]RLF DQG VHOHFWHG (DUO\ &DPEULDQPLFURIRVVLOV DQG PLFURIRVVLOOLNH REMHFWV ,Q 6FKRSI -: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS ² &DPEULGJH 8QLYHUVLW\ 3UHVV1HZ0LFURELRWD RI WKH &KLFKNDQ )RUPDWLRQ /LWWOH
-: 6FKRSI .DUDWDX 5DQJH 6RXWK .D]DNKVWDQ @ 3DOHRQWRO =KXU ²LQ5XVVLDQ 2KPRWR+.DNHJDZD7DQG/RZH'5 ELOOLRQ\HDUROG ELRJHQLF S\ULWHV IURP %DUEHUWRQ 6RXWK $IULFD 6XOIXU LVRWRSH HYLGHQFH1DWXUH 2¶/HDU\ 0+ &DUERQ LVRWRSLF IUDFWLRQDWLRQ LQ SODQWV 3K\WRFKHP ² 2OVHQ *- DQG :RHVH &5 5LERVRPDO 51$ $ NH\ WR SK\ORJHQ\ )$6(% - 3DUGXH -: 6FDODQ 56 9DQ %DDOHQ & DQG 3DUNHU 3/ 0D[LPXP FDUERQ LVRWRSH IUDFWLRQDWLRQ LQ SKRWRV\QWKHVLV E\ EOXHJUHHQ DOJDH DQG D JUHHQ DOJD *HRFKLP &RVPRFKLP $FWD ² 5XEH\(* -DQQDVFK+:DQG'HXVHU:* )UDFWLRQDWLRQ RI VWDEOH FDUERQ LVRWRSHV GXULQJ FKHPRDXWRWURSKLF JURZWK RI VXOIXUR[LGLQJEDFWHULD $SS(QY0LFURELRO ² 6FKLGORZVNL 0 +D\HV -0 DQG .DSODQ ,5 ,VRWRSLF LQIHUHQFHV RI DQFLHQW ELRFKHPLVWULHV &DUERQ VXOIXU K\GURJHQ DQGQLWURJHQ ,Q 6FKRSI-: HG (DUWK V (DUOLHVW%LRVSKHUH ,WV 2ULJLQDQG(YROXWLRQSS ²3ULQFHWRQ8QLYHUVLW\3UHVV 3ULQFHWRQ1- 6FKRSI -: 0LFURIORUD RI WKH %LWWHU 6SULQJV )RUPDWLRQ /DWH3UHFDPEULDQ FHQWUDO$XVWUDOLD-3DOHRQWRO² 6FKRSI-: 3UHFDPEULDQ PLFURRUJDQLVPV DQG HYROXWLRQDU\ HYHQWVSULRUWRWKHRULJLQRIYDVFXODUSODQWV %LRO5HY&DPEULGJH 3KLO6RF² 6FKRSI -: %LRVWUDWLJUDSKLF XVHIXOQHVV RI VWURPDWROLWLF 3UHFDPEULDQ PLFURELRWDV $ SUHOLPLQDU\ DQDO\VLV 3UHFDPEULDQ 5HV² 6FKRSI -: D +LVWRULFDO GHYHORSPHQW RI 3URWHUR]RLF PLFURSDOHRQWRORJ\ ,Q 6FKRSI -: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS ² &DPEULGJH8QLYHUVLW\3UHVV1HZ
&KDSWHU 7KH )RVVLO5HFRUG&\DQREDFWHULDO/LQHDJH 6FKRSI -: D &\DQREDFWHULD 3LRQHHUV RI WKH HDUO\ (DUWK 1RYD+HGZLJLD ² 6FKRSI -: E 0HWDEROLF PHPRULHV RI (DUWK V HDUOLHVW ELRVSKHUH ,Q 0DUVKDOO &5 DQG 6FKRSI-:HGV (YROXWLRQDQG WKH 0ROHFXODU 5HYROXWLRQ SS ² -RQHV %DUWOHWW %RVWRQ0$ 6FKRSI-:,Q3UHVV 7UDFLQJWKHURRWVRIWKHXQLYHUVDOWUHHRIOLIH ,Q%UDFN$HG 7KH0ROHFXODU2ULJLQVRI/LIH$VVHPEOLQJWKH 3LHFHV&DPEULGJH8QLYHUVLW\3UHVV1HZ
:RHVH &5 .DQGOHU2 DQG :KHOOLV 0/ 7RZDUGV DQDWXUDO V\VWHP RI RUJDQLVPV 3URSRVDO IRU WKH GRPDLQV $UFKDHD %DFWHULDDQG(XFDU\D3URF1DW$FDG6FL86$² :ROIH56 0LFURELDOIRUPDWLRQRIPHWKDQH $GY 0LFURELDO 3K\VLRO ² :RQJ : 6DFNHWW :0 DQG %HQHGLFW &5% ,VRWRSH IUDFWLRQDWLRQ LQ SKRWRV\QWKHWLF EDFWHULD GXULQJ FDUERQ GLR[LGH DVVLPLODWLRQ 3ODQW3K\VLRO²
&KDSWHU &\DQREDFWHULDLQ*HRWKHUPDO+DELWDWV 'DYLG0:DUG 5LFKDUG:&DVWHQKRO] 'HSDUWPHQW RI 0LFURELRORJ\ 0RQWDQD 6WDWH 8QLYHUVLW\ %R]HPDQ 07 86$ 2 'HSDUWPHQW RI %LRORJ\ 8QLYHUVLW\ RI 2UHJRQ (XJHQH 25 86$
1
6XPPDU\ , ,QWURGXFWLRQ ,, 'LVWULEXWLRQ RI 7KHUPRSKLOLF &\DQREDFWHULD%DVHGRQ 0RUSKRORJ\DQG(QULFKPHQW &XOWXUH $*HRJUDSKLF'LVWULEXWLRQ %'LVWULEXWLRQ 'HWHUPLQHGE\&KHPLVWU\ & :HOO6WXGLHG 0DW 6\VWHPV +XQWHU¶V+RW6SULQJ 2UHJRQ 2FWRSXV 6SULQJ
6XPPDU\ 5HFHQW DGYDQFHVLQ PROHFXODU ELRORJ\ HQDEOHG D PXFK PRUHGHWDLOHG YLHZ RI F\DQREDFWHULD WKDW LQKDELW ZHOO VWXGLHG KRW VSULQJKDELWDWV :KDW VHHPHG RQ WKH EDVLV RI PLFURVFRS\ DQG FXOWXUH PHWKRGV WR EH DUHODWLYHO\ VLPSOHVWRU\EHFDPH RQH ZKLFK LV PRUHFRPSOH[\HWPRUHRUGHUHG )RUH[DPSOH 6 U51$VWXGLHVRI RQH ZHOOVWXGLHG
B A Whitton and M Potts (eds). The Ecology of Cyanobacteria. 2000 Kluwer Academic Publishers Printed in the Netherlands
'0:DUGDQG5:&DVWHQKRO]
LQ SDUWIURPDGDSWLYHUDGLDWLRQRI VSHFLDOL]HGHFRW\SHV 7KHVHUHVXOWVFRUUHVSRQGZLWK SUHYLRXVGHVFULSWLRQVRI Synechococcus WHPSHUDWXUH³VWUDLQV´FXOWLYDWHGIURPDZHOOVWXGLHG2UHJRQKRWVSULQJPDWUDLVLQJLQWHUHVWLQJ TXHVWLRQV ZKLFKPD\EH DQVZHUHGWKURXJKPROHFXODUDQDO\VLV )RUH[DPSOHDUH WKHVDPHF\DQREDFWHULD IRXQG LQ JHRJUDSKLFDOO\ LVRODWHG VSULQJV RU GRHV OLPLWHG GLVSHUVDO DOVR FDXVH GLYHUJHQFH RI F\DQREDFWHULD ZLWK VXEVHTXHQW DGDSWLYH UDGLDWLRQV LQ LQGHSHQGHQW OLQHDJHV " $GYDQFHV LQ PLFURHOHFWURGH WHFKQRORJ\ SURYLGHG GHWDLOHGYLHZVRI WKHYHUWLFDOGLVWULEXWLRQDQGG\QDPLFVRI OLJKW R[\JHQDQGVXOILGH ZLWKLQ WKHSKRWLF]RQHLQ VXFKPDWVZKLFKPD\ UDQJHIURPaPPWR !FP LQWKLFNQHVV 7KHVH PHWKRGV LQFUHDVHGRXUNQRZOHGJH RI GLHO FKHPLFDO FKDQJHV ZLWKLQ WKH PDWV LQFOXGLQJ WKH EDVLV IRU G\QDPLFPLJUDWLRQV RI PRWLOH F\DQREDFWHULD 0LFURHOHFWURGHVWXGLHVRI RQH
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
,, 'LVWULEXWLRQ RIWKHUPRSKLOLF F\DQREDFWHULDEDVHGRQ PRUSKRORJ\DQG HQULFKPHQW FXOWXUH 2EYLRXV GLIIHUHQFHV LQ F\DQREDFWHULDO SRSXODWLRQV EHWZHHQ VSULQJV FDQ EH HDVLO\ GLVFHUQHG E\ VLPSOH PDFUR RU PLFURVFRSLF H[DPLQDWLRQ EDVHG RQ GLVWLQFWLYHPRUSKRORJLFDO )LJ RU SK\VLRORJLFDO 7DEOH FKDUDFWHULVWLFV 'LIIHUHQFHV REVHUYHG Abbreviations: F'1$ FRPSOHPHQWDU\ GHR[\ULERQXFOHLF DFLG '**( GHQDWXULQJ JUDGLHQW JHO HOHFWURSKRUHVLV +3/& KLJK SHUIRUPDQFHOLTXLGFKURPDWRJUDSK\ +7)KLJKWHPSHUDWXUHIRUP 3&5SRO\PHUDVHFKDLQUHDFWLRQ 895XOWUDYLROHWUDGLDWLRQ
DPRQJ VSULQJV D IHZ PHWHUV RUNLORPHWHUVIURPHDFK RWKHU DUH OLNHO\ GXH WR GLIIHUHQFHV LQ FKHPLFDO FRPSRVLWLRQ WHPSHUDWXUH RU H[SRVXUH WR VRODU LUUDGLDQFH 7KRVHGLIIHUHQFHVREVHUYHGDPRQJ ZLGHO\ VHSDUDWHG VSULQJV PD\ EH WKH FRQVHTXHQFH RI JHRJUDSKLFLVRODWLRQDQG OLPLWDWLRQV RI GLVVHPLQDWLRQ
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ynechococcus )LJ O D DQG E DUHFOHDUO\ UHVWULFWHG LQ JHRJUDSKLFDO GLVWULEXWLRQ $OO IRUPV RI WKHUPRSKLOLF Synechococcus DUH DEVHQW IURP ,FHODQGLF KRW VSULQJV DOWKRXJK QXPHURXV VSULQJV H[LVW WKDW DSSHDU FKHPLFDOO\ VXLWDEOH 0RUSKRORJLFDOO\ GLIIHUHQW Synechococcus H[LVW LQ 1HZ =HDODQG DQG (XURSHDQ VSULQJVEXW RQO\ IRUPV ZKLFK JURZXS WRWHPSHUDWXUHOLPLWVRI DQG & UHVSHFWLYHO\&DVWHQKRO] 7DEOH )RUPV RI Synechococcus WKDW JURZ LQ QDWXUH XS WR & LQ WKH ZHVWHUQ FRQWLJXRXV
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV
Fig. 1. 5HSUHVHQWDWLYH F\DQREDFWHULD RI KRW VSULQJV A. &RFXOWXUH RI Synechococcus FI lividus WKHUPRW\SH , FXOWXUH +V IURP +XQWHU¶V+RW6SULQJV 25 SOXVILODPHQWRXVJUHHQQRQVXOIXUEDFWHULXP Chloroflexus aurantiacus FXOWXUH-IO :LGWKRISynechococcus LVaP B. &ROOHFWHGFHOOVIURP&DW.DKQHHWD+RW6SULQJV25 7KHQDUURZURGVaPZLGH DUHSynechococcus FI lividus. 7KH ZLGHU VTXDW FHOOV DUH Cyanothece (Synechococcus) FI minervae. C. Oscillatoria FI terebriformis FROOHFWHG IURP +XQWHU¶V +RW 6SULQJV 25 7ULFKRPH ZLGWK LV a P D. &ROOHFWLRQ RI Spirulina FI labyrinthiformis IURPa& DW 0DPPRWK +RW 6SULQJV
8QLWHG 6WDWHV VR IDU QRW IRXQG LQ $ODVND DQG SUREDEO\VRXWKLQWR&HQWUDODQG6RXWK$PHULFDDUH DEVHQWLQWKHJHRJUDSKLFUHJLRQV PHQWLRQHG DERYH DOWKRXJKWKH\DSSDUHQWO\H[WHQGLQWRHDVWHUQ$VLDDQG SRVVLEO\ FHQWUDO $IULFD &DVWHQKRO] ,Q FRQWUDVW VRPHWKHUPRSKLOLFF\DQREDFWHULD VXFK DV Mastigocladus FI laminosus )LJ I DQGWKHKLJK WHPSHUDWXUHIRUP+7) RI³Chlorogloeopsis´)LJ J DSSHDUWREHFRVPRSROLWDQLQGLVWULEXWLRQ7DEOH &DVWHQKRO]
% 'LVWULEXWLRQ'HWHUPLQHGE\&KHPLVWU\ ,Q KRW VSULQJV ZRUOGZLGH F\DQREDFWHULD DUH QRW REVHUYHG EHORZ S+ DQG WKHLU GLYHUVLW\ VHHPV TXLWH OLPLWHG EHORZ S+ %URFN ,Q
'0:DUGDQG5:&DVWHQKRO]
Table 1. 5HSUHVHQWDWLYHF\DQREDFWHULDRIKRWVSULQJVEDVHGRQPRUSKRORJ\DQGSK\VLRORJ\
)LHOG REVHUYDWLRQV QRW IURP VWXGLHV RI VXOILGH HIIHFWV RQ JURZWK RI SXUH FXOWXUHV
7KH KLJK WHPSHUDWXUH IRUP RI Synechococcus FI lividus PD\ FRQVLVW RI PRUH WKDQ RQH VSHFLHV 6WXGLHV ZHUH FDUULHG RXW LQ
0DQ\ ORRNDOLNH
VSHFLHV RI ORZHU WHPSHUDWXUH IRUPV DSSDUHQWO\ H[LVW DOWKRXJK VRPH VKRZ VRPH PRUSKRORJLFDO GLIIHUHQFHV )LJ LQ &DVWHQKRO]
)RUPHUO\ UHIHUUHG WR DV "Mastigocladus +7) $OWKRXJK WKLV ZLGHVSUHDG F\DQREDFWHULXP KDV D PRUSKRORJLFDO OLIH F\FOH VLPLODU WR &KORURJORHRSVLVIULWVFKLL 6 U51$ VHTXHQFH GDWD VXJJHVW WKDW D VHSDUDWH JHQHULF GHVLJQDWLRQ ZLOO EH UHTXLUHG $ :LOPRWH SHUV FRPP
$FFRUGLQJ WR 6 U51$ VHTXHQFH GDWD PRUH WKDQ RQH VSHFLHV PD\ EH LQYROYHG 5 5LSSND SHUV FRPP
7KLV LV QRW D FORVH SK\ORJHQHWLF UHODWLYH RI RWKHU UHSUHVHQWDWLYHV RI WKH ODUJH IRUPJHQXV
Oscillatoria :LOPRWWH $OVR LW LV DSSDUHQWO\ QRW UHODWHG WR WKH SODQNWRQLF Oscillatoria (Limnothrix) amphigranulata, LWV PRUSKRORJLFDO QDPHVDNH 6HH &DVWHQKRO]
6HYHUDO DSSDUHQW VSHFLHV RI Calothrix RFFXU LQ RU DURXQG WKH HGJHV RI KRW VSULQJV 7KH VSHFLHV LQ +XQWHU V +RW 6SULQJV JURZV XS WR D WHPSHUDWXUH RI a & :LFNVWURP DQG &DVWHQKRO]
JURXSJUHZDWPD[LPDOUDWHVRQO\DWS+OHYHOVDERYH S+ DQGWKXVDSSHDUHGDFLGRWROHUDQWQRWDFLGRSKLOLF .DOODVDQG&DVWHQKRO] DE 7KHDFWXDOS+ ZLWKLQF\DQREDFWHULDOPDWVPD\YDU\DERYHDQGEHORZ WKHS+RIRYHUO\LQJZDWHUGXHWRSKRWRV\QWKHWLF&2 XSWDNHDQGUHVSLUDWRU\&2 SURGXFWLRQVHHEHORZ 7HPSHUDWXUH LQFRPELQDWLRQZLWKDYDLODELOLW\RI FRPELQHG QLWURJHQ DQGRU FRQFHQWUDWLRQ RI IUHH VXOILGH LH +6 +6 6 DOVR GHWHUPLQHV F\DQREDFWHULDO VSHFLHV FRPSRVLWLRQ :KHQ WKH RXWIORZV RI QHXWUDO WR DONDOLQH QRQVXOILGLF KRW
VSULQJVWKDWFRQWDLQFRPELQHGQLWURJHQKDYHFRROHGWR & WKHUH LV WKH OLNHOLKRRG DW OHDVW LQ VRPH JHRJUDSKLF UHJLRQV WKDW D KLJK WHPSHUDWXUH IRUP +7) RISynechococcus ZLOOEHSUHVHQWDVDELRILOP RUPDWZKLFKPD\ LQWXUQ LQIOXHQFHWKHFKHPLVWU\ GRZQVWUHDP ZKHUH RWKHU VSHFLHV RI F\DQREDFWHULD HQWHU WKH WKHUPDO JUDGLHQW )RU H[DPSOH WKH FRPELQHG QLWURJHQ XVXDOO\ DV 1+ LQ WKH VSULQJ VRXUFHPD\EHODUJHO\UHPRYHGE\WKHQRQQLWURJHQ IL[LQJ Synechococcus ZKLFKPD\WKHQEHVXFFHHGHG GRZQVWUHDP EHORZ a& E\ 1IL[LQJ
&KDSWHU &\DQREDFWHULDLQ*HRWKHUPDO +DELWDWV F\DQREDFWHULD PRVW FRPPRQO\ KHWHURF\VWRXV Mastigocladus (Fischerella) FI laminosus EXW SRVVLEO\ ³+7) Chlorogloeopsis” EHORZ a& RU Calothrix VSS )LJ OK EHORZ a& 1LWURJHQ IL[DWLRQ KDV EHHQ PHDVXUHG EHORZ a& LQ KRW VSULQJ F\DQREDFWHULDO PDWV DQG HVSHFLDOO\ DW ORZHU WHPSHUDWXUHV ZKHUH QLWURJHQIL[LQJ Calothrix RFFXUV 6WHZDUW :LFNVWURP ,Q FRQWUDVW D VSULQJ PD\ EHULFK HQRXJK LQ FRPELQHG QLWURJHQ WR H[FOXGH Mastigocladus LQ IDYRU RI Synechococcus HJ DQG RWKHU QRQ1 IL[LQJ F\DQREDFWHULD Phormidium: )LJ OH ZKLFK PD\GRPLQDWH DWORZHU WHPSHUDWXUHV ,QRWKHUFDVHVFRPELQHGQLWURJHQPD\ EH YHU\ ORZ DWWKH VRXUFH HYHQ LQ KLJKWHPSHUDWXUH VSULQJV DQGKHWHURF\VWRXVF\DQREDFWHULD PD\EHWKH XSSHUWHPSHUDWXUH VSHFLHV DW WKH VSHFLILF XSSHU WHPSHUDWXUH OLPLW IRU JURZWK HJ +7) Chlorogloeopsis DW&Mastigocladus DW& 0DQ\ QHXWUDO WR DONDOLQH JHRWKHUPDO VSULQJV FRQWDLQ SULPDU\ VROXEOH VXOILGH LQ WKH VRXUFH ZDWHU 6XOILGHLVDQHIIHFWLYHLQKLELWRURISKRWRV\QWKHVLV DQG SRVVLEO\RWKHUSK\VLRORJLFDOSURFHVVHVLQWKHPDMRULW\ RIF\DQREDFWHULDEXWPD\EHXVHGDVDSKRWRV\QWKHWLF HOHFWURQ GRQRU LQ VXOILGHWROHUDQW VSHFLHV &RKHQ HW DO &DVWHQKRO] 8WNLOHQ 6HH &KDSWHU 3UHVHQWHYLGHQFH LQGLFDWHV WKDW QR WKHUPRSKLOLF F\DQREDFWHULD ZLWK WKH FDSDFLW\ WR JURZ DERYH & DUHFDSDEOHRIJURZLQJ LQZDWHUVZLWKPRUHWKDQ a 0 VXOILGH &DVWHQKRO] *DUFLD3LFKHO DQG &DVWHQKRO] ,Q VXOILGHULFK VSULQJV RI 1HZ =HDODQG WKH XSSHUWHPSHUDWXUH VXOILGHWROHUDQW DQG VXOILGHXWLOL]LQJ F\DQREDFWHULXP LV DQ Oscillatoria FI amphigranulata PRUSKRW\SH &DVWHQKRO] ,QVRPHKRWVSULQJVRI0DPPRWK 7HUUDFHV
VXUIDFH ZDWHU VXOILGH GLVDSSHDUV a& &DVWHQKRO]E*LRYDQQRQLHW DO ,Q D IHZ LQWHUHVWLQJ ,FHODQGLF PDWV VXOILGHR[LGL]LQJ SKRWRV\QWKHWLF EDFWHULD RQ WKH PDW VXUIDFH ZKLFK UHVHPEOH Chloroflexus aurantiacus, UHPRYH VXOILGH DQG SHUPLW VXOILGHVHQVLWLYHF\DQREDFWHULDWR JURZ EHQHDWK WKHP LQ WKH ORZHU SDUW RI WKH SKRWLF ]RQH -¡UJHQVHQ DQG 1HOVRQ
& :HOO6WXGLHG0DW 6\VWHPV +XQWHU V+RW6SULQJ2UHJRQ 7KH SULYDWHO\ RZQHG +XQWHU V +RW 6SULQJ NP QRUWK RI /DNHYLHZ 2UHJRQ HOHY P ZHUH VWXGLHG SHULRGLFDOO\ E\ &DVWHQKRO] DQG VWXGHQWV VLQFH WKH HDUO\ V 7KH\ FRQVLVW RI D VHULHV RI VSULQJV LQLWLDWLQJ IURP DQ DFWLYH GHHS IDXOW V\VWHP PDQ\ LVVXLQJ DW WHPSHUDWXUHV DERYH & DW D S+ RI 7KH PDMRU LRQV DUH VRGLXP a PJ / FKORULGH a PJ / VLOLFDWH a PJ / ELFDUERQDWH a PJ / DQG VXOIDWH PJ 7KH / VSHFLILF FRQGXFWDQFH RI a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ynechococcus lividus &RSHODQG)LJ D RFFXUV DV WKH WRS FRYHU RI WKHPDWLQ PRVW RI WKH+XQWHU V GUDLQZD\V WR DERXW & ZKHUH LW LV DEUXSWO\ UHSODFHG E\ D GDUN UHG EURZQ FRYHU RI Oscillatoria FI terebrijormis )LJ F D GLVWLQFWLYH IRUP LGHQWLILHG E\ PRUSKRORJ\ SK\VLRORJ\ DQG HFRORJ\ &DVWHQKRO] )LJ DQG 3ODWH D 7KH XQLIRUP JUHHQ FRYHU RI Synechococcus, KRZHYHU ZDV IRXQG WR FRQVLVW RI DW OHDVW VWDEOH WHPSHUDWXUHGHILQHG HFRW\SHV ZKLFK
'0:DUGDQG5:&DVWHQKRO]
Fig. 2. 2Q WKH OHIW LGHDOL]HG KRUL]RQWDO GLVWULEXWLRQ RI PDW VXUIDFH RUJDQLVPV LQ RXWIORZV RI +XQWHU V +RW 6SULQJV 2UHJRQ ZLWK V\PEROV GHSLFWLQJ WKH SUHGRPLQDQW VSHFLHV 2Q WKH ULJKW WKH DSSUR[LPDWH JURZWK UDWHV LQ FXOWXUH RI WKH SULQFLSDO F\DQREDFWHULDO VSHFLHV UHODWLYH WR WHPSHUDWXUH 7KH URPDQ QXPHUDOV RI 6 FI lividus UHIHU WR WKH GHVLJQDWLRQV LQ 3HDU\ DQG &DVWHQKRO] Potamocypris VS LV WKH DV \HW XQQDPHG WKHUPRSKLOLF RVWUDFRG LQ :LFNVWURP DQG &DVWHQKRO]
DSSHDUHG PRUSKRORJLFDOO\ LGHQWLFDO 3HDU\ DQG &DVWHQKRO] )LJ ,Q FXOWXUH WKH PRVW WKHUPRWROHUDQW FORQH JUHZ DW D PD[LPXP UDWH DW ZLWK WHPSHUDWXUHV RYHU WR EHLQJ VXSUDRSWLPDO 0HHNV DQG &DVWHQKRO] )LJ ,Q FXOWXUH ZLWK FRQWLQXRXV LOOXPLQDWLRQ WKH PD[LPXP JURZWK UDWH RI WKLV VWUDLQ ZDV DERXW GRXEOLQJV SHU K D UDWH FRQVLGHUDEO\ VORZHU WKDQ WKHORZHUWHPSHUDWXUHVWUDLQV)LJ Oscillatoria FI terebriformis KDV DQ XSSHU JURZWK WHPSHUDWXUH OLPLW RI & LQ FXOWXUH DQG JURZV DW D PD[LPDO UDWH WR & RI WKLVOLPLW&DVWHQKRO] D )LJ 6LQFH LW LV KLJKO\ PRWLOH E\ JOLGLQJ WKH XSSHU HGJH RI WKH PDW DGMXVWV LWVSRVLWLRQWR LWV XSSHU OLPLW3ODWH D DQG E 6LQFH WKH PDW LV JHQHUDOO\ WKLFN HQRXJK WR DEVRUE RYHU RI YLVLEOH UDGLDWLRQ LWV JURZWK DEUXSWO\ OLPLWV WKH YLVLEOH ORZHU HGJH RI WKH Synechococcus FRYHU ZKLFK LV RQO\ VOLJKWO\ PRWLOH SUHVXPDEO\ EHFDXVH O. FI terebriformis RXWFRPSHWHV RWKHU F\DQREDFWHULD IRU OLJKW $OO FORQHV RI O. FI terebriformis KDG UHVSHFWDEOH JURZWK UDWHV ZHOO EHORZ & DOWKRXJK LQ PRVW RI WKH VSULQJV RI (
2UHJRQ DQG 1 1HYDGD WKH Oscillatoria PDW HQGHG DEUXSWO\ DW DERXW& )LJ 3ODWHD 7KLV LV D UHVXOW RI YRUDFLRXV SRSXODWLRQV RI WKH WKHUPRSKLOLF DQG KHUELYRURXV RVWUDFRG Potamocypris VS WKDW KDYH DQ DOPRVW XELTXLWRXV GLVWULEXWLRQ LQ WKH VDPH JHRJUDSKLF UHJLRQ &DVWHQKRO] D :LFNVWURP DQG &DVWHQKRO] )LJ 3ODWHV E F Potamocypris FDQ VXUYLYH DQG UHSURGXFH DW WHPSHUDWXUHV DV KLJK DV & :LFNVWURP DQG &DVWHQKRO] DQG ZLOO HOLPLQDWH WKH GHOLFDWH WULFKRPHV RI Oscillatoria DW D UDSLG UDWH $V D FRQVHTXHQFH WKH F\DQREDFWHULDO SRSXODWLRQV EHORZ WKLV WHPSHUDWXUH DUH FRPSRVHG SULPDULO\ RI WKH PRUH JUD]HUUHVLVWDQW VSHFLHV Pleurocapsa VS DQG Calothrix VS )LJ OK 7KH WDSHUHG ILODPHQWV RI Calothrix DUH HPEHGGHG ZLWKLQ WKH VHHPLQJO\ DPRUSKRXV PDVV RI Pleurocapsa FHOOV :LFNVWURP DQG &DVWHQKRO] 7KH RVWUDFRGV DSSHDU WR JUD]H SULPDULO\ RQ WKH H[SRVHG ODZQ RI Calothrix ILODPHQW WLSV ZKLFK FRQWLQXH WR JURZ WKURXJK WKH Pleurocapsa DV ZHOO DV RQ WULFKRPH PDVVHV RI FI terebriformis ZKLFK DUH IUHTXHQWO\ ZDVKHG
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
2. Octopus Spring, Yellowstone National Park 2FWRSXV 6SULQJ LV ORFDWHG LQ WKH :KLWH &UHHN GUDLQDJH RIWKH /RZHU *H\VHU %DVLQ
WHPSHUDWXUHV WR D PD[LPXP RI VHYHUDO FP DV WHPSHUDWXUH GHFUHDVHV DORQJ WKH HIIOXHQW FKDQQHO DOWKRXJK PDW IRUPDWLRQ DQG GHFRPSRVLWLRQ LV SUREDEO\ PD[LPL]HG LQ WKH XSSHU IHZ PLOOLPHWHUV %URFN :DUG HW DO %HORZ a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
PDW SRSXODWLRQV ZHUH DOVR H[DPLQHG 6LQFH F\DQREDFWHULD DUH SUHGRPLQDQW PHPEHUV RI WKHVH PDW FRPPXQLWLHV LW ZDV QRW VXUSULVLQJ WKDW WKHLU 6 U51$ VHTXHQFHV DQG OLSLG ELRPDUNHUV ZHUH UHDGLO\ GHWHFWHG LQ PROHFXODU DQDO\VHV ,GHQWLILFDWLRQ RI 6 U51$ VHTXHQFHV DV EHLQJ RI F\DQREDFWHULDO RULJLQ ZDV IDFLOLWDWHG E\ WKH PRQRSK\OHWLF HYROXWLRQDU\ GHVFHQW RI WKH R[\JHQLF SKRWRWURSKLF SURNDU\RWHV *LRYDQQRQL HW DO :LOPRWWH
$ 'HWHFWLRQ RI 6U51$'HILQHG &\DQREDFWHULDO 3RSXODWLRQV $W WKH RXWVHW RI PROHFXODU VWXGLHV LW ZDV WKRXJKW WKDW D VLQJOH ZLGHVSUHDG 6\QHFKRFRFFXV VSHFLHV 6 =LYLGXV PD\ KDYH FRQVWUXFWHG WKH 6\QHFKRFRFFXV W\SH KRW VSULQJ PDWV ZKLFK DUH FRPPRQ LQ ZHVWHUQ 1RUWK $PHULFD %URFN &DVWHQKRO] D *HQHWLFDOO\VWDEOH WHPSHUDWXUHVSHFLDOL]HG ³VWUDLQV´ ZHUH FXOWLYDWHG IURP +XQWHU¶V 6SULQJV VHH 6HFWLRQ ,,& EXWWKH JHQHWLF GLIIHUHQFHV DPRQJ WKH ³VWUDLQV´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
'0:DUGDQG 5:&DVWHQKRO] % *HQHWLF'LYHUVLW\$PRQJ 6U51$ 'HILQHG &\DQREDFWHULDO 3RSXODWLRQV 7KH JHQHWLF GLYHUVLW\ DPRQJ 6 U51$GHILQHG 2FWRSXV 6SULQJ F\DQREDFWHULDO SRSXODWLRQV LV LPSUHVVLYH WKH YDULDWLRQ DPRQJ WKHLU 6 U51$ VHTXHQFHV LV VLPLODU WR WKDW REVHUYHG IRU DOO FXOWXUHG F\DQREDFWHULDO VSHFLHV )LJ 6L[ SRSXODWLRQV W\SHV $ $ $
$
% DQG %¶ DUH FORVHO\ UHODWHG H[KLELWLQJOHVVWKDQ GLIIHUHQFH LQ WKHLU 6 U51$ VHTXHQFHV \HW DUH HFRORJLFDOO\ GLVWLQFW VHH EHORZ 7KHVH RFFXU DV WZR GLVWLQFW SK\ORJHQHWLF FOXVWHUV UHIHUUHG WR DV $OLNH W\SH $ $ $
DQG $
DQG % OLNH W\SHV % DQG % F\DQREDFWHULDO VHTXHQFHV *LYHQWKDW 6 U51$ VHTXHQFHV DUHKLJKO\FRQVHUYHG :RHVH LW LV SRVVLEOH WKDW RWKHU WKHUPRSKLOLF F\DQREDFWHULDO HFRW\SHV PD\ H[LVW ZKLFK FDQ RQO\ EH GLVFRYHUHG XVLQJ FRPSDULVRQV RI PRUH UDSLGO\ HYROYLQJJHQHWLFHOHPHQWV 7KH SUHYLRXV LGHD WKDW WKHUPRSKLOLF Synechococcus H[KLELWHG OLPLWHG JHQHWLF GLYHUVLW\ PD\ KDYH UHVXOWHG IURP WKH VWURQJ VHOHFWLRQ IRU D VLQJOH JHQRW\SH LQ OLTXLG HQULFKPHQWFXOWXUH HJ VWUDLQ & LQ 7DEOH $V VKRZQ E\ )HUULV HW DO E DOO FXOWXUH FROOHFWLRQ VWUDLQV RI WKHUPRSKLOLF Synechococcus FI lividus KDG YHU\ VLPLODU LI QRW LGHQWLFDO 6 U51$ VHTXHQFHV 6XFFHVVIXO HQULFKPHQW DQG FXOWLYDWLRQ RI D VWUDLQ ZLWK WKH VDPH JHQRW\SH & YHULILHG LWV SUHVHQFH LQ WKH 2FWRSXV 6SULQJ PDW DSSDUHQWO\ DW D GHQVLW\ WRR ORZ WR EH GHWHFWHG E\ GLUHFW PROHFXODU DQDO\VLV )HUULV HW DO E 'LOXWLRQ RI LQRFXOXP WR QHDU H[WLQFWLRQ EHIRUH HQULFKPHQW SUHYHQWHG FRPSHWLWLYHH[FOXVLRQRI RWKHU Synechococcus VSS E\ WKH & SRSXODWLRQ DQG OHG WR WKH UHFRYHU\ RI PRUH QXPHULFDOO\ UHOHYDQW DQG QRYHO 6\QHFKRFRFFXV JHQRW\SHV )HUULV HW DO E :DUG HW DO ,VRODWHV IURP KLJKO\ GLOXWHG LQRFXOD FRQWDLQLQJ DSSUR[LPDWHO\ WR 6\QHFKRFRFFXV FHOOV LQFOXGHG WKUHH VWUDLQV 6HTXHQFHV IURP WKHVH WKUHH VWUDLQV ZHUH GHWHFWHG LQ WKH PDW % % 3 YHULI\LQJ WKDW PXOWLSOH Synechococcus SRSXODWLRQV H[LVW LQ WKH PDW 2QH Synechococcus LVRODWH & ZDV FXOWLYDWHG IURP DQ HQULFKPHQW LQRFXODWHG ZLWK DSSUR[LPDWHO\ Synechococcus FHOOV DQG PD\ EH D VXEGRPLQDQW SRSXODWLRQ LWV 6 U51$ VHTXHQFH KDV \HW WR EH GHWHFWHG LQ WKH PDW WKURXJK PROHFXODU DQDO\VLV 7KH FXOWLYDWLRQ RI VWUDLQV & DQG & UDLVHG WR HOHYHQ WKH QXPEHU RI JHQRW\SLFDOO\ XQLTXH F\DQREDFWHULDO SRSXODWLRQV GHWHFWHG LQ WKH 2FWRSXV 6SULQJ PDW DW OHDVW ILYH RI ZKLFK DUH NQRZQ WR H[KLELW D S. FI 7KH IRXU lividus-like PRUSKRORJ\ 7DEOH
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV
Table 2. 8QLTXHF\DQREDFWHULDO6U51$VHTXHQFHW\SHVIRXQGLQWKH2FWRSXV6SULQJODPLQDWHGF\DQREDFWHULDOPDWRULQF\DQREDFWHULD LVRODWHVIURPWKHPDW
6HTXHQFHGHWHFWHGE\FORQLQJRU'**( 5HIHUHQFHV )HUULVHWDO D )HUULVDQG:DUG :DUGHWDO 5DPVLQJHWDO LQ SUHSDUDWLRQ )HUULVHW DO :DUGHWDO :HOOHUHWDO 5XII5REHUWVHWDO )HUULVHWDOD 5DPVLQJHWDO VWUDLQORVWEXWOLPLWHG'1$PD\EHDYDLODEOHLQWKHODERI'0: SDUWLDO 6U51$VHTXHQFHLGHQWLFDOWRWKRVHRIDOOWKHUPRSKLOLF6\QHFKRFRFFXVFXOWXUHFROOHFWLRQVWUDLQVVHH)HUULVHWDO EIRU$77& 1RV
XQFXOWLYDWHG $OLNH SRSXODWLRQV PD\ H[KLELW WKLV PRUSKRORJ\ EHFDXVHRQO\S. FI OLYLGXVOLNHFHOOVZHUH REVHUYHG E\ PLFURVFRS\ LQ WKH PDW DW WKH KLJK WHPSHUDWXUHV ZKHUH WKHVH SRSXODWLRQV ZHUH GHWHFWHG 6WUDLQV & DQG & ZHUH JHQHWLFDOO\ GLVVLPLODU IURP SRSXODWLRQV GHWHFWHG E\ PROHFXODU PHDQV 6\QHFKRFRFFXV VWUDLQ & LVFXUUHQWO\WKHPRVW GHHSO\ GLYHUJHQW F\DQREDFWHULXP NQRZQ WKRXJK WKH VWDWLVWLFDO VXSSRUW IRU WKLV LQIHUHQFH LV QRW VWURQJ )HUULV HW DO E
& 'LVWULEXWLRQRI6U51$'HILQHG &\DQREDFWHULDO 3RSXODWLRQV 7KH GLVWULEXWLRQV RI VRPH RI WKHVH F\DQREDFWHULDO SRSXODWLRQV ZHUH VWXGLHG XVLQJ ROLJRQXFOHRWLGH
K\EULGL]DWLRQSUREHVDQG'**(
7HPSHUDWXUH'LVWULEXWLRQ $QDO\VHV XVLQJ '**( VXJJHVW WKDW ILYH FORVHO\ UHODWHG F\DQREDFWHULDO SRSXODWLRQV $
$ $ % DQG % RFFXU DORQJWKHWKHUPDO JUDGLHQWIURPKLJK WR ORZ WHPSHUDWXUH UHVSHFWLYHO\ )LJ +\EULGL]DWLRQ SUREHVWXGLHVVXJJHVWHGWKDWWKHW\SH-SRSXODWLRQZDV DOVR IRXQG DW WKHORZHVWWHPSHUDWXUHV5XII5REHUWVHW DO +\EULGL]DWLRQ SUREH H[SHULPHQWV DOVR GHPRQVWUDWHG WKDW GXULQJ D RQHZHHN LQ VLWX LQFXEDWLRQ LQ ZKLFK VDPSOHV ZHUH VKLIWHGIURPORZ WR KLJK WHPSHUDWXUH $OLNH 6 U51$ LQFUHDVHG ZKLOH %OLNH DQG - 6 U51$ GHFUHDVHG VXJJHVWLQJ WKDW WKHVH GHULYH IURP SRSXODWLRQV ZLWK GLIIHUHQW
'0:DUGDQG5:&DVWHQKRO]
Fig. 3 'LVWDQFH PDWUL[ SK\ORJHQHWLF WUHH RI F\DQREDFWHULDO 6 U51$ VHTXHQFHV GHWHFWHG LQ WKH 2FWRSXV 6SULQJ F\DQREDFWHULDO PDW RU F\DQREDFWHULDO LVRODWHV IURP WKH PDW WKLFN OLQHV FRPSDUHG WR UHSUHVHQWDWLYH F\DQREDFWHULDO OLQHDJHV WKLQ OLQHV $ WUHH ZLWK DOO OLQHV H[FHSW2FWRSXV 6SULQJW\SHV $ $ $
$
DQG% ZDV ILUVWFRQVWUXFWHGXVLQJQXFOHRWLGHVWKDW DOLJQZLWK ( FROLSRVWLRQV WR WRWRDQG WR WKHWUHHZDVURRWHGZLWK WKH 6 U51$VHTXHQFHV RIThermotoga maritima, Chlorobium vibrioforme DQG Escherichia coli. 7KH FRQVHQVXV YDOXHV DWWKHIRUNVLQGLFDWHWKHQXPEHURIWLPHV WKHJURXSFRQVLVWLQJ RIWKH VHTXHQFHV WR WKH ULJKW RI WKH IRUNRFFXUUHG DPRQJ WUHHV LQIHUUHG IURPWKHERRWVWUDSSHG GDWDVHWRIQXFOHRWLGHV $WUHH IRUWKHUHPDLQLQJ VHTXHQFHV ZKLFK DOVR LQFOXGHG WKHLU FORVHVW UHODWLYH WKH W\SH % VHTXHQFH ZDV FRQVWUXFWHG XVLQJ ( FROL SRVLWLRQV WR 7KH WZR WUHHV ZHUH FRQQHFWHG DW WKH SRLQWZKHUH WKH W\SH % VHTXHQFHGLYHUJHV IURP RWKHUVHTXHQFHV LQ WKH ILUVW WUHH DUURZ %DULQGLFDWHV IL[HG SRLQW PXWDWLRQVSHUVHTXHQFHSRVLWLRQ
WHPSHUDWXUH RSWLPD 5XII5REHUWV HW DO 7KHVH GDWD UHLQIRUFHG WKH FRQFHSW RI WHPSHUDWXUH DGDSWHG ³VWUDLQV´ LQWURGXFHGE\ 3HDU\ DQG&DVWHQKRO] )LJ :KHWKHU WKHVH VKRXOG EH FRQVLGHUHG VWUDLQV RI RQH VSHFLHV RUGLIIHUHQW VSHFLHV GHSHQGV RQ RQH¶V FRQFHSW RI ³VSHFLHV´ &DVWHQKRO] 7HPSHUDWXUH GLVWULEXWLRQ GDWD VXJJHVW WKDW WKHVH SRSXODWLRQV UHSUHVHQW GLIIHUHQW HFRW\SHV DQG HFRORJLFDOO\ GLVWLQFW DVH[XDO SRSXODWLRQV DUH VRPHWLPHV FRQVLGHUHG WR EH VHSDUDWH VSHFLHV 0D\U 7KHRQHVXUYLYLQJWHPSHUDWXUH³VWUDLQ´ 2+ V 3&& DQG $7&& FXOWLYDWHG IURP +XQWHU¶V 6SULQJV E\ 3HDU\DQG&DVWHQKRO] KDG D Synechococcus &OLNH JHQRW\SH )HUULV HW DO E 2QH SRVVLEOH LQIHUHQFH LV WKDW WHPSHUDWXUH DGDSWHG HFRW\SHV RI WKH Synechococcus & JHQRW\SH H[LVW LQ +XQWHU¶V +RW 6SULQJV PDWV 7KLV ZRXOG EH VLJQLILFDQW DV LW ZRXOG LQGLFDWH WKDW WHPSHUDWXUH DGDSWHG SRSXODWLRQV HYROYHG LQ WZR ZLGHO\VHSDUDWHG F\DQREDFWHULDO OLQHDJHV )LJ $OWHUQDWLYHO\ DV LQ WKH 2FWRSXV6SULQJPDW Synechococcus JHQRW\SH & PD\ EH D PLQRU SRSXODWLRQ LQ +XQWHU¶V +RW 6SULQJV PDWV DQG LWV UHFRYHU\ PD\ EH IDYRUHG E\ OLTXLG HQULFKPHQW FXOWXUH IURP ORZGLOXWLRQ LQRFXOD :KLOH DGDSWLYH UDGLDWLRQ WR WHPSHUDWXUH PD\ DW OHDVW
SDUWLDOO\ H[SODLQ WKH HYROXWLRQ RI WKH $ DQG %OLNH JHQRW\SHV WKH HYROXWLRQDU\ SUHVVXUHV ZKLFK FDXVHG GLYHUJHQFHV DPRQJ WKH PRUH GHHSO\ EUDQFKLQJ OLQHV RI WKHUPRSKLOLF F\DQREDFWHULDO GHVFHQW HJ $% OLNH & & , - DQG 3 UHPDLQ REVFXUH ZLWKRXW IXUWKHU GLVWULEXWLRQ DQDO\VLV RU SKHQRW\SLF FKDUDFWHUL]DWLRQ
9HUWLFDO'LVWULEXWLRQ 6HYHUDOW\SHVRI REVHUYDWLRQVXJJHVWWKDWRQHSRVVLEOH HYROXWLRQDU\VWLPXOXV IRUDGDSWDWLRQLVLUUDGLDQFHHJ 3LHUVRQ DQG &DVWHQKRO] LQFOXGLQJ ERWK LQWHQVLW\ DQG TXDOLW\ RI YLVLEOH DQGRU XOWUDYLROHW UDGLDWLRQ Synechococcus ULFK LQ FKORURSK\OO D RFFXU XQGHUQHDWK WKH XSSHUPRVW Synechococcus OD\HU %URFN ZKLFK LV PRUH \HOORZ 0LFURVFRS\ VWXGLHV RI YHUWLFDO WKLQ VHFWLRQV RI WKLV OD\HU LQ WKH & PDW RI 0XVKURRP 6SULQJ UHYHDOHG YHUWLFDO VWUDWLILFDWLRQ RISynechococcus SRSXODWLRQV 3ODWH 6WHHS OLJKW JUDGLHQWV ZHUH PHDVXUHG LQ VXFK PDWV )LJ &\DQREDFWHULDDVPHDVXUHG E\ FKORURSK\OOD DQG SK\FRF\DQLQ DEVRUSWLRQ WURXJKV DQG LQWHUYHQLQJ SHDNV DSSHDUHG WR RFFXU PDLQO\ LQ WKH XSSHU PP 7KH SUHVHQFH RI EDFWHULFKORURSK\OO cFRQWDLQLQJ
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV
7HPSHUDWXUH& Fig. 4 7HPSHUDWXUHGLVWULEXWLRQRIF\DQREDFWHULDO6 U51$ VHTXHQFHW\SHV GHWHFWHG LQ WKH 2FWRSXV 6SULQJF\DQREDFWHULDO PDWE\ '**( DQDO\VLV ,QWHQVLWLHVRI'**(EDQGVZHUHGHWHUPLQHGE\LQWHJUDWLQJVFDQQHGJHOLPDJHVDQGDGGLQJWKHLQWHQVLW\ RIKRPRGXSOH[ DQGRQH KDOIRIWKHLQWHQVLW\RIKHWHURGXSOH[EDQGVFRQWDLQLQJHDFKVHTXHQFHW\SH >VHH)HUULVDQG:DUG@
RUJDQLVPV HJ &KORURIOH[XV VSS RU JUHHQ VXOIXU EDFWHULD DQG EDFWHULFKORURSK\OO a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
$QRWKHUSRSXODWLRQ W\SH& SK\ORJHQHWLFDOO\ UHODWHG WR JUHHQ QRQVXOIXUEDFWHULDHJ &KORURIOH[XV ZDV GHWHFWHG LQ LQFUHDVLQJ DPRXQWV EHORZ a PP 5DPVLQJHWDOXQSXEOLVKHGGDWD 2WKHU NLQGV RI HYLGHQFH DUH FRQVLVWHQW ZLWK WKH SRVVLELOLW\ WKDW JHQHWLFDOO\ GLVWLQFW Synechococcus SRSXODWLRQV DUH DGDSWHG WR GLIIHUHQW OLJKW LQWHQVLWLHV )RU H[DPSOH K\EULGL]DWLRQ SUREH VWXGLHV UHYHDOHG GLIIHUHQWLDO UHVSRQVHV RI PDW SRSXODWLRQV WR UHGXFHG OLJKW LQWHQVLW\ 5XII5REHUWV HW DO $OVR JHQHWLFDOO\ GLVWLQFWSynechococcus FI lividus LVRODWHV H[KLELWHG GLIIHUHQW SKRWRWDFWLF PRYHPHQWV LQ UHVSRQVHWROLJKWRIGLIIHUHQWLQWHQVLW\5DPVLQJHWDO )XUWKHUPRUH OLJKWDGDSWHG HFRW\SHV RI WKH PRGHUDWHO\ WKHUPRSKLOLF F\DQREDFWHULXP Plectonema VS ZHUH GHVFULEHG 6KHULGDQ ,W LV K\SRWKHVL]HG WKDW Synechococcus SRSXODWLRQV ZKLFK DUH DGDSWHG WR GLIIHUHQW OLJKW LQWHQVLWLHV DQGRU XOWUDYLROHW UDGLDWLRQ H[LVW DORQJ WKH WKHUPDO JUDGLHQW
'0:DUGDQG 5: &DVWHQKRO]
Fig. 5 . /RJDULWKPLF SORW RI GRZQZDUG LUUDGLDQFH LQ & 2FWRSXV 6SULQJ ODPLQDWHG Synechococcus PDW PHDVXUHG ZLWK D /,&RU VSHFWURUDGLRPHWHU DQG D PP GLDPHWHU ILEHU RSWLF WLS VHH 3LHUVRQ HW DO 'HSWK RIHDFK WUDFH PDUNHG LQ PP EHORZ VXUIDFH 'HSUHVVLRQV LQVSHFWUXPDW aQPSK\FRF\DQLQ a QP FKORURSK\OOD aQPEDFWHULRFKORURSK\OO F a DQG QP EDFWHULFKORURSK\OOD >NLQGO\SURYLGHGE\%.3LHUVRQ@
VR WKDW DGDSWDWLRQVWRWHPSHUDWXUHDQG LUUDGLDQFH PD\ H[SODLQ PXFK RI WKH JHQRW\SLF GLYHUVLW\ IRXQG LQ 6\QHFKRFRFFXV PDWV ,W LV DOVR SRVVLEOH WKDW RWKHU SDUDPHWHUV ZKLFK YDU\ YHUWLFDOO\ LQ WKH PDW ZHUH UHVSRQVLEOH IRU WKH VSHFLDOL]DWLRQ RI PDW F\DQREDFWHULDO SRSXODWLRQV
7HPSRUDO'LVWULEXWLRQ ,W ZDV VXJJHVWHG WKDW WKHUPRSKLOLF F\DQREDFWHULD DGDSWHG WR GLIIHUHQW OLJKW LQWHQVLWLHV PD\ VKLIW GRPLQDQFH LQ PDWV LQ GLIIHUHQW VHDVRQV 6KHULGDQ ,Q FRQWUDVW WKH SDWWHUQ RI WHPSHUDWXUH GLVWULEXWLRQ ZLWKLQ WKH 2FWRSXV 6SULQJ PDW FRPPXQLW\ )LJ ZDV VLPLODU LQ DOO VHDVRQV )HUULV DQG :DUG ZKLFK LPSOLHV WKDW WKHUH PD\ EH
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
ZHUH HIIHFWLYH FRORQLVWV 7KH EDVLV IRU VSHFLDOL]DWLRQ DV D FRORQLVWUHPDLQV XQNQRZQ
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV
Fig. 6. 9HUWLFDOGLVWULEXWLRQ RIF\DQREDFWHULDO 6 U51$VHTXHQFHW\SHV GHWHFWHGE\'**(DQDO\VLV RI PWKLFNFU\RWRPHVHFWLRQV RI WKHVDPHPDWVKRZQLQ)LJ ,QWHQVLWLHV RI'**(EDQGV ZHUHGHWHUPLQHGE\ LQWHJUDWLQJVFDQQHG JHO LPDJHV DQG DGGLQJ WKHLQWHQVLW\RI KRPRGXSOH[DQGRQHKDOIRIWKHLQWHQVLW\RIKHWHURGXSOH[EDQGVFRQWDLQLQJHDFKVHTXHQFHW\SH >IURP5DPVLQJHWDO@
'LVWULEXWLRQ$PRQJ6SULQJV $QRWKHU SRVVLEOH HYROXWLRQDU\ GULYLQJ IRUFH IRU GLYHUVLW\ DPRQJ WKHUPRSKLOLF F\DQREDFWHULD LV S+ 6LQFHS+PD\YDU\RYHUDGLHOSHULRGZLWKLQ DVLQJOH PDW RYHU D UDQJH RI a WR XQLWV VHH EHORZ DGDSWDWLRQV WR S+ PD\ EH OLPLWHG WR H[WUHPHV RIWKH S+ UDQJH IRXQG LQ GLIIHUHQW VSULQJV LQKDELWHG E\ WKHUPRSKLOLF F\DQREDFWHULD VHH 7DEOH 7KH 6 U51$ RI D6 FI lividus LVRODWH & JHQRW\SH ZDV UHDGLO\ GHWHFWHG LQ WKH S+ &OHDUZDWHU 6SULQJV SRRO IURP ZKLFK LW ZDV RULJLQDOO\ FXOWLYDWHG DQG ZKHUH %OLNH SRSXODWLRQV ZHUH DOVR GHWHFWHG 5XII 5REHUWVHWDO 6SULQJVDWORZHUS+HJGRZQ WRS+ WKDWFRQWDLQHG SynechococcusVKDSHG FHOOV GLGQRWFRQWDLQGHWHFWDEOH 6U51$RI WKH& RU$ OLNH %OLNH RU - F\DQREDFWHULDO SRSXODWLRQV ZKLFK PD\VXJJHVWWKHSRVVLEOHH[LVWHQFHRIORZS+DGDSWHG Synechococcus SRSXODWLRQV 5HODWLYHO\ OLWWOH LV NQRZ DERXW WKH JHRJUDSKLFDO GLVWULEXWLRQ RI F\DQREDFWHULDO SRSXODWLRQV GHILQHG WKURXJK PROHFXODU DQDO\VHV LQ QHXWUDO WR DONDOLQH VSULQJVZKLFKPD\RUPD\QRWGLIIHULQRWKHUFKHPLFDO SURSHUWLHV $V PHQWLRQHG DERYH '**( DQDO\VLV UHYHDOHG WKH VLPLODULW\ RIF\DQREDFWHULDO SRSXODWLRQV IRXQG LQ a& PDWV RI 2FWRSXV 6SULQJ DQG 0XVKURRP6SULQJERWKS+ ZKLFKDUHVHSDUDWHG E\ RQO\ D IHZ KXQGUHG PHWHUV '1$ K\EULGL]DWLRQ
DQDO\VLV UHYHDOHG WKDW QHXWUDO WR DONDOLQH VSULQJV VHSDUDWHG E\ XS WR WHQV RI NLORPHWHUV FRQWDLQHG $ OLNH S+ DQG %OLNH S+ F\DQREDFWHULDO SRSXODWLRQV DW WHPSHUDWXUHV FRPSDUDEOHWR WKRVH DWZKLFK WKH\ZHUH IRXQG LQ WKH 2FWRSXV 6SULQJ PDW W\SH- F\DQREDFWHULDO SRSXODWLRQVZHUHREVHUYHGRQO\LQWR&PDWVLQ 2FWRSXV DQG 7ZLQ %XWWH 9LVWD 6SULQJV S+ ZKLFK DUH VHSDUDWHG E\ DSSUR[LPDWHO\ PHWHUV 5XII5REHUWV HW DO :DUG HW DO D 6R IDU QR PROHFXODU DQDO\VHV ZHUH SHUIRUPHG RQ KRW VSULQJ F\DQREDFWHULDO SRSXODWLRQV RI JUHDWHU JHRJUDSKLFDO VHSDUDWLRQ VXFK JHQRW\SLF LQIRUPDWLRQ PD\ SURYH WR EH HVVHQWLDO IRU WKH HYDOXDWLRQ RI WKH LPSRUWDQFH RI DOORSDWULF VSHFLDWLRQ LQ WKHUPRSKLOLF F\DQREDFWHULD
' /LSLG%LRPDUNHU6WXGLHV /LSLG ELRPDUNHUV DSSHDU WR EH PXFK OHVV VSHFLILF WKDQ 6U51$VHTXHQFHV +RZHYHU EHFDXVHRIWKH JUHDWHU SHUVLVWHQFH RI OLSLGV RYHU JHRORJLFDO WLPH WKH\PD\EHXVHIXOIRUUHFRJQL]LQJF\DQREDFWHULDDVD JURXS DV RSSRVHG WR VSHFLILF F\DQREDFWHULDO HFRW\SHV LQ GHSRVLWV RIJHRORJLFDOO\ VLJQLILFDQW DJH )RU LQVWDQFH WKH PLGFKDLQEUDQFKHG PRQR DQG GLPHWK\O\DONDQHVIRXQGLQ3UHFDPEULDQRLOVDSSHDUWR EH W\SLFDO RI F\DQREDFWHULD IRXQG LQ WKHUPDO PDWV
'0:DUGDQG5:&DVWHQKRO]
6KLHDHWDO )UHHOLSLGV'REVRQHWDO 6KLHDHWDO SRODUOLSLGIDWW\DFLGV=HQJ HWDO DDQGE:DUGHWDO E DQGFRPSOH[ SRODUOLSLGV:DUGHWDOE W\SLFDORIWKRVHLQ6 FI lividus, GRPLQDWHOLSLGH[WUDFWVIURPWKH2FWRSXV 6SULQJPDW 7KHVHOLSLGVZHUHGLIILFXOWWRGLVWLQJXLVK IURP OLSLGV RI JUHHQ QRQVXOIXU EDFWHULD HJ& aurantiacus), EXW WKH SUHVHQFH RI VXOIRTXLQRYRV\O GLJO\FHULGHV DQG WKH SUHGRPLQDQFH RI& RYHU & IDWW\ DFLGV VXJJHVWHG WKDW F\DQREDFWHULD ZHUH WKH PDLQVRXUFH )DWW\DFLGFKDLQOHQJWKDQGGHJUHHRI VDWXUDWLRQYDULHGZLWKPDWWHPSHUDWXUH ,WLVXQFHUWDLQ ZKHWKHUWKLVUHIOHFWHGSRSXODWLRQYDULDWLRQDORQJWKH WKHUPDOJUDGLHQW)LJ DFFOLPDWLRQWRWHPSHUDWXUH RUERWK
Synechococcus FHOOV VXJJHVWWKDWWKHVHDUHOLNHO\WR EH WKH SUHGRPLQDQW 2FWRSXV 6SULQJ PDW F\DQREDFWHULD &KDQJHV LQ VSHFLILF $ DQG %OLNH F\DQREDFWHULDOSRSXODWLRQVDORQJWKHUPDODQGYHUWLFDO JUDGLHQWV DSSDUHQWO\ UHIOHFW VSHFLDOL]DWLRQ WR WHPSHUDWXUHDQGOLJKWRURWKHUSDUDPHWHUVWKDWYDU\LQ WKHSKRWLF]RQH 2WKHUF\DQREDFWHULDOSRSXODWLRQV H[LVWLQWKHPDW VRPHRIZKLFKHJ&DQG3 PD\ EHVXEGRPLQDQWEDVHGRQUHFRYHU\IURPHQULFKPHQWV FRQWDLQLQJWRSynechococcus FHOOV VRPHPD\ HYHQEH]\PRJHQRXV HJ&
( &\DQREDFWHULDO0DW&RPPXQLW\ 6WUXFWXUH
(DUOLHU ZRUN RQ WKH DFWLYLWLHV DQG EHKDYLRU RI WKHUPRSKLOLF F\DQREDFWHULD ZDV UHYLHZHG E\ &DVWHQKRO] DQG %URFN +HUH ZH HPSKDVL]H PRUH UHFHQW ZRUN ZKLFK UHVXOWHG IURP PLFURSUREH DQG GHWDLOHG UDGLRODEHOOLQJ DQDO\VHV ([SHULPHQWVZHUHFRQGXFWHG RQQDWXUDOPDWHULDORU RQ FXOWLYDWHG F\DQREDFWHULD DQG WKH GDWD PXVW EH LQWHUSUHWHG ZLWK FDXWLRQ JLYHQ RXU LQFUHDVLQJ XQGHUVWDQGLQJ RI WKH SRSXODWLRQ ELRORJ\ RI PDW F\DQREDFWHULD
7KH JUHDWHU DEXQGDQFH RI OLSLGV FKDUDFWHULVWLF RI F\DQREDFWHULDDQGChloroflexus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³JXLOG´ LH F\DQREDFWHULD 2QH PXVWKRZHYHUEHFDXWLRXVLQLQWHUSUHWLQJPROHFXODU GDWD TXDQWLWDWLYHO\ EHFDXVH SULPHUGHSHQGHQW UHDFWLRQV OLNH F'1$ V\QWKHVLV DQG 3&5 PD\ EH VHOHFWLYH 5H\VHQEDFK HW DO 5DLQH\ HW DO DQG WKHUH LV QR JXDUDQWHH WKDW 3&5 LV TXDQWLWDWLYH 6X]XNL DQG *LRYDQQRQL )HUULV DQG:DUG 1HYHUWKHOHVVWKHREVHUYDWLRQVWKDW $ RU %OLNH F\DQREDFWHULDO 6 U51$ VHTXHQFHV SUHGRPLQDWHF'1$ DQG3&5EDVHGFORQHOLEUDULHV :DUGHWDO D DQG'**(SURILOHVFRPELQHG ZLWK WKH HQULFKPHQW DQG LVRODWLRQ RI % DQG %¶ Synechococcus FXOWXUHVIURPVDPSOHVGLOXWHGWRQHDU H[WLQFWLRQ LQRFXOD FRQWDLQHG DV IHZ DV
,9 3K\VLRORJLFDODQG %HKDYLRUDO (FRORJ\RI&\DQREDFWHULDRI *HRWKHUPDO +DELWDWV
$3KRWRV\QWKHVLV 5DWHV RI R[\JHQLF SKRWRV\QWKHVLV LQ
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV KRW VSULQJV SUREDEO\UHIOHFWV WKH FRRUGLQDWH DFWLYLWLHV RI D PL[WXUH RI FRH[LVWLQJ Synechococcus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ynechococcus SRSXODWLRQV LQ WKHVH PDWV 6XFK D WUHQG LV W\SLFDO RI R[\JHQLF SKRWRV\QWKHVLV LQ RWKHU WKHUPDO PDWV VXFK DV WKDW LQ +XQWHU V +RW 6SULQJV VHH EHORZ EXW LW ZDV DOWHUHG LQ 2FWRSXV 6SULQJ PDWV E\ UHPRYDO RI WKH XSSHU Synechococcus OD\HU DQG VXEVHTXHQW UHFRORQL]DWLRQ
Fig. 7. 0LFURHOHFWURGH PHDVXUHPHQWV RI R[\JHQLF SKRWRV\QWKHVLV LQ D ODPLQDWHG Synechococcus PDW IURP WKH VKRXOGHUUHJLRQRI 2FWRSXV 6SULQJ LQFXEDWHG DW GLIIHUHQW WHPSHUDWXUHV $ 'HSWKLQWHJUDWHGUDWHV % UDWHV LQ WKH VXUIDFH P P]RQH & UDWHV DW PP GHSWK >NLQGO\ SURYLGHG E\ 0 .XKO DQG )*DUFLD3LFKHO@
'0:DUGDQG 5: &DVWHQKRO]
)LJ 0LFURHOHFWURGH PHDVXUHPHQWV LOOXVWUDWLQJ YHUWLFDO GLVWULEXWLRQV RI $ R[\JHQLF SKRWRV\QWKHVLV % R[\JHQ DQG & S+ IRU ODPLQDWHG2FWRSXV6SULQJSynechococus PDWLQIXOO PLGGD\VXQOLJKW>IURP5HYVEHFKDQG:DUGD @
)HUULV HW DO $IWHU VXFK D GLVWXUEDQFH WKH SKRWLF ]RQH EHFDPH WKLFNHU DSSUR[LPDWHO\ PP FDXVLQJ JUHDWHU OLJKW SHQHWUDWLRQ DQG D ELPRGDO YHUWLFDO SURILOH RI R[\JHQLF SKRWRV\QWKHVLV ZDV REVHUYHG 7KHVH UHVXOWV PD\ VXJJHVW SRVVLEOH OLJKW DGDSWDWLRQ E\ GLIIHUHQW F\DQREDFWHULDO SRSXODWLRQV $OVR UHVXOWV IURP OLJKW UHGXFWLRQ H[SHULPHQWV DUH FRQVLVWHQW ZLWK HLWKHU DFFOLPDWLRQ 0DGLJDQ DQG %URFN DQGRU DGDSWDWLRQ 5XII5REHUWV HW DO RISynechococcus SRSXODWLRQVWRUHGXFHGOLJKW LQWHQVLW\ )XUWKHU HYLGHQFH RI GHSWK VWUDWLILFDWLRQ RI F\DQREDFWHULDO SRSXODWLRQV GHULYHV IURP REVHUYDWLRQV RI GLIIHUHQFHV LQ WHPSHUDWXUH RSWLPD IRU R[\JHQLF SKRWRV\QWKHVLVQHDUWKHVXUIDFHRUDWPPGHSWKLQ WKH 2FWRSXV 6SULQJ PDW )LJ E DQG F DQG IURP PROHFXODUDQDO\VLV)LJ ,W LV LQWHUHVWLQJ WR VSHFXODWH WKDW FRH[LVWLQJ F\DQREDFWHULDO SRSXODWLRQV DGDSWHG WR GLIIHUHQW WHPSHUDWXUH RU OLJKW UHJLPHV PD\ KHOS VWDELOL]H SKRWRV\QWKHVLV GXULQJ IOXFWXDWLQJ FRQGLWLRQV RI OLJKW
DQG WHPSHUDWXUH 7KH GLIIHUHQW VSHFLDOL]HG F\DQREDFWHULDO SRSXODWLRQV FDQ EH VDLG WR EHORQJ WR WKH VDPH ³JXLOG´ LH R[\JHQLF SKRWRV\QWKHVLV VR WKDW LQWUDJXLOG GLYHUVLW\ PLJKW VWDELOL]H JXLOG IXQFWLRQ ZKLFKLQWXUQ PD\EHQHILWRWKHUFRPPXQLW\ PHPEHUVZKLFKDUHGHSHQGHQWRQSKRWRV\QWKHVLV
% (IIHFWRI8OWUDYLROHW5DGLDWLRQ RQ 3KRWRV\QWKHVLV :% QP DQG :$ QP DUH GHWULPHQWDO WR SKRWRV\QWKHVLV HLWKHU E\ GLUHFW OHVLRQV RI PROHFXODU WDUJHWV RU LQGLUHFWO\ E\ SURGXFWLRQ RI UHDFWLYH R[\JHQ PHWDEROLWHV 3DUWLDO SURWHFWLRQ LV DIIRUGHG E\ 895VFUHHQLQJ FRPSRXQGV VXFK DV VF\WRQHPLQ DQG SUREDEO\ P\FRVSRULQHOLNH DPLQR DFLGVUHDFWLYHR[\JHQPHWDEROLWH³TXHQFKHUV´VXFKDV FDURWHQRLGV VXSHUR[LGH GLVPXWDVH FDWDODVH RU SHUR[LGDVHVDQGE\YDULRXVV\QWKHWLFDQG'1$UHSDLU SURFHVVHV VXFK DV H[FLVLRQ UHSDLU DQG
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV SKRWRUHDFWLYDWLRQ &KDSWHU ,Q KRW VSULQJV RQO\ Calothrix VSS DQG Pleurocapsa VS ZKLFK RFFXU DW WHPSHUDWXUHV EHORZ a & DUH NQRZQ WR SRVVHVV VF\WRQHPLQ &\DQREDFWHULD WKDW JURZ DW KLJKHU WHPSHUDWXUHV VXFK DV Synechococcus VSS DQG Mastigocladus FI laminosus, DSSHDU WR ODFN VFUHHQLQJ FRPSRXQGV DQG DUWLILFLDO UHPRYDO RI 895 E\ ILOWHUV JUHDWO\ HQKDQFHG SKRWRV\QWKHVLV 0LOOHU HW DO XQSXEOLVKHG GDWD 4XHQFKLQJ DQG SRVVLEO\ GDUN UHSDLU SURFHVVHV DUH SUREDEO\ QHFHVVDU\ IRU VXUYLYDO RI WKH KLJKWHPSHUDWXUH IRUPV XQGHU QRUPDO FRQGLWLRQV
& 3DUWLWLRQLQJRI3KRWRV\QWKDWHLQ 6\QHFKRFRFFXV &HOOV 0RVW RI WKH SKRWRDXWURSKLFDOO\ IL[HG FDUERQ UHWDLQHG E\
KRXUV LQ SXUH FXOWXUH DV UHSRUWHG E\ 3HDU\ DQG &DVWHQKRO] +RZHYHU WKH ILHOG H[SHULPHQWV PLJKW QRW KDYH SURYLGHG DFFXUDWH PHDVXUHV RI in situ JURZWK UDWHV *URZWK UDWHV HVWLPDWHG IURP UDWHV RI GHFUHDVH RI Synechococcus FHOOV DIWHU GDUNHQLQJ WKH PDWV %URFN DQG %URFN UHTXLUHG WKH assumption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ynechococcus VSS PD\ HQWHU LQWR DQ DFWLYH EXWQRQJURZLQJ RU YHU\ VORZ JURZLQJ SKDVH VLPLODU WR WKDW IRXQG LQ RWKHU KLJK GHQVLW\ EDFWHULDO SRSXODWLRQV HJ ELROXPLQHVFHQW EDFWHULD LQ OLJKW RUJDQV 5XE\ DQG $VDWR DQG 1IL[LQJ EDFWHULD LQ URRW QRGXOHV :HUQHU ,Q WKHVH FDVHV FRQWURO RI SK\VLRORJLFDO VWDWH DSSHDUV WR EH OLQNHG WR DXWRUHJXODWRU\ VXEVWDQFHV )XTXD HW DO 3HUKDSV WKH VDPH LV WUXH RI F\DQREDFWHULDO PDWV
Fig. 9. 3DUWLWLRQLQJ RI SKRWRDXWRWURSKLFDOO\IL[HG &2 LQWR PDFURPROHFXODU DQGORZ PROHFXODU ZHLJKW PHWDEROLWHV LQ ODPLQDWHG 2FWRSXV 6SULQJ Synechococcus PDW H[SRQHQWLDOO\ JURZLQJ 6\QHFKRFRFFXV VS &FHOOV DQG PDW GD\V IROORZLQJ UHPRYDO RI WKH F\DQREDFWHULDO VXUIDFH OD\HU >IURP GDWD RI 1ROG DQG :DUG DQG )HUULV HW DO @
' 3KRWRH[FUHWLRQ )HUPHQWDWLRQDQGWKH &URVV)HHGLQJ RI+HWHURWURSKV 3KRWRDXWRWURSKV PXVW SOD\ D PDMRU UROH LQ FURVV IHHGLQJ RI IL[HG FDUERQ WR KHWHURWURSKLF PDW SRSXODWLRQV &KDSWHU ,Q SXUHFXOWXUH6 FI lividus LVRODWHV IURP KRW VSULQJV LQ 2UHJRQ DQG
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
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV
Fig. 10. 0LFURHOHFWURGH PHDVXUHPHQWVRIYHUWLFDO GLVWULEXWLRQV RIR[\JHQ VXOILGHDQGS+LQ D WR& F\DQREDFWHULDO PDWLQ +XQWHU V +RW6SULQJVDWGLIIHUHQWWLPHVRIGD\DQGOLJKWLQWHQVLWLHV DVLQGLFDWHG >IURP5HYVEHFKDQG:DUG E@
RVFLOODWRULDQ F\DQREDFWHULDHJ&DVWHQKRO]HWDO *DUFLD3LFKHOHW DO 0RVW RIWKHVH F\DQREDFWHULDZLOOUHWXUQWRWKHVXUIDFHLIWKHPDWLV FRPSOHWHO\GDUNHQHG +RZHYHU LQ Oscillatoria FI terebriformis, WKHUHLVDVHFRQGUHWUHDWGRZQZDUGLQWR VXOILGHULFKXQGHUPDWVDIWHUGXVNZLWKDUHWXUQWRWKH VXUIDFHLQHDUO\PRUQLQJ5LFKDUGVRQDQG&DVWHQKRO] E-¡UJHQVHQHWDO 1HLWKHUWKHFXHVQRU WKHUHDVRQVIRUWKLV XQXVXDOQLJKWWLPHEHKDYLRU DUH IXOO\ XQGHUVWRRG 2WKHU VWXGLHV VKRZHG WKDW WKH GD\WLPHDYRLGDQFHRIKLJKVRODULUUDGLDQFHHVSHFLDOO\ WKH895FRPSRQHQWSUHYHQWHGWKHSKRWRLQKLELWLRQRU SKRWRGHVWUXFWLRQRIWKHVHPLJUDWLQJVSHFLHV*DUFLD 3LFKHO HW DO *DUFLD3LFKHODQG&DVWHQKRO] .UXVFKHO DQG &DVWHQKRO] XQSXEOLVKHGGDWD ,Q RQHVWXG\ LQYROYLQJ K\SHUVDOLQH PDW VSHFLHV RI Oscillatoria DQG Spirulina, WKHUHZDVHYLGHQFHWKDW QRWRQO\SKRWRLQKLELWRU\LQWHQVLWLHVZHUHDYRLGHGEXW WKDW H[SRVXUH WR RSWLPDO LQWHQVLWLHV RI SKRWRV\QWKHWLFDOO\DFWLYHOLJKWZDVDFKLHYHGGXULQJ PRVW RIWKH GD\ E\ GRZQZDUG PRYHPHQW *DUFLD 3LFKHOHWDO 9HUWLFDOO\PLJUDWLQJILODPHQWRXV F\DQREDFWHULD DOVR UHWDLQ D KLJK FRQWHQW RI OLJKW KDUYHVWLQJSLJPHQWVZKLFKXQGHUKLJKOLJKWFDQDOVR DFWDV OHWKDOSKRWRVHQVLWL]LQJSLJPHQWV +RZHYHU WKLV KLJK SLJPHQW FRQWHQW DOVR DOORZV PRWLOH
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
3KRUPLGLXP LQ 9HUWLFDO6WUXFWXUHV %RWK FRQLFDO &RQRSK\WRQOLNH DQG FROXPQDU YHUWLFDOVWUXFWXUHVFRPSRVHG ODUJHO\RIPhormidium
VSRUVSS RFFXULQVORZIORZLQJWKHUPDOVWUHDPVRU LQ KRW VSULQJSRROV 3ODWH F :DOWHU HW DO 0DQ\ DUHVLOLFLILHGRU FDOFLILHG DQG LW ZDV VXJJHVWHG WKDW WKH\ IRUP DV D UHVXOW RI XSZDUG SKRWRRULHQWHG PRYHPHQWV RI WKH SULQFLSDO ILODPHQWRXV VSHFLHV RI Phormidium SUHVHQW 7KHXSZDUG RULHQWDWLRQ XVXDOO\ EHJLQV RQ VPDOO QRGXOHVRUEOLVWHUVLQWKH PDW ZKLFK DUH RIWHQ IRUPHG E\ 2 EXEEOHV GXULQJGD\WLPH ,W ZDV DOVR VXJJHVWHG WKDW WKH FRQLFDO DQG FROXPQDU VWUXFWXUHV ZHUH KHOLRWURSLF SRLQWLQJ VRPHZKDW VRXWKZDUGLQWKHQRUWKHUQKHPLVSKHUH$ZUDPLNDQG 9DQ\R 7KH IRUPDWLRQ RI FROXPQDU DQG FRQLFDO F\DQREDFWHULDO VWUXFWXUHV DV ZHOO DV ODPLQDWLRQV LQ KRW VSULQJ PDWV ZHUH UHYLHZHG E\ %URFN
6\QHFKRFRFFXV 6RPHWKHUPRSKLOLFSynechococcus VWUDLQVDSSHDUWR H[KLELW DWZLWFKLQJMHUN\ RU SLYRWLQJJOLGLQJPRWLRQ &DVWHQKRO] D %URFN 8QLFHOOV RI F\DQREDFWHULDJOLGH DQG VKRZ SKRWRWDFWLFUHVSRQVHV HJ Synechococcus VSS 6WDQLHU HW DO EDHRF\WHV RI VRPH 3OHXURFDSVDOHV :DWHUEXU\ DQG 6WDQLHU 5HFHQWO\ LW ZDV GHPRQVWUDWHG TXDQWLWDWLYHO\ E\ XVLQJ WLPHODSVH GLJLWDO LPDJH DQDO\VLV WKDW VRPH 2FWRSXV 6SULQJ Synechococcus LVRODWHV ZHUH FDSDEOH RI VORZ PRYHPHQW FD WR PV DQG VKRZHG SRVLWLYHSKRWRWD[LV5DPVLQJ HW DO 7KH PHFKDQLVP RI PRYHPHQWLQ WKHVH QRQIODJHOODWHG XQLFHOOV LV XQNQRZQ WKRXJK LW UHTXLUHV FRQWDFW EHWZHHQ FHOOV DQG D VROLG VXUIDFH DQGRU RWKHU FHOOV 'LIIHUHQWSynechococcus VWUDLQV H[KLELWHG GLIIHUHQW UHVSRQVHV WR OLJKW RI GLIIHUHQW LQWHQVLW\ )RU LQVWDQFH VWUDLQ & ZDV SRVLWLYHO\ SKRWRWDFWLF DW OX[ ZKHUHDV VWUDLQ & ZDV SRVLWLYHO\ SKRWRWDFWLF DW OX[ EXW QHJDWLYHO\ SKRWRWDFWLF DWOX[ 1DWLYH PDW Synechococcus SRSXODWLRQV DOVR H[KLELWHG OLJKWGLUHFWHG PRWLOLW\ 7KH VSHHG RI PRYHPHQW ZDV VXIILFLHQW WR SHUPLW PLJUDWLRQWKURXJKWKHSKRWLF]RQHZLWKLQDIHZ KRXUV 6KDGLQJ H[SHULPHQWV OHG WR WKH REVHUYDWLRQ WKDW Synechococcus PDWVEHFDPHGDUN JUHHQ ZLWKLQ KRXUV RI D UHGXFWLRQ LQ OLJKW LQWHQVLW\0DGLJDQDQG %URFN %URFN 7KLVPD\ UHIOHFW XSZDUGYHUWLFDO PLJUDWLRQ RI FKORURSK\OOULFK ORZOLJKW DGDSWHG SRSXODWLRQVZKLFKQRUPDOO\UHVLGHGHHSHULQ PDWV
9&RQFOXVLRQ 7KH ZRUN GHVFULEHGKHUHVKRZV WKH YDOXH RI IRFXVLQJ LQYHVWLJDWLRQV RQ D OLPLWHG JURXS RI RUJDQLVPV WKDW RFFXU ZLWKLQ DOLPLWHGVHW RI KDELWDWV :H DOVRKRSH
'0:DUGDQG 5: &DVWHQKRO] WKDW LW LOOXVWUDWHV D SRLQW PDGH E\ WKH RUJDQLF JHRFKHPLVW *HRIIUH\ (JOLQWRQ LQ KLV DFFHSWDQFH VSHHFK XSRQ UHFHLSW RI WKH $OIUHG 7UHLEV $ZDUG ZKR IRXQG WKHUHODWLRQVKLS EHWZHHQ WKH ZRUN RI SDVW DQG SUHVHQWVWXGHQWVRIWKHVDPHV\VWHPV akin to that of the masons who built the great cathedrals of Europe... [which] took, in many instances, several hundred years to build. The successive master masons gave their whole lives to them, building on, adding to, modifying, and restoring the work of their predecessors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
$FNQRZOHGJPHQWV :H WKDQN 1LHOV 5DPVLQJ 0LFKDHO .XKO )HUUDQ *DUFLD3LFKHO DQG %HYHUO\ 3LHUVRQ IRU NLQGO\ VKDULQJ WKHLU XQSXEOLVKHG ILJXUHV DQG 0DU\ %DWHVRQ IRU KHOS ZLWK ILJXUHFRQVWUXFWLRQ :H WKDQNWKH 86 1DWLRQDO 3DUN6HUYLFHIRUWKHLUVXSSRUWRI DQG DVVLVWDQFH ZLWK RXU ZRUN 7KH SUHSDUDWLRQ RIWKLV FKDSWHU DQG PDQ\ RI LWV FRQWHQWV ZDV IXQGHG E\ JUDQWV IURP WKH 1DWLRQDO 6FLHQFH )RXQGDWLRQ %65 WR '0: DQG ,%1 DQG ,%1 WR 5:& DQG WKH 1DWLRQDO $HURQDXWLFV DQG 6SDFH $GPLQLVWUDWLRQ 1$*: WR'0:
5HIHUHQFHV $QGHUVRQ ./ 7D\QH 7$ DQG :DUG '0 )RUPDWLRQ DQG IDWHRIIHUPHQWDWLRQSURGXFWV LQ KRWVSULQJF\DQREDFWHULDO PDWV $SSO (QYLURQ 0LFURELRO $ZUDPLN 60 DQG 9DQ\R -3 +HOLRWURSLVP LQ PRGHP VWURPDWROLWHV6FLHQFH
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scillatoria terebriformis LQKRWVSULQJV -3K\FRO &DVWHQKRO] 5: 7KHUPRSKLOLF EOXHJUHHQ DOJDH DQG WKH WKHUPDOHQYLURQPHQW %DFWHULRO5HY &DVWHQKRO] 5: D (FRORJ\ RI EOXHJUHHQ DOJDH LQ KRW VSULQJV ,Q &DUU 1* DQG :KLWWRQ %$ HGV 7KH %LRORJ\ RI %OXHJUHHQ $OJDH SS %ODFNZHOO 6FLHQWLILF 3XEOLFDWLRQV2[IRUG &DVWHQKRO] 5: E 7KH SRVVLEOH XVH RI VXOILGH E\ WKH ILODPHQWRXV SKRWRWURSKLF EDFWHULD RI KRW VSULQJV /LPQRO 2FHDQRJU &DVWHQKRO]5: 7KHHIIHFWRIVXOILGHRQWKHEOXHJUHHQDOJDH RIKRWVSULQJV,1HZ=HDODQGDQG,FHODQG-3K\FRO &DVWHQKRO] 5: 7KH HIIHFW RI VXOILGH RQ WKH EOXHJUHHQ DOJDH RIKRW VSULQJV ,,
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ynechococcus SRSXODWLRQV LQ DVLQJOH KRW VSULQJ PLFURELDO PDW KDELWDW $SSO (QYLURQ 0LFURELRO )HUULV 0- DQG :DUG '0 6HDVRQDO GLVWULEXWLRQV RI 6 U51$GHILQHG SRSXODWLRQV LQ D KRW VSULQJ PLFURELDO PDW H[DPLQHG E\ GHQDWXULQJ JUDGLHQW JHO HOHFWURSKRUHVLV $SSO (QYLURQ0LFURELRO )XTXD& :LQDQV 6& DQG *UHHQEHUJ (3 4XRUXP VHQVLQJ LQ EDFWHULD WKH /X[5/X[, IDPLO\ RI FHOO GHQVLW\UHVSRQVLYH WUDQVFULSWLRQDOUHJXODWRUV -%DFWHULRO *DUFLD3LFKHO ) DQG &DVWHQKRO] 5: &RPSDUDWLYH DQR[\JHQLF SKRWRV\QWKHWLF FDSDFLW\ LQ strains RI D WKHUPRSKLOLFF\DQREDFWHULXP$UFK0LFURELRO *DUFLD3LFKHO)DQG &DVWHQKRO]5: 2QWKH VLJQLILFDQFHRI VRODUXOWUDYLROHWUDGLDWLRQ IRUWKHHFRORJ\RIPLFURELDO PDWV ,Q 6WDO /DQG&DXPHWWH3 HGV 0LFURELDO 0DWV SS 1$72 $6,6HULHV9RO*6SULQJHU9HUODJ%HUOLQ+HLGHOEHUJ *DUFLD3LFKHO ) 0HFKOLQJ 0 DQG &DVWHQKRO] 5: 'LHO PLJUDWLRQV RIPLFURRUJDQLVPVZLWKLQDEHQWKLF K\SHUVDOLQHPDW FRPPXQLW\ $SSO(QYLURQ0LFURELRO *LRYDQQRQL 6- 5HYVEHFK 13 :DUG '0 DQG &DVWHQKRO] 5: 2EOLJDWHO\ SKRWRWURSKLF Chloroflexus: SULPDU\ SURGXFWLRQ LQ DQDHURELF KRW VSULQJ PLFURELDO PDWV $UFK 0LFURELRO *LRYDQQRQL 6- 7XUQHU 6 2OVHQ *- /DQH '- DQG 3DFH 15 (YROXWLRQDU\ UHODWLRQVKLSV DPRQJ F\DQREDFWHULD DQG JUHHQFKORURSODVWV -%DFWHULRO +DGHU '3 (IIHFWV RI :% RQ PRWLOLW\ DQG SKRWRRULHQWDWLRQ LQ WKH F\DQREDFWHULXP Phormidium XQFLQDWXP $UFK 0LFURELRO -RUJHQVHQ %% DQG 1HOVRQ '& %DFWHULDO ]RQDWLRQ SKRWRV\QWKHVLV DQG VSHFWUDO OLJKW GLVWULEXWLRQ LQ KRW VSULQJ PLFURELDOPDWVRI,FHODQG 0LFURE(FRO -RUJHQVHQ %% &DVWHQKRO] 5: DQG 3LHUVRQ %. 7KH PLFURHQYLURQPHQW ZLWKLQ PRGHP PLFURELDO PDWV ,Q 6FKRSI -: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS &DPEULGJH 8QLYHUVLW\ 3UHVV&DPEULGJH .DOODV 7 DQG &DVWHQKRO] 5: D ,QWHUQDO S+ DQG $73$'3 SRROV LQ WKH F\DQREDFWHULXP Synechococcus VS GXULQJ H[SRVXUHWRJURZWKLQKLELWLQJORZS+-%DFWHULRO .DOODV 7 DQG &DVWHQKRO] 5: E 5DSLG WUDQVLHQW JURZWK DW ORZ S+ LQ WKH F\DQREDFWHULXP Synechococcus VS - %DFWHULRO
.RQRSND $ $FFXPXODWLRQ DQG XWLOL]DWLRQ RI SRO\VDFFKDULGH E\ KRWVSULQJ SKRWRWURSKV GXULQJ D OLJKWGDUN WUDQVLWLRQ )(060LFURE(FRO .RSF]\QVNL('%DWHVRQ00DQG:DUG'0 5HFRJQLWLRQ RIFKLPHULF VPDOOVXEXQLWULERVRPDO51$ VHTXHQFHV FRPSRVHG RI JHQHV IURP XQFXOWLYDWHG PLFURRUJDQLVPV $SSO (QYLURQ 0LFURELRO 0DGLJDQ07DQG%URFN 7' 4XDQWLWDWLYHHVWLPDWLRQ RIWKH EDFWHULRFKORURSK\OO c LQWKHSUHVHQFHRIFKORURSK\OOa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lectonema notatum &\DQRSK\WD - 3K\FRO 6KLHD - %UDVVHOO 6 DQG :DUG '0 0LGFKDLQ EUDQFKHG PRQR DQGGLPHWK\ODONDQHVLQKRWVSULQJF\DQREDFWHULDOPDWV D GLUHFW ELRJHQLF VRXUFH IRU EUDQFKHG DONDQHV LQ DQFLHQW VHGLPHQWV 2UJ*HRFKHP 6KLHD- %UDVVHOO6DQG:DUG'0 &RPSDUDWLYHDQDO\VLVRI IUHH OLSLGV LQ KRW VSULQJ F\DQREDFWHULDO DQG DQR[\JHQLF SKRWRV\QWKHWLFEDFWHULDOPDWV2UJ*HRFKHP 6WDQLHU5< .XQLVDZD5 0DQGHO 0 DQG &RKHQ%D]LUH* 3XULILFDWLRQ DQG SURSHUWLHV RI XQLFHOOXODU EOXHJUHHQ DOJDH 2UGHU&KURRFRFFDOHV %DFWHULRO5HY 6WHZDUW :'3 1LWURJHQ IL[DWLRQ E\ EOXHJUHHQ DOJDH LQ
&KDSWHU &\DQREDFWHULDLQ *HRWKHUPDO +DELWDWV PHPEHUV LQ KRW VSULQJ F\DQREDFWHULDO PDWV 6\PS 6RF *HQ 0LFURELRO :DUG '0 :HOOHU 5 6KLHD - &DVWHQKRO] 5: DQG &RKHQ < D +RW VSULQJ PLFURELDO PDWV DQR[\JHQLF DQG R[\JHQLF PDWV RI SRVVLEOH HYROXWLRQDU\ VLJQLILFDQFH ,Q &RKHQ < DQG 5RVHQEXUJ ( HGV 0LFURELDO 0DWV 3K\VLRORJLFDO (FRORJ\ RI %HQWKLF 0LFURELDO &RPPXQLWLHV SS $P62& 0LFURELRO :DVKLQJWRQ '& :DUG'0 %DWHVRQ00:HOOHU 5DQG 5XII5REHUWV$/D 5LERVRPDO 51$ DQDO\VLV RI PLFURRUJDQLVPV DV WKH\ RFFXU LQ QDWXUH $GY0LFURELDO(FRO :DUG '0 %DXOG - &DVWHQKRO] 5: DQG 3LHUVRQ %. E 0RGHP SKRWRWURSKLF PLFURELDO PDWV DQR[\JHQLF LQWHUPLWWHQWO\ R[\JHQLFDQR[\JHQLF WKHUPDO HXFDU\RWLF DQG WHUUHVWULDO ,Q 6FKRSI-: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS &DPEULGJH 8QLYHUVLW\3UHVV&DPEULGJH :DUG': )HPV 0- DQG%DWHVRQ00 LQ SUHVV 2UJDQL]DWLRQ RI QDWLYH SRSXODWLRQV ZLWKLQ KRW VSULQJ PLFURELDO PDW FRPPXQLWLHV QHHG IRU D PRUHHFRORJLFDO DSSURDFK ,Q ,60( 6\PSRVLXP3URFHHGLQJV6DQWRV%UD]LO :DUG'0)HUULV0-1ROG6&%DWHVRQ00.RSF]\QVNL('DQG 5XII5REHUWV $/ D 6SHFLHV GLYHUVLW\ LQ KRW VSULQJ PLFURELDO PDWV DV UHYHDOHG E\ ERWK PROHFXODU DQG HQULFKPHQW FXOWXUH DSSURDFKHVUHODWLRQVKLS EHWZHHQ ELRGLYHUVLW\ DQG FRPPXQLW\ VWUXFWXUH ,Q 6WDO /- DQG &DXPHWWH 3 HGV 0LFURELDO PDWV 6WUXFWXUH 'HYHORSPHQW DQG (QYLURQPHQWDO 6LJQLILFDQFH 6HULHV*(FRORJLFDO6FLHQFHV YRO1$72$6, 6HULHVSS²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¶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
59
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² =HQJ <% :DUG '0 %UDVVHOO 6 DQG (JOLQWRQ * D %LRJHRFKHPLVWU\ RI KRW VSULQJ HQYLURQPHQWV /LSLG FRPSRVLWLRQV RI
&KDSWHU &\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV /XFDV-6WDO 1HWKHUODQGV ,QVWLWXWH RI (FRORJ\ &HQWUH RI (VWXDULQH DQG &RDVWDO (FRORJ\ 32 %R[ 1/ $&
B A Whitton and M Potts (eds). The Ecology of Cyanobacteria. © 2000 Kluwer Academic Publishers Printed in the Netherlands
/- 6WDO
; 3KRVSKRUXV LQ 0LFURELDO 0DWV ;, &RQFOXVLRQV
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µPDWOLNH¶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
5RVHQEHUJ 6WDO DQG &DXPHWWH 'HWDLOHG LQIRUPDWLRQ RQ VWURPDWROLWHV FDQ EH IRXQG LQ ERRNV HGLWHG E\ :DOWHU 6FKRSI DQG 6FKRSI DQG .OHLQ $ JHQHUDO RYHUYLHZ RI EHQWKLF F\DQREDFWHULD LQ PDULQH OLWWRUDO HQYLURQPHQWV FDQ EH IRXQG LQ WKH ERRN FKDSWHU E\ :KLWWRQ DQG 3RWWV 7KLV UHYLHZ ZLOO GLVFXVV WKH PHWDEROLF DFWLYLWLHV RI F\DQREDFWHULD WKDW DOORZ WKHP WR IRUP PLFURELDO PDWVDQG VWURPDWROLWHV
,, 0LFURELDO 0DWV 6WURPDWROLWHV DQG WKHLU (QYLURQPHQWV $ :KDWDUH 0LFURELDO0DWVDQG 6WURPDWROLWHV " 6RPH 'HILQLWLRQV .UXPEHLQ UHIHUULQJWRWKHZRUNRI.DONRZVN\ SURSRVHG WKH IROORZLQJ GHILQLWLRQ RI D VWURPDWROLWH “Stromatolites are laminated rocks, the origin of which can clearly be related to the activity of microbial communities, which by their morphology, physiology, and arrangement in space and time interact with the physical and chemical environment to produce a laminated pattern which is retained in the final rock structure”. 7KLV GHILQLWLRQ LQFOXGHV IRVVLO DV ZHOO DV UHFHQW IRUPDWLRQV 0RGHUQ VWURPDWROLWHV WKDW ILW WKH GHILQLWLRQ RI .UXPEHLQ DUH UDUH $ZUDPLN DQG 0DUJXOLV LQ :DOWHU GHILQHG VWURPDWROLWHV DV “Organosedimentary structures produced by sediment trapping, binding and/or precipitation as a result of the growth and metabolic activity of microorganisms, principally cyanophytes”. 7KLV GHILQLWLRQ LQFOXGHV IRVVLO DQG UHFHQW FRQVROLGDWHG VWURPDWROLWHV DV ZHOO DV XQFRQVROLGDWHG PLFURELDO PDWV %RWK GHILQLWLRQV KRZHYHU HPSKDVL]H WKH UROH RI PLFURELDO PDWV DQG WKHDFWLYLW\RIWKHRUJDQLVPVWKHUHLQIRUWKHIRUPDWLRQ RI VWURPDWROLWHV :DOWHU IRUPXODWHG WKH IROORZLQJ FRQGLWLRQV QHFHVVDU\ WR IRUP D PLFURELDO PDW L WKH HQYLURQPHQWDO FRQGLWLRQV PXVW DOORZ JURZWK RIWKHPDWEXLOGLQJPLFURRUJDQLVPV LL WKH JURZWK UDWH RI WKH PDWEXLOGLQJ RUJDQLVPV PXVW EH IDVWHU WKDQ FRQVXPSWLRQ E\ JUD]LQJ RUJDQLVPV LLL VHGLPHQWDWLRQ UDWHV VKRXOG QRW EH H[FHHGLQJO\ KLJK WR DOORZ VWDELOL]HG FRORQL]DWLRQ RI WKH VXUIDFH E\WKHPDWEXLOGLQJRUJDQLVPV LY GHVWUXFWLYHIRUFHV VXFK DV EXUURZLQJRUJDQLVPV DQG PHFKDQLFDO DQG FKHPLFDO HURVLRQ PXVW EH DEVHQWRUDWOHDVWQRWSUHYHQWDFFUHWLRQRIRUJDQLVPV
,Q RUGHU WR SURGXFH D VWURPDWROLWH SUHVHUYDWLRQ RI WKHVWUXFWXUHPXVWRFFXU,QPDQ\HQYLURQPHQWVWRGD\ XQFRQVROLGDWHG PLFURELDO PDWV DUH IRUPHG LH V\VWHPV WKDWGR QRWKDYH WKHSRWHQWLDO WR SUHVHUYH LWV VWUXFWXUH GHILQHG E\ .UXPEHLQ DV “Unconsolidated laminated systems, clearly related to the activity of microbial communities, often called recent Stromatolites or living Stromatolites are defined as potential stromatolites”. ,QGHHG VWURPDWROLWHV sensu .UXPEHLQ DUH VWLOO IRUPHG WRGD\ ([FHOOHQW H[DPSOHV RI FRQVROLGDWHG ZHOOODPLQDWHG VWURPDWROLWHV WKDW VWLOO JURZ WKURXJK WKH JURZWK DQG PHWDEROLFDFWLYLW\RIDPLFURELDOPDWDUHIRXQGLQWKH ([XPD &D\V %DKDPDV 5HLG DQG %URZQH 3LQFNQH\HWDO 3ODWH 6LQFH WKHLU GLVFRYHU\ LQ LQ 6KDUN %D\ :HVWHUQ $XVWUDOLD SRRUO\ ODPLQDWHG FRQVROLGDWHG FDOFDUHRXV VWURPDWROLWHV KDYH EHHQ SUHVHQWHG DV VWULNLQJO\ VLPLODU WR 3UHFDPEULDQ VWURPDWROLWHV /RJDQ 5HFHQW FDOFDUHRXV 6WURPDWROLWHV DUH DOVRIRUPHGLQ3RO\QHVLDQDWROOV.RSDUD 'pIDUJHHW DO D E DQG RWKHU ODFXVWULQH DQG SHULPDULQH VHWWLQJV .HPSH HW DO .HPSH DQG .D]PLHUF]DN 6XFK VWURPDWROLWHV FDQ EH FDOOHG µUHFHQWVWURPDWROLWHV¶ WR GLVWLQJXLVKWKHPIURP IRVVLO IRUPDWLRQV ,W LV QRW QHFHVVDU\ WR QDPH XQFRQVROLGDWHG PLFURELDO PDWV µSRWHQWLDO VWURPDWROLWHV¶ VLQFHWKH\ DUH QRW VWURPDWROLWHV sensu .UXPEHLQ DQG PRVW RI WKHVH PLFURELDO PDWV ZLOO QHYHUEHFRPHRQH 7KHWHUP µSRWHQWLDO¶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in situ FDOFLILFDWLRQ 6XFK FORWWHG DQG SRRUO\ ODPLQDWHG VWURPDWROLWHV DUH DOVR NQRZQ DV WKURPEROLWHV .HQQDUG DQG-DPHV 7KH VWURPDWROLWHV RIWKH ([XPD&D\V%DKDPDVLQ FRQWUDVW ZLWK RWKHU UHFHQW IRUPDWLRQV SRVVHVV D ILQH ODPLQDWHG VWUXFWXUH ,Q DGGLWLRQWRWUDSSLQJDQGELQGLQJRIFDUERQDWHVDQG in
/- 6WDL situ SUHFLSLWDWLRQ RI FDOFLXP FDUERQDWH SURGXFHV GLVWLQFWOD\HUVRIFHPHQW5HLGDQG %URZQH
% 0LFURELDO0DWV DQG6WURPDWROLWHV WKH *HRORJLFDO(YLGHQFHVHHDOVR&KDSWHU 'XULQJ WKH +DGHDQ HUD IURP WKH RULJLQ RI WKH (DUWK [ \HDUVEHIRUHSUHVHQWWR [ WKHUHLVQR URFN UHFRUG 7KH ROGHVW URFN NQRZQ IURP WKH HDUO\ $UFKHDQ LV SUREDEO\ QRW RI ELRJHQLF RULJLQ 7KH ROGHVW VWURPDWROLWHV DUH NQRZQ IURP WKH $UFKHDQ DERXW [ \HDUV DJR 2QO\ D IHZ H[DPSOHV DUH NQRZQ IURP WKLV HUD EXW WKH\ ZHUH XQGRXEWHGO\ ELRORJLFDOO\ SURGXFHG 0LFURIRVVLOV KDYH EHHQ IRXQG LQ WKHVH VWURPDWROLWHV EXW LW LV SUHPDWXUH WR LGHQWLI\ WKHPDVF\DQREDFWHULD )URPPHDVXUHPHQWVRIFDUERQ LVRWRSH FRPSRVLWLRQV LQ WKHVH URFNV LW KDV EHHQ GHGXFHG WKDW DXWRWURSKLF PLFURRUJDQLVPV PXVW KDYH EHHQ DFWLYHDWWKDWWLPH +RZHYHU LWFRXOGKDYHEHHQ FKHPRDXWRWURSKV WKDW IL[HG WKH &2 UDWKHU WKDQ SKRWRDXWRWURSKV 7KH PRUSKRORJ\ RIWKH PLFURIRVVLOV IURPWKHVH ROGHVW VWURPDWROLWHV DOVR GRHV QRW JLYH DQ XQHTXLYRFDO FOXH DERXWWKH LGHQWLW\ RIWKHRUJDQLVPV &\DQREDFWHULD DUH D JURXS RI R[\JHQLF SKRWRWURSKLF RUJDQLVPV DQG LWLV ZHOO HVWDEOLVKHG WKDWWKH $UFKHDQ DWPRVSKHUHGLGQRWFRQWDLQR[\JHQ 'XULQJ WKH 3URWHUR]RLF ZKLFK VWDUWHG DERXW [ \HDUV DJR VWURPDWROLWHV EHFDPH DEXQGDQW )LJ DQG RFFXU LQ D ZLGH YDULHW\ RIIDFLHV 7KH\ RFFXSLHG HYHU\ PDMRU HFRORJLFDO QLFKH PDULQH DQG ODFXVWULQH VKDOORZ DQG GHHS ZDWHU 0RVW RI OLPHVWRQHVGRORPLWHVDQGPDJQHVLWHVDVZHOODVPDQ\ SKRVSKRULWHV DQG LURQ IRUPDWLRQV RI WKH 3URWHUR]RLF FRQWDLQ VWURPDWROLWHV /LNH PRGHUQ PLFURELDO PDWV LW VHHPV FHUWDLQ WKDWWKH 3URWHUR]RLF VWURPDWROLWHV ZHUH SURGXFHG E\ JURZWK DQG PHWDEROLF DFWLYLW\ RI F\DQREDFWHULD 7KH 3URWHUR]RLF VWURPDWROLWHV FRQWDLQ D ZHDOWK RI YHU\ ZHOO SUHVHUYHG PLFURIRVVLOV WKDW VKRZ DQ DVWRQLVKLQJO\ JRRG VLPLODULW\ WR SUHVHQW GD\ F\DQREDFWHULD 2YHU PLFURIRVVLOV KDYH EHHQ GHVFULEHGIURP VWURPDWROLWHIRUPDWLRQV:DOWHU HW DO 0DQ\ RIWKHVH IRVVLOV DUH SUHVHUYHG LQWKH FKHUW\ SDUWV RI VWURPDWROLWHV 7KH EHVW SUHVHUYDWLRQ RFFXUUHG IROORZLQJ HDUO\ VLOLFLILFDWLRQ RI WKH VWURPDWROLWHV %HFDXVH RI WKH DEVHQFH RI GLDWRPV GXULQJ WKLV HUDLW LV WKRXJKWWKDW VHDZDWHUPLJKWKDYH FRQWDLQHGPXFKKLJKHUFRQFHQWUDWLRQVRIVLOLFD 2[\JHQZDVSUHVHQWLQWKHDWPRVSKHUHDW[ \HDUVEHIRUHSUHVHQW ,WLV QRZ ZHOO DFFHSWHG WKDWWKH R[\JHQDWLRQ RI WKH DWPRVSKHUH ZDV WKH UHVXOW RI R[\JHQLF SKRWRV\QWKHVLV ,W PLJKW KDYH WDNHQ
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Fig. 1. 7KH UHODWLYH DEXQGDQFH RI VWURPDWROLWHV SORWWHG DJDLQVW WLPHDIWHU$ZUDPLN
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
7KH PRUSKRORJ\ RI VWURPDWROLWHV RI WKH [ ROG ,QVX]L JURXS RI 6RXWK $IULFD KLQW DW WKH LQYROYHPHQW RI ILODPHQWRXV RUJDQLVPV WKDW PLJKW KDYH KDG D SKRWRWDFWLF UHVSRQVH 0DVRQ DQG 9DQ %UXQQ +RZHYHU D FKHPRWDFWLF UHVSRQVH FRXOG DOVR H[SODLQ WKH VWUXFWXUH RI WKLV IRUPDWLRQ 6FKRSI DQG :DOWHU 2I WKH PDQ\ GLIIHUHQW PRUSKRORJLFDO IRUPV RI PLFURIRVVLOV PDQ\ FDQ EH WUDFHG EDFN IURP SUHVHQW GD\ F\DQREDFWHULD VXFK DV Oscillatoria DQG Lyngbya 6FKRSI DQG :DOWHU 7KHVH RUJDQLVPV DUH FRPPRQ LQ PRGHUQ PLFURELDO PDWV SDUWLFXODUO\ LQ V\VWHPV WKDWIL[QLWURJHQ ,W LV RI FRXUVH QRW SRVVLEOH WR GHWHUPLQH ZKHWKHU WKHVH DQFLHQW PDWV ZHUH GLD]RWURSKLF +RZHYHU PLFURIRVVLOV UHVHPEOLQJ F\DQREDFWHULD RI WKHKHWHURF\VWRXVJHQHUDNostoc DQG Scytonema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in situ SUHFLSLWDWLRQDVFHPHQW
LL in situ SUHFLSLWDWLRQ DV PLFULWH HLWKHU DFFUHWHG SDVVLYHO\ IURPVXVSHQVLRQRU WKURXJKWUDSSLQJDQG ELQGLQJ RI WKHJUDLQVE\ WKH PDWPLFURRUJDQLVPV LLL SUHFLSLWDWLRQ RI PLFULWH LPSRUWHG IURPDGMDFHQW HQYLURQPHQWV 7KH ILQH DQG GLVWLQFW ODPLQDWLRQ RI 3URWHUR]RLF VWURPDWROLWHV KLQWV DW in situ SUHFLSLWDWLRQ 0RVW 3KDQHUR]RLF VWURPDWROLWHV VKRZ SRRU RU QR ODPLQDWLRQ 7KHVHIRUPDWLRQVDUHSUREDEO\SURGXFHG E\ WUDSSLQJ DQG ELQGLQJ RI FDUERQDWH DQG VDQG JUDLQV &ORXG DQG 6HPLNKDWRY
& 6WUDWLILFDWLRQDQG6WUXFWXUHRI0LFURELDO 0DWVDQG6WURPDWROLWHV 0LFURELDO PDWV DUH FKDUDFWHUL]HG E\ WKH YHUWLFDO VWUDWLILFDWLRQ RI GLIIHUHQW IXQFWLRQDO JURXSV RI PLFURRUJDQLVPV 7KLV VWUXFWXUH LV WKH UHVXOW RI WKH SK\VLFRFKHPLFDO JUDGLHQWV WKDW DUH SUHVHQW LQ PDWV DQG LQ IDFWSURGXFHGE\ WKHPHWDEROLFDFWLYLW\RI WKH PDW RUJDQLVPV WKHPVHOYHV -¡UJHQVHQ HW DO 7KH W\SLFDO VWUXFWXUH RI D PLFURELDO PDW EXLOG E\ F\DQREDFWHULDLVVFKHPDWLFDOO\GHSLFWHGLQ )LJ
Fig. 2. 6FKHPDWLFGUDZLQJRIDEXLOGLQJRIDW\SLFDOPLFURELDOPDW IRUPHG E\ F\DQREDFWHULD 7KH OD\HU RI JUHHQ VXOIXU EDFWHULD KDV EHHQREVHUYHGRQRQO\DIHZRFFDVLRQV
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¡UJHQVHQ DQG 'HV 0DUDLV 7KLV ELRORJLFDO VWUDWLILFDWLRQ LV WKXV WKH UHVXOW RI JUDGLHQWV RI OLJKW R[\JHQ DQG VXOILGH DQG LV IRXQG LQ YLUWXDOO\ DOO F\DQREDFWHULDO PDWV )LJ ,Q VRPHUDUH FDVHV D OD\HU RI JUHHQ DQR[\JHQLF SKRWRWURSKLF EDFWHULD FDQ EHIRXQG XQGHUQHDWK WKHSXUSOHEDFWHULD 1LFKROVRQ HW DO $ GLVWLQFW OD\HU RI R[LGL]HG LURQ PD\ EH SUHVHQW EHWZHHQ WKH F\DQREDFWHULD DQG WKH SXUSOH VXOIXU EDFWHULD 6WDO ,W LV QRWFOHDU KRZ WKLV OD\HU LV IRUPHG ,W PD\ EH IRUPHG E\ FKHPLFDO R[LGDWLRQ E\
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Fig. 3. 9HUWLFDO SURILOHV RI R[\JHQ VXOILGH DQG S+ DW QLJKW OHIW SDQHO DQG GXULQJ WKH GD\ ULJKW SDQHO LQ D PDW RI Microcoleus chthonoplastes IURP6RODU/DNH6LQDL5HGUDZQIURP5HYVEHFKHWDO
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
Fig. 4. 6SHFWUDO OLJKW SHQHWUDWLRQ WKURXJK D PP PDW RI Microcoleus chthonoplastes RI WKH 1RUWK 6HD LVODQG RI 0HOOXP *HUPDQ\ )DUUHGOLJKWLVOHDVWDWWHQXDWHGE\WKHPDW7KHHIIHFWV RI OLJKW DEVRUSWLRQ E\ FKORURSK\OO a DQG SK\FRELOLSURWHLQV DW UHVSHFWLYHO\DQGQPDUHFOHDUO\SUHVHQW
Fig. 5. 7KH PDW RI Microcoleus chthonoplastes RI K\SHUVDOLQH SRQGRIWKHVDOWHUQRI([SRUWDGRUDGH6DO 6$*XHUUHUR1HJUR %DMD &DOLIRUQLD 6XU 0H[LFR VKRZLQJ WKH W\SLFDO PXOWLODPLQDWHG VWUXFWXUH
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
/- 6WDO DQR[\JHQLF SKRWRWURSKLF EDFWHULD DQG HYHQ SHUIRUP R[\JHQLF SKRWRV\QWKHVLV WKHUH 0LFURELDO PDWV PD\ EH DOVR MXVW µLQYHUWHG¶ ZLWK F\DQREDFWHULD RFFXUULQJ XQGHUQHDWK WKH OD\HU RI DQR[\JHQLF SKRWRWURSKLF EDFWHULD 9DQ *HPHUGHQ HW DO 7KLV W\SH RI PLFURELDO PDW FDQGHYHORS ZKHQ PXFK RUJDQLF PDWWHU LV GHSRVLWHG RQ WKH VHGLPHQW DQG LWV GHJUDGDWLRQ UHVXOWVLQYHU\ KLJK UDWHV RI VXOILGHSURGXFWLRQ 7KH MRLQW PHWDEROLF DFWLYLW\ RI PLFURRUJDQLVPV LQ PLFURELDO PDWV UHVXOWV LQ VWHHS SK\VLFRFKHPLFDO JUDGLHQWV HJ RI OLJKW R[\JHQ VXOILGH FDUERQ GLR[LGH DQG S+WKHVHJUDGLHQWV VKLIWPDUNHGO\ GXULQJ D K F\FOH )LJ -¡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µKLVWRULFDO¶ ODPLQDWLRQ 0RQW\ ,Q PDQ\ FRDVWDO HQYLURQPHQWVWKH RUJDQLF PDWWHU LV IXOO\ GHJUDGHG DQG D KLVWRULFDO ODPLQDWLRQ LV DEVHQW +RZHYHU LQ RWKHU PDWV SDUWLFXODUO\ WKRVH JURZLQJ LQ K\SHUVDOLQH HQYLURQPHQWV GHJUDGDWLRQ PD\ EH LQFRPSOHWH ([DPSOHV RI VXFK PDWV DUH WKH ZHOOLQYHVWLJDWHG K\SHUVDOLQHPDWV RIµSRQG¶ LQ *XHUUHUR1HJUR %DMD &DOLIRUQLD 0H[LFR DQG WKRVH RI 6RODU /DNH 6LQDL (J\SW '¶$QWRQL '¶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¡UJHQVHQ DQG &RKHQ .UXPEHLQ HW DO 7KH VRODU /DNH PLFURELDO PDW LV DERXW P WKLFN DQG WKH KLVWRULFDO ODPLQDWLRQ JRHV EDFN DOPRVW
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV ' (QYLURQPHQWVLQ ZKLFK &\DQREDFWHULDO 0DWV RFFXU &RDVWDO0LFURELDO0DWV 0LFURELDO PDWV LQFOXGLQJ WKRVH SURGXFHG E\ F\DQREDFWHULD XVXDOO\IRUPLQHQYLURQPHQWV LQ ZKLFK JUD]LQJ DFWLYLWLHV E\ PHWD]RD DUH ODUJHO\ H[FOXGHG ,W LV RIWHQWKHFDVH WKDW WKH PDWIRUPLQJ F\DQREDFWHULD LQ WKHVH HQYLURQPHQWV DUH QRW WKULYLQJ RSWLPDOO\ 7KH\ PD\ DOVR JURZ ZHOO XQGHU PRUH PRGHUDWH FRQGLWLRQV EXW LQ VXFK FDVHV JUD]LQJ ZLOO SUHYHQW WKH DFFXPXODWLRQ RI VXIILFLHQW ELRPDVV DQG WKH DFFUHWLRQ RI VHGLPHQW &RDVWDO WLGDO VDQG IODWV RIWHQ DUH H[FHOOHQW HQYLURQPHQWV IRU PLFURELDO PDWV WR GHYHORS SDUWLFXODUO\ ZKHQ WKH IODWV H[WHQG RYHU D ODUJH DUHD DQG ZKHQ WKH VORSH RI WKH IODW LV ORZ )LJ D /DUJH DUHDV ZLOO EH FRYHUHG E\ ZDWHU IRU RQO\DVKRUW SHULRG GXULQJ WKH WLGH DQG RIWHQ WKH VHGLPHQW LV QRW LQXQGDWHG IRU VHYHUDO GD\V GXULQJ QHDS WLGHV 6XFK VHGLPHQWV RIWHQ H[SHULHQFHODUJHIOXFWXDWLRQVLQ ZDWHU FRQWHQW VDOLQLW\ DQG WHPSHUDWXUHUHVXOWLQJLQ H[WUHPH FRQGLWLRQV WKDW OLPLW WKH UDQJH RI RUJDQLVPV DEOH WR LQKDELWWKLV HQYLURQPHQW 7KH QHDU DEVHQFH RI JUD]LQJ RUJDQLVPV DOORZV PDWEXLOGLQJ RUJDQLVPV VXFK DV F\DQREDFWHULD WR DFFXPXODWH &RDVWDO VDQG IODWV DUH XVXDOO\ QXWULHQWSRRU EXW WKH SKRWRWURSKLF F\DQR EDFWHULD KDYH ORZ QXWULHQWGHPDQGV 0RUHRYHU PRVW F\DQREDFWHULD UHVLVW ORQJ SHULRGV RI GURXJKW WROHUDWLQJ ODUJH IOXFWXDWLRQV RI VDOLQLW\ DQG WHPSHUDWXUH 2IWHQ WKHVH FRDVWDO PLFURELDO PDWV DUH FRPSRVHG RI ILODPHQWRXV F\DQREDFWHULD WKDW IRUP D GHQVH HQWDQJOHG PDVV ZKLFKWUDSV DQG ELQGV VHGLPHQW SDUWLFOHV 6XFK PDWV DUH FOHDUO\ YLVLEOH WR WKH QDNHG H\H DV PDVVLYH VWUXFWXUHV WKDWWR D ODUJH H[WHQW UHVLVW HURVLRQ )LJ E 7KHLUVHGLPHQWVWDELOL]LQJHIIHFWLV RI JUHDW LPSRUWDQFH IRU FRDVWDO PRUSKRJHQHVLV 7\SLFDO H[DPSOHV DUH IRXQG LQ WLGDO VDQG IODWV RI LVODQGV RI WKH VRXWKHUQ 1RUWK 6HD HJ 0HOOXP *HUPDQ\ DORQJ WKH HDVW FRDVW RI 1RUWK $PHULFD HJ *UHDW 6LSSHZLVVHWW 0DUVK &DSH &RG 0DVVDFKXVHWWV %LUG 6KRDO 1RUWK &DUROLQD 3DFLILF &RDVW HJ *XHUUHUR 1HJUR 0H[LFR 3ODWH DQG 6KDUN %D\DQG 6SHQFHU *XOI LQUHVSHFWLYHO\:HVWHUQ DQG 6RXWK $XVWUDOLD $ PRUH FRPSOHWH OLVW LV JLYHQ E\ 3LHUVRQ 0RVW VXFK FRDVWDO PLFURELDO PDWV DUH QRW VWURPDWROLWHV EXWH[DPSOHVRI VWURPDWROLWHV LQ FRDVWDO VHGLPHQWV FDQ EH IRXQG LQ (O +DPLUD %D\ 6LQDL ZKHUH VWURPDWROLWLF EHDFKURFN LV IRUPHG .UXPEHLQ D DQG DORQJ WKH ([XPD&D\V RI WKH %DKDPDV LQWHUWLGDO DQG VXEWLGDO VWURPDWROLWHV DUH
IRUPHG 5HLG DQG %URZQH 3ODWH Schizothrix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¶$QWRQL'¶$PHOLR HW DO 'HV0DUDLV HW DO 6DOWVDQG EULQHV DUH GLVFXVVHGPRUHIXOO\E\ 2UHQ LQ &KDSWHU
+RW6SULQJ0DWV RI&\DQREDFWHULD 7KHUPDO KRW VSULQJV DUH HQYLURQPHQWVLQ ZKLFK WKH FRPELQDWLRQ RIKLJK WHPSHUDWXUH LQ FRPELQDWLRQ ZLWK +6 RU DFLGLF FRQGLWLRQV GHFUHDVHV ELRGLYHUVLW\ HQRUPRXVO\ &\DQREDFWHULDO PDWV DUH PRVW FRPPRQ LQ KRW VSULQJVDW QHDU QHXWUDO RU DONDOLQHFRQGLWLRQV DQG DUHGHVFULEHGPRUHIXOO\LQ &KDSWHU 7KHUPDO VSULQJV WKDW FRQWDLQ VXOILGH PD\ OLPLW WKH IRUPDWLRQ PDWV RI VLQFH WKHUPRSKLOLF F\DQREDFWHULD GR QRW
/- 6WDO
D
E Fig. 6. a: 7KHH[WHQGHGWLGDOVDQGIODWRIWKHLVODQGRI0HOOXUQ6RXWKHUQ1RUWK6HD*HUPDQ\ DWORZWLGHFRYHUHGZLWKPLFURELDOPDWV b: 0DWXUHPDWV RIMicrocoleus chthonoplastes RIWKH LVODQG RI0HOOXPKDYH DFFUHWHG DQG IL[HG VR PXFK VHGLPHQW WKDW WKH\ DUH RIWHQ QRW VXEPHUJHGDWKLJKWLGH 7KLVGHFUHDVHVWKHJUD]LQJSUHVVXUH
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
WROHUDWHWKHFRPELQDWLRQRIKLJK WHPSHUDWXUH DQG KLJK OHYHOV RI VXOILGH &DVWHQKRO] $W PRGHUDWH FRQFHQWUDWLRQV RI VXOILGH PDWV RI Oscillatoria VSS KDYH EHHQ VKRZQ WR ORZHU WKH VXOILGH FRQFHQWUDWLRQ WKURXJK DQR[\JHQLF SKRWRV\QWKHVLV E\ WKH F\DQREDFWHULXP &RKHQ HW DO :DUG HW DO $QRWKHU VWUDWHJ\ LV IRXQG LQ WKH VRFDOOHG LQYHUWHG PDWV &DVWHQKRO] +HUH PDWV RI WKHDQR[\JHQLFSKRWRWURSK Chloroflexus RYHUOD\ WKH F\DQREDFWHULDO PDW &DVWHQKRO] $QR[\JHQLF SKRWRV\QWKHVLV VFDYHQJHV WKH VXOILGH DQG SURWHFWV WKH XQGHUO\LQJ PDW RI WKH R[\JHQLF KHWHURF\VWRXV F\DQREDFWHULXP Chlorogloeopsis VS -¡UJHQVHQ DQG 1HOVRQ
7HUUHVWULDO&\DQREDFWHULDO0DWV 7HUUHVWULDO F\DQREDFWHULDO PDWV FDQ EH IRXQG LQ D YDULHW\ RI GLIIHUHQW HQYLURQPHQWV 'H :LQGHU HW DO D E GHVFULEHG D F\DQREDFWHULDODOJDO FUXVW LQ FRDVWDO GXQHV 6DQG GXQHV KDYH D SRRU FDSDFLW\ IRU UHWDLQLQJ ZDWHU DQG DUH WKHUHIRUH H[WUHPHO\ GU\ HQYLURQPHQWV WKDW DUH FKDUDFWHUL]HG E\ D ORZ ELRGLYHUVLW\ 8QGHU FHUWDLQ FRQGLWLRQV D PDW RI Crinalium epipsammum, D XQLTXH EDQGVKDSHG ILODPHQWRXV F\DQREDFWHULXP GHYHORSV )LJ LWKDV DQ XQXVXDO FHOO HQYHORSH H[FHSWLRQDOO\ ZHOODGDSWHG WR GHVLFFDWLRQ 'H:LQGHUHW DO 7KLV VSHFLHV ZDV IRXQG WR EH LPSRUWDQW LQ WKH 1HWKHUODQGV LQ VWDELOL]LQJ DQG IL[LQJ WKH GXQH VDQG DQG SURWHFWLQJ LW IURP ZLQG HURVLRQ 2QFH WKLV RUJDQLVP KDV HVWDEOLVKHG D PDWUL[ WKH FRPPXQLW\ LV WDNHQ RYHU E\ WKH JUHHQ DOJD Klebsormidium flaccidum. 'HVLFFDWLRQ DQG OLIH XQGHU ORZ ZDWHU SRWHQWLDO DUH DOVR WKH FRQWUROOLQJ IDFWRUV IRU WKH GHYHORSPHQW RI F\DQREDFWHULDO PDWV DQG VWURPDWROLWHV LQ WKH KRW GHVHUW Microcoleus chthonoplastes, ZKLFK RFFXUV LQ VRPH GHVHUW FUXVWV %URFN KDV D SRO\VDFFKDULGH VKHDWK ZKLFK SOD\V DQ LPSRUWDQW UROH LQ SURWHFWLRQ IURP GHVLFFDWLRQ $IWHU UHZHWWLQJ WKH VKHDWK DEVRUEV ZDWHUDQGWKHF\DQREDFWHULXPUHVXPHV DFWLYLW\ LPPHGLDWHO\ &DPSEHOO 7KH VKHDWKV RI WKH XQLFHOOXODU GHVHUW Chroococcus VS DQG Chroococcidiopsis VS DOVR KDYH WKLV IXQFWLRQ DQG WKHVH VSHFLHV DUH IRXQGK\SROLWKLFDOO\ RQ URFNV LQ WKH 1HJHY GHVHUW3RWWVDQG )ULHGPDQQ 3RWWVHW DO &DLRODHW DO &\DQREDFWHULDOPDWV DUH SDUWLFXODUO\ ZHOO LQYHVWLJDWHG LQ WKH 1HJHY GHVHUW LQ,VUDHO )ULHGPDQQ HW DO %HUQHU DQG -HQVHQ .UXPEHLQ DQG *LHOH GHPRQVWUDWHG FDOFLILFDWLRQ LQ D PDW RI D XQLFHOOXODU F\DQREDFWHULXP SURGXFLQJ VWURPDWROLWLF VWUXFWXUHV LQ WKH GHVHUW
Fig. 7. $ VFDQQLQJ HOHFWURQ PLFURJUDSK RI Crinalium epipsammum, D QHZ VSHFLHV RI D F\DQREDFWHULXP WKDW IRUPV SKRWRWURSKLF PLFURELDO FUXVWV RQ FRDVWDO GXQHV 7KLV RUJDQLVP LV XQXVXDO EHFDXVH RI WKH IODW WULFKRPHV & epipsammum SRVVHVVHV DQXQXVXDOFHOOZDOOFRQWDLQLQJFHOOXORVH 7KHVFDOHEDULVP
&\DQREDFWHULDO PDWV DOVRVHHPWREH LQYROYHG LQ WKH IRUPDWLRQ RI URFN YDUQLVK LQ WKHGHVHUW 'HVHUWURFN YDUQLVK LV FRPSRVHG RI LURQ DQG PDQJDQHVH R[LGHV WKDW DUHSUHFLSLWDWHGE\ WKH PHWDEROLF DFWLYLW\ RI PDW PLFURRUJDQLVPV 7KH F\DQREDFWHULDO PDW LV XVXDOO\ SUHVHQW XQGHUQHDWK WKLV KDUG EURZQLVK OD\HU ZKHUH WKH\ DUH SURWHFWHG IURP GLUHFW VXQOLJKW DQG DUH FDSDEOH RI UHWHQWLRQ RI VRPH ZDWHU .UXPEHLQ DQG -HQV 7KH XQLFHOOXODU QLWURJHQIL[LQJ Gloeothece Gloeocapsa IRUPV PDWV RQ WKH ZDOOV RI FDUERQDWH FDYHV ZKHUH GD\OLJKW LV ORZ &R[ HW DO *ULIILWKV HW DO $QRWKHU H[DPSOH RI VXFK D ORZOLJKW WHUUHVWULDO HQYLURQPHQW LV WKH PDWV RI Leptolyngbya VS GHVFULEHG E\ $OEHUWDQR DQG .RYDFLN RQ WKH ZDOOV RI &DVD $XUHXP LQ 5RPH Nostoc RFFXUV DW WKHHQWUDQFHRI WKHFDUERQDWH FDYHV ZKHUH Gloeothece RFFXUV *ULIILWKV HW DO 7HUUHVWULDOPDWV RI Nostoc KDYH EHHQUHSRUWHG IURP DYDULHW\ RI GHVHUW HQYLURQPHQWV &KDSWHU LQFOXGLQJ WKHFROGGHVHUWLQ $QWDUFWLFD 'DYH\ 'DYH\ DQG 0DUFKDQW
,,, 7KH 2UJDQLVPV &\DQREDFWHULDWKDW EXLOG0LFURELDO 0DWV &\DQREDFWHULD WKDW EXLOG PLFURELDO PDWV EHORQJ WR D UDQJH RI VSHFLHV ILODPHQWRXV DV ZHOO DV XQLFHOOXODU 7KH ILODPHQWRXV QRQKHWHURF\VWRXV Microcoleus chthonoplastes LV WKH GRPLQDQW LQ PDULQH LQWHUWLGDO PLFURELDO PDWV DOO RYHU WKH ZRUOG 6WDO HW DO
/-6WDO
D b
F d
Fig. 8. 6FDQQLQJ(OHFWURQ0LFURVFRS\SKRWRJUDSKVRIDPDWRIMicrocoleus chthonoplastes RIWKHLVODQGRI0HOOXP*HUPDQ\ a: 2YHUYLHZRIWKHPDWVKRZLQJWKHODUJHSRO\VDFFKDULGHHQVKHDWKHGEXQGOHVRIWULFKRPHV6FDOHEDU P b: 'HWDLOVKRZLQJRQHHQGRID EXQGOHRIM. chthonoplastes. 7KHLQQHUURRPRIWKHEXQGOHLV FRPSRVHGRIGLIIHUHQWFRPSDUWPHQWV VHSDUDWHGE\SRO\VDFFKDULGHZDOOV 7KLV EXQGOHFRQWDLQV DODUJHQXPEHURIWULFKRPHV 6FDOHEDU P c: $VLGHYLHZRI DEXQGOHRIM. chthonoplusfes. 7KHRXWVLGHRI WKHEXQGOH LV FRORQL]HG ZLWKRWKHU PLFURRUJDQLVPV DPRQJ ZKLFK DUH GLDWRPV 6FDOHEDU P d $ GHWDLO RI WKH RSHQLQJ RI WKH EXQGOH ZLWK WKH LQGLYLGXDOWULFKRPHVVWLFNLQJRXW7KHWULFKRPHVFDQPRYHIUHHO\LQDQGRXWRIWKHEXQGOH6FDOHEDU P
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chthonoplastes, 6$* &ROOHFWLRQ RI $OJDH 8QLYHUVLW\ RI *RWWLQJHQ *HUPDQ\ DOVR NQRZQ DV VWUDLQ DQG LVRODWHG IURP D WLGDO PDW RI WKH 1RUWK
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV 6HD LVODQG 0HOOXP 6WDO DQG .UXPEHLQ DQG PI[ RULJLQDOO\ LVRODWHGIURPD PDW LQ$XVWUDOLD GLG QRW EHORQJ WR WKLV JURXS 7KH PRVW FRQVSLFXRXV FKDUDFWHULVWLF RI M. chthonoplastes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chthonoplastes, IRUPHG EXQGOHV EXWWKHVH ZHUH QRWVXUURXQGHG E\ DFRPPRQ VKHDWK 7KLV VWUDLQ ZDV UHDVVLJQHG DV Symploca VS 5 5LSSND SHUV FRPP D JHQXV WKDW VKRZV JUHDW VLPLODULW\ ZLWK Schizothrix, FRPPRQ LQ PLFURELDO PDWV $Q LQWHUHVWLQJ IHDWXUH RI M. chthonoplastes Symploca VWUDLQ 3HDUVRQ LV LWV FDSDFLW\ IRU QLWURJHQ IL[DWLRQ 3HDUVRQ HW DO 6WUDLQ6$*LV FDSDEOH RI LQGXFLQJ QLWURJHQDVH XQGHU DQR[LF FRQGLWLRQV 6WDO DQG .UXPEHLQ ,W KDV EHHQ QRWHG WKDW PDWV RI M. chthonoplastes IL[ QLWURJHQ3RWWVHW DO DQG LQ HDUO\ZRUNRQ PLFURELDO PDWV LW KDV EHHQ VXJJHVWHG WKDW WKH FRPPRQ VKHDWK RI WKLV QRQKHWHURF\VWRXV F\DQREDFWHULXP ZRXOG KHOS WR SURYLGH DQ DQR[LF IRU WKH R[\JHQVHQVLWLYH PLFURHQYLURQPHQW QLWURJHQDVH 3HDUVRQ HW DO VKRZHG WKDWEXQGOH IRUPDWLRQ LQ M. chthonoplastes ZDV QRW LQYROYHG LQ QLWURJHQ IL[DWLRQ 0RUHRYHU VWUDLQ 6$* ZDV LVRODWHG IURP D GLD]RWURSKLF PLFURELDO PDW EXW RQO\ FDSDEOH RI QLWURJHQ IL[DWLRQ XQGHU DQR[LF FRQGLWLRQV 1RQH RI WKH VWUDLQV LQYHVWLJDWHG E\ *DUFLD3LFKHO HW DO DQG DVVLJQHG E\ WKHP WR WKH WLJKWO\ GHOLPLWHG JURXS RI M. chthonoplastes ZDV FDSDEOH RI V\QWKHVL]LQJ QLWURJHQDVH XQGHU DHURELF RU DQDHURELF FRQGLWLRQV 9LOOEUDQGW DQG 6WDO XQSXEOLVKHG 6WUDLQ 3&& EHORQJV WR WKH M. chthonoplastes JURXS sensu *DUFLD3LFKHO HW DO DQG ZDV RULJLQDOO\ LVRODWHGIURPDVDOW PDUVKLQ:RRGV +ROH 86$ LW LV DOVR LQFDSDEOH RI LQGXFLQJ QLWURJHQDVH 5LSSND DQG :DWHUEXU\ 7KLV LV UHPDUNDEOH VLQFH RI WKH VWUDLQV RI WKH /33JURXS (Lyngbya, Plectonema, Phormidium), WR ZKLFK M. chthonoplastes EHORQJV KDYH WKH FDSDFLW\ RIV\QWKHVL]LQJ QLWURJHQDVH 7KLV LV DQRWKHU SURSHUW\ ZKLFK VHSDUDWHV VWUDLQ 6$* IURP WKH FRUH JURXS RI Microcoleus chthonoplastes. )XWXUH LQYHVWLJDWLRQV PD\ UHYHDO WKDW 6$* VKRXOG EH DVVLJQHG WR Symploca. Symploca DQG Schizothrix DUH ERWK FRVPRSROLWDQ PDWIRUPLQJ
F\DQREDFWHULD WKDW VKDUH VRPH PRUSKRORJLFDO FKDUDFWHULVWLFV ZLWK M. chthonoplastes. $ WUXH M. chthonoplastes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pirulina RU Merismopedia 3DOLQVND HW DO 2IWHQ WKH PDWV ZHUH FRPSRVHG RI PL[WXUHV RI GLIIHUHQW VSHFLHV )LJ $Q LPSRUWDQW VSHFLHV WKDW
Fig. 9. D 6FDQQLQJHOHFWURQ PLFURVFRSLF 6(0 SKRWRJUDSK RID QLWURJHQIL[LQJ PLFURELDO PDW RI Oscillatoria VSS DQG RWKHU F\DQREDFWHULD E 6(0 SKRWRJUDSK RI D GHWDLO RI D PDW RI Oscillatoria OLPRVD ZLWK WKH W\SLFDO FRLOHG ILODPHQW RI Spirulina subsalsa ZKLFK LV DW\SLFDO FRPSRQHQWRIWKHVH PDWV DQG D VLQJOH WULFKRPH RIMicrocoleus chthonoplastes.
ZDV DOZD\V SUHVHQW DQG ZDV IUHTXHQWO\GRPLQDQW ZDV Oscillatoria limosa 6WDO DQG.UXPEHLQ ZKLFK ZDV VKRZQ WR EH WKHGLD]RWURSKLFFRPSRQHQWRI WKHVH PDWV 7KLV VWUDLQ ZDV FDSDEOH RI DHURELF QLWURJHQ IL[DWLRQ LQ FXOWXUH 6WDO DQG .UXPEHLQ 7KH WD[RQRPLF SRVLWLRQ RI 0. limosa VWUDLQ LVRODWHG IURP 0HOOXP LV DOVR XQFHUWDLQ 7KH PRUSKRW\SH limosa LV REVHUYHG IUHTXHQWO\ LQ PLFURELDO PDWV DOO RYHU WKH ZRUOG DQG DV IDU DV NQRZQ DOO RI WKHVH PDWV DUH GLD]RWURSKLF +RZHYHU RIWHQ WKLV F\DQREDFWHULXP LV LGHQWLILHG DV Lyngbya aestuarii. L. aestuarii LV D ILODPHQWRXV F\DQREDFWHULXP ZLWK GLVNVKDSHG FHOOV DQG UHODWLYHO\ ZLGH WULFKRPHV P 7KH WULFKRPHV KDYH D WKLFNSRO\VDFFKDULGHVKHDWKZKLFK LV RIWHQ SLJPHQWHG 3ODWH 7KH RUJDQLVP LV QRW PRWLOH 5LSSND HW DO DVVLJQ WKLV RUJDQLVP WR WKH JURXS /33 $ ZKLFK GLIIHUV IURP Oscillatoria PDLQO\ E\ WKH DEVHQFH RI D VKHDWK DQG PRWLOLW\ LQ WKH ODWWHU 6WUDLQ SRVVHVVHV DWKLQVKHDWK DQG LV VOLJKWO\ PRWLOH 0DWV RI Lyngbya/Oscillatoria FDQ EH IRXQG LQ JHRJUDSKLFDOO\ GLVWDQWORFDWLRQV DQG DUH FKDUDFWHUL]HG E\ KLJK UDWHV RI QLWURJHQIL[DWLRQ)LJ $OWKRXJK PDQ\ PLFURELDO PDWV DUH HYLGHQWO\ JURZLQJ GLD]RWURSKLFDOO\ KHWHURF\VWRXV IRUPV DUH UDUHO\ WKH GRPLQDQW FRPSRQHQW +RZHYHU WKH IDFW WKH\ FDQ VRPHWLPHVEHLVRODWHGSURYHVWKHLUSUHVHQFH EXW DSSDUHQWO\ IDFWRUV DUH SUHVHQW WKDW SUHYHQW WKHLU SUROLIHUDWLRQ $ IHZH[FHSWLRQVDUHNQRZQ ,QSDUWV RI WKH WLGDO IODW LQ *XHUUHUR 1HJUR %DMD&DOLIRUQLD 6XU 0H[LFR H[WHQVLYH PDWV RI WKH KHWHURF\VWRXV Culothrix VS DUH SUHVHQW 6WDO 3ODWH ,Q D FRDVWDO ODJRRQ QHDU %RUGHDX[ )UDQFH PDWV RI Anabaena VRPHWLPHV GHYHORS 9LOOEUDQGW DQG 6WDO 0DQJURYHV RIWHQ VXSSRUW H[WHQVLYH PDWV RI WKH KHWHURF\VWRXV Scytonema VS 3RWWV
Fig. 10. 0L[HG FRPPXQLW\ RI GLIIHUHQW VSHFLHV RI Oscillatoria IURP D QLWURJHQIL[LQJ PLFURELDO PDW LQ D FRDVWDO ODJRRQ RQ WKH 0HGLWHUUDQHDQFRDVWRI)UDQFH
/- 6WDO Calothrix VS LVDOVRNQRZQ WR IRUP PDWV RQURFNVLQ WKH VSUD\ ]RQH DW WKH VHDVKRUH :KLWWRQ DQG 3RWWV 7KH GHYHORSPHQW RI WKHVH KHWHURF\VWRXV GLD]RWURSKLF PDWV LVGLVFXVVHGIXUWKHULQ 6HFWLRQ 9, +RWVSULQJPLFURELDOPDWV VXFK DV 2FWRSXV6SULQJ LQ WKH /RZHU *H\VHU%DVLQ RI
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV VDOLQLWLHV XS WR R $W KLJKHU VDOLQLW\ WKH XQLFHOOXODU Aphanothece halophytica XVXDOO\ GRPLQDWHV WKH PDWV 7KLV VSHFLHV RFFXUV XS WR D VDOLQLW\ RI 'RU DQG 3D] 7KH VDOLQLW\ WROHUDQFHV IRU F\DQREDFWHULD VHHP WR EH KLJKHU LQ PDWV RI WKH 6DENKD ZKHUHA. halophytica RFFXUV DW D VDOLQLW\ RI ZKLFK LV FORVH WR VDWXUDWLRQ ZKLOH S. subsalsa LV SUHVHQW DW D VDOLQLW\ RI 'RU DQG 3D] 7KH UHDVRQIRUWKLV GLIIHUHQFHLV XQFOHDU 6DOLQLW\ WROHUDQFH PD\ EH LQIOXHQFHG E\ WHPSHUDWXUH $OWKRXJK WKH VHGLPHQW VXUIDFH RI WKH 6DENKD PD\ EHFRPH KRW IURPWKH VRODUUDGLDWLRQ WKH VXEPHUJHG PDWV LQ 6RODUSRQGV PD\ DOVR EH H[SRVHG WR KLJK WHPSHUDWXUHV /LWKLILHG PLFURELDO PDWV IRXQG LQ WKH ([XPD&D\V %DKDPDV DUH EXLOW E\ WKH ILODPHQWRXV F\DQREDFWHULXP Schizothrix VS 3LQFNQH\ HW DO 3ODWH 7KLV F\DQREDFWHULXP IRUPV WKLQ WULFKRPHV RI DERXW P ZLGH ZLWK FHOOV WLPHV DV ORQJ DV ZLGH 7KH WULFKRPHV PD\ EH HQFORVHG E\ D WKLFN VKHDWK &RPPXQLWLHV RI Schizothrix VS PD\ IRUP GHQVH DQG WRXJK PDWV WKDW DUH RIWHQ DVVRFLDWHG ZLWK FDOFLXP FDUERQDWH SUHFLSLWDWLRQ 7KH OLWKLILHG PLFURELDOPDWV RI ([XPD&D\V %DKDPDV UHVXOW LQ WKH IRUPDWLRQ RI VWURPDWROLWHV D SURFHVV ZKLFK LV FRQWLQXRXV WRGD\ 5HLG DQG %URZQH 3LQFNQH\ HWDO
,9 0RWLOLW\ &KHPR DQG 3KRWRWD[LV of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
H[FHHG WKHDPRXQWWKDWFRXOG SRVVLEO\EHSURYLGHGE\ WKHRUJDQLVP+ROWRQDQG )UHHPDQ 0RUHRYHU WKH KLJKO\ PRWLOH Phormidium uncinatum GRHV QRW SRVVHVV SRUHV DOWKRXJK LW VHFUHWHV SRO\VDFFKDULGH +DGHU E 7KH VHFUHWLRQ RI PXFLODJH DV DPRGHO IRU WKH SURSXOVLRQ RI F\DQREDFWHULD DOVR GRHV QRW H[SODLQ WKH URWDWLRQ DORQJ WKH ORQJ D[LV LQ VRPH 2VFLOODWRULDFHDH +DOIHQ DQG &DVWHQKRO] DQG &DVWHQKRO] VXJJHVWHG WKDW FRQWUDFWLOH PLFURILEULOV WKDW WKH\ IRXQG LQ WKH H[WHUQDO OD\HUV RI WKH F\DQREDFWHULXP Oscillatoria princeps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hormidium DQG Anabaena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chthonoplastes LQGLFDWHG WKDW WKH VWUDWHJ\ RI WKLV RUJDQLVP LV WR PLQLPL]H PRYHPHQW ZKHQ FRQGLWLRQV DUH IDYRUDEOH ,QVWHDG RI YDU\LQJ WKH VSHHG RI PRYHPHQW SKRWRNLQHVLV LW PRYHV OHVV IUHTXHQWO\ M. chthonoplastes DOVR UHYHUVHV LWV PRYHPHQW IUHTXHQWO\ 7KLV LV QRW D SKRWRSKRELF UHVSRQVH EHFDXVH WKLV ZRXOG LPSO\ D VWHSGRZQ RU VWHSXS FKDQJH LQ OLJKWLQWHQVLW\ ZKLFK LV QRW WKH FDVH 5DPVLQJ DQG 3UXIHUW%HERXW IXUWKHU REVHUYHG WKDW M. chthonoplastes EHQGV PRUH IUHTXHQWO\DWRSWLPXPOLJKWLQWHQVLW\,QWKHORQJWHUP WKLV FRXOG OHDG WR FXUOLQJ RIWULFKRPHV LQWR EXQGOHV 0RWLOLW\ LQ VXFK EXQGOHV LV OLNHO\ WR EH UHVWULFWHG 6XFK F\DQREDFWHULD DUH OLNHO\ WR EH FRQILQHG WR D IL[HGSRVLWLRQLQWKHPDW)LJ *DUFLD3LFKHOHWDO GHPRQVWUDWHG WKDWPDW IRUPLQJF\DQREDFWHULD Oscillatoria VS DQG Spirulina subsalsa PLJUDWHG XS DQGGRZQ LQWKHPDWLQ DGDLO\ PDQQHU )LJ $W VXQVHW WKHVH F\DQREDFWHULD PRYHG WRZDUGV WKH PDW VXUIDFH DQG VWD\HG WKHUH WKURXJKRXW WKH QLJKW $W VXQULVH WKH\ PLJUDWHG GRZQZDUGV 7KH GHSWK WR ZKLFK WKH\ PLJUDWHG DSSHDUHG WR EHUHODWHG WR WKH OLJKW LQWHQVLW\ UHDFKLQJ WKH PD[LPXP GHSWK LQ WKH PDW DW PLGGD\ ZKHQ WKH OLJKW LQWHQVLW\ ZDV KLJKHVW ,QWHUHVWLQJ ZDV DOVR WKDW Oscillatoria VS DQG S. subsalsa FRQWDLQHG XQXVXDOO\ KLJK DPRXQWV RI FKORURSK\OO a GZW $ XQLFHOOXODU F\DQREDFWHULXP LQ WKLV PDW ZDV QRQ PRWLOH7KLVVSHFLHVFRQWDLQHGRQO\ FKORURSK\OO a *DUFLD3LFKHOHWDO ,WZDVFDOFXODWHGWKDWLI Oscillatoria DQG S. subsalsa GLG QRW PLJUDWH WKH\ ZRXOGEHSKRWRLQKLELWHGIRUPRVWRIWKHWLPHZKHUHDV WKH GDLO\ PRYHPHQW JXDUDQWHHG RSWLPXP SKRWR V\QWKHVLV WKURXJKRXW WKH OLJKW SHULRG 0DQ\ F\DQREDFWHULD PRYH GHHSHU LQWR WKH VHGLPHQW DWKLJK OLJKWLQWHQVLWLHV3HQWHFRVW :KDOHDQG:DOVE\ 5LFKDUGVRQDQG&DVWHQKRO]D )LJ
/-6WDO
Fig. 11. 0RYHPHQWV RI WKH OD\HU RI F\DQREDFWHULD LQ PLFURELDO PDWV GXULQJ D K SHULRG ,Q WKH XSSHUSDQHO DQ H[DPSOH RIWKH GDLO\ VLQXVRLGDO OLJKW FXUYH LV VKRZQ ([DPSOH , LV D PDW ZKLFK GRHVQRWGLVSODFHLWVHOIGXULQJDGD\QLJKWF\FOH 7KLV LVHLWKHUWKH FDVHZLWKXQLFHOOXODUF\DQREDFWHULDWKDWDUHQRWPRWLOHDQGJURZDW RSWLPDO OLJKW LQWHQVLW\ RU E\ VSHFLHV WKDW PLQLPL]H PRYHPHQW ZKHQ FRQGLWLRQV DUH IDYRUDEOH ZKLFK PD\ EH WKH FDVH LQ VRPH SRSXODWLRQV RI Microcoleus chthonoplastes. ([DPSOH ,,LV D PDW WKDWPRYHV WRZDUG WKHVXUIDFHDWVXQVHWDQG PRYHV GRZQLQWR WKH VHGLPHQWGXULQJWKHGD\8SZDUGVPRYHPHQWPD\EHFRQWUROOHGE\ FKHPLFDOIDFWRUVVXFKDVR[\JHQRUVXOILGH'RZQZDUGVPRYHPHQW LV LQ PRVW FDVHV DWWULEXWHG WR QHJDWLYH SKRWRWD[LV 0DWV RI Oscillatoria VSSRIWHQVKRZWKLVW\SHRIGLVSODFHPHQWGXULQJD KF\FOH ([DPSOH ,,, LV H[KLELWHG E\ WKH KRWVSULQJ Oscillatoria terebriformis. ,Q WKH GDUN WKH RUJDQLVP PRYHV UDQGRPO\ EXW PRWLOLW\ LV LQKLELWHG E\ VXOILGH ZKLFK HYHQWXDOO\ UHVXOWV LQ WKH DFFXPXODWLRQRIWKHSRSXODWLRQLQWKHVXOILGHULFKOD\HUGHHSLQWKH VHGLPHQW ,Q WKH OLJKW LW VKRZV D SRVLWLYH SKRWRWD[LV DW ORZ OLJKW DQG QHJDWLYHSKRWRWD[LV DWKLJK OLJKW 7KLV IRUFHV WKH RUJDQLVP WR PRYHGHHSHULQWRWKHVHGLPHQWGXULQJPLGGD\
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scillatoria VS ZDV VWULFWO\ GHSHQGHQW RQ OLJKW DQG JOLGLQJ VWRSSHG LQ WKH GDUN DIWHU D VKRUW SHULRG SUHVXPDEO\ EHFDXVH HQHUJ\ UHVHUYHV ZHUH H[KDXVWHG 7KHVH DXWKRUV GHPRQVWUDWHGUHVSRQVHV RIOscillatoria VS WRR[\JHQDHURWD[LV DQG&2 DQGELFDUERQDWH $
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icrocoleus chthonoplastes :KDOH DQG :DOVE\ Oscillatoria VS DQG S. subsalsa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scillatoria terebriformis. 7KLV F\DQREDFWHULXP RFFXUV LQ KRW VSULQJ PLFURELDO PDWV DQG KDV D OLJKWRULHQWHG PRWLOLW\ ,Q WKH GDUN WKH RUJDQLVP FRQWLQXHV WR PRYH EXW UDQGRPO\ ,W PD\ WKXV KDSSHQ WKDW LW PRYHV GRZQ LQWR WKH VHGLPHQW UHDFKLQJ WKH VXOILGH OD\HU 6XOILGH VHHPV WR LQKLELW PRWLOLW\ LQ O. terebriformis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terebriformis VKRZV
QHJDWLYH SKRWRWD[LV DQG PRYHV GHHSHU LQWR WKH VHGLPHQW LQ RUGHU WR SUHYHQW SKRWRR[LGDWLYH GDPDJH )LJ 7KH PDMRULW\ RI PRWLOH PDWIRUPLQJ F\DQREDFWHULD ZLOO SUHIHU ORZ OLJKW LQWHQVLWLHV DQG PRYH GHHSHU LQWR WKH VHGLPHQW GXULQJ WKH GD\ 7KH WUDSSLQJ RI O. terebriformis LQ WKH VXOILGH OD\HU GXULQJ WKH GDUN LV XQXVXDO EXW HVVHQWLDO IRU WKLV RUJDQLVP WR VXUYLYH WKH GDUN SHULRG ,Q WKH SUHVHQFH RI R[\JHQ GDUN UHVSLUDWLRQ ZLOO FDXVH D UDSLG GHSOHWLRQ RI WKH HQGRJHQRXV VWRUDJH FDUERK\GUDWH ZKLFK ZLOO UHVXOW LQ WKH GHDWK RI WKH RUJDQLVP LQ D PDWWHU RI KRXUV 7KH VXOILGHOD\HULV RI FRXUVH DQR[LF O. terebriformis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terebriformis LV FDSDEOH RI IHUPHQWLQJH[WUDFHOOXODU FRPSRXQGV VXFK DV IUXFWRVH DQG JOXFRVH 5LFKDUGVRQ DQG &DVWHQKRO] E &KHPRWDFWLF UHVSRQVHV RI F\DQREDFWHULD WR RUJDQLF FRPSRXQGV DUH ODUJHO\ XQNQRZQ )HFKQHU UHSRUWHG D QHJDWLYH FKHPRWDFWLFUHVSRQVHWRRUJDQLFDFLGVDQG5LFKDUGVRQ DQG &DVWHQKRO] REVHUYHG LQKLELWLRQ RI JOLGLQJ RI 0. terebriformis E\ IUXFWRVH 7KLV HIIHFW ZDV VLPLODU WR WKDWREVHUYHGIRUVXOILGH *OXFRVHWKHRWKHU VXEVWUDWHIRUWKLVRUJDQLVPGLGQRWKDYHDQHIIHFWQRU ODFWDWH ZKLFK LV RQH RI WKH IHUPHQWDWLRQ SURGXFWV SURGXFHG E\ O. terebriformis. ,Q FRQWUDVW WR ZKDW KDG EHHQ DVVXPHG LW ZDV GHPRQVWUDWHG E\ *DUFLD3LFKHO DQG %HERXW WKDW XOWUDYLROHW UDGLDWLRQ SHQHWUDWHV ZHOO LQ PLFURELDO PDWV 7KH DPRXQW RISHQHWUDWLRQ YDULHV ZLWK WKH W\SH RI VHGLPHQW RQ ZKLFK PLFURELDO PDWV GHYHORSHG 6LOW\ PXG DEVRUEHG UV OLJKW PRVW DQG TXDUW] VDQG WKH OHDVW 0DWV WKDW DUH PDLQO\ RUJDQLF WDNH DQ
/- 6WDO
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scillatoria VS DQG Spirulina subsalsa GHHS LQ WKH VHGLPHQW 7KLV LQWHQVLW\ LV RQO\ RI WKH OHYHO WKDW WKHVH RUJDQLVPV ZRXOG H[SHULHQFH DW PLGGD\ 7KHVH F\DQREDFWHULD UHVSRQGHG E\ QHJDWLYH SKRWRWD[LV ,Q DQRWKHU VWXG\ RI PLFURELDO PDWV LQ 6RODU /DNH 6LQDL LW ZDV VKRZQ WKDW M. chthonoplastes UHVSRQGV FOHDUO\ WR 89% OLJKW QP ([SRVXUH RI WKH PDW WR : P ZDV VXIILFLHQW WR FDXVH D GRZQZDUGV PLJUDWLRQ RI WKH F\DQREDFWHULD 7KH HIIHFW RI 89% ZDV DERXW WZR RUGHUV RI PDJQLWXGH VWURQJHU WKDQ QRUPDO YLVLEOH OLJKW $OVR 89$ KDG WKLV HIIHFW EXW ZDV DERXW WLPHV OHVV HIILFLHQW WKDQ 89% %HERXW DQG *DUFLD 3LFKHO ,W ZDV FRQFOXGHG IURP WKHVH H[SHULPHQWV WKDW M. chthonoplastes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
OD\HUV PRUH ELRPDVV DFFXPXODWHV JURVV SKRWR V\QWKHVLV LV HYHQ KLJKHU EXW GXH WR WKH ORZ OHYHO RI SKRWRV\QWKHWLF DFWLYH UDGLDWLRQ 3$5 ELRPDVV VSHFLILF SKRWRV\QWKHVLV LV ORZ 1RW DOO F\DQREDFWHULD H[KLELW QHJDWLYH SKRWRWD[LV ZLWK UHVSHFW WR 89 OLJKW 'RQNHU DQG +DGHU DQG 'RQNRU HW DO VKRZHG WKDW PRWLOLW\ LQ WKH F\DQREDFWHULD WKH\ LQYHVWLJDWHG ZDV UDWKHU LPSDLUHG E\ 89% QP 7KLV PD\ DOVR KDYH EHHQ WKH FDVH LQ D PDW RI M. chthonoplastes RI WKH WHPSHUDWH VRXWKHUQ 1RUWK 6HD ZKHUH SKRWRV\QWKHVLV ZDV VWURQJO\ LQKLELWHG E\ 89% UDGLDWLRQ DQG GLG QRW UHFRYHU GXULQJ WKH VXEVHTXHQW K ZKHQ 89 ZDV H[FOXGHG *DUFLD 3LFKHO DQG &DVWHQKRO] 7KH UHVSRQVHV RI F\DQREDFWHULD WR XOWUDYLROHW LUUDGLDWLRQ DUH GLVFXVVHG LQ PRUHGHWDLOLQ &KDSWHU
9 &DUERQ 0HWDEROLVP $ ,QWURGXFWLRQ &\DQREDFWHULD DUH WKH SULQFLSDO SULPDU\ SURGXFHUV LQ WKH PDMRULW\ RI PLFURELDO PDWV 2[\JHQLF SKRWRV\QWKHVLV DQG VRPHWLPHV DQR[\JHQLF SKRWRV\QWKHVLV GULYHV &2 IL[DWLRQ &\DQREDFWHULD HQULFK WKH PLFURELDO PDW ZLWK RUJDQLF PDWWHU &2 IL[DWLRQ SULPDULO\ UHVXOWV LQ WKH IRUPDWLRQ RI WKH VWUXFWXUDO ELRPDVV RIWKH F\DQREDFWHULD 7KLV RUJDQLF PDWWHU PD\ EHFRPH DYDLODEOH WR RWKHU RUJDQLVPV LQ WKH PDW E\ WKH GHDWK DQG VXEVHTXHQW O\VLV RI WKH F\DQREDFWHULD +RZHYHU LW DSSHDUV WKDW LQ VSLWH RI WKH KLJK UDWHV RI SKRWRV\QWKHVLV XVXDOO\ REVHUYHG LQ PLFURELDO PDWV QHW JURZWK RI WKH F\DQREDFWHULD LV RIWHQ ORZ LQ PDWXUH PDWV 1ROG DQG :DUG +HQFH RWKHU SURFHVVHV PXVW EH LQYROYHG LQ RUGHU WR GLYHUW SKRWRV\QWKDWH WR WKH PDW FRPPXQLW\ $V ZLOO EH GLVFXVVHG EHORZ WKHVH SURFHVVHV PD\ LQFOXGH WKH H[FUHWLRQ RI JO\FRODWH GXULQJ SKRWRUHVSLUDWLRQ WKH H[FUHWLRQ RI FRPSDWLEOHVROXWHVDIWHUDQRVPRWLFGRZQ VKRFN WKH H[FUHWLRQ RI IHUPHQWDWLRQ SURGXFWV GXULQJ GDUN DQR[LF FRQGLWLRQV DQG WKH VHFUHWLRQ RI H[WUDFHOOXODU SRO\PHULF VXEVWDQFHV (36
% 2[\JHQLF3KRWRV\QWKHVLV 2[\JHQLF SKRWRV\QWKHVLV UHTXLUHVWKHSUHVHQFHRI WZR SKRWRV\VWHPV 36 , DQG &\DQREDFWHULD FRQWDLQ FKORURSK\OO a LQ WKH UHDFWLRQ FHQWHUV RI ERWK 36 , DQG ,, EXWWKHIRUPHUFRQWDLQV DERXW WLPHV DV PDQ\ PROHFXOHV RI FKORURSK\OO a. 7KLV FKORURSK\OO PD\ DOVR FRQWULEXWH WR WKH OLJKW KDUYHVWLQJ EXW WKH SK\FRELOLSURWHLQVDUHIDUPRUHLPSRUWDQWSLJPHQWV DV
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¡UJHQVHQ DQG 'HV 0DUDLV3LHUVRQHW DO )LJ 6HGLPHQW DQG F\DQREDFWHULDO PDWV DUH UHODWLYHO\WUDQVSDUHQWWR OLJKW RI ZDYHOHQJWKV DERYH QP ZKLFK H[SODLQV WKH RFFXUUHQFH RI FRPPXQLWLHV RI EDFWHULRFKORURSK\OO aFRQWDLQLQJ DQR[\JHQLF SKRWRWURSKLF SXUSOHVXOIXU EDFWHULD 3LHUVRQ HW DO 7KHVH ILQGLQJV ZHUH FRQILUPHG E\ XVLQJ QHZO\ GHYHORSHG ILEHURSWLF PLFURSUREHV FRQQHFWHG WR D GLRGH DUUD\ GHWHFWRU .KO DQG -¡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able 1. &RPSDULVRQ RI SULPDU\ SURGXFWLYLW\ LQ PLFURELDO PDWV ZLWKRWKHUHFRV\VWHPV
(FRV\VWHP 0LFURELDO0DW 0HOOXP1RUWK6HD 6RODU/DNH6LQDL 6HDDQG2FHDQ 0HGLWHUUDQHDQ &RDVWDO8SZHOOLQJV 2FHDQ
3ULPDU\SURGXFWLYLW\ PJ&P G
/DNHV 2OLJRWURSKLFODNHV (XWURSKLF ODNHV +\SHUWURSKLFODNHV 0DQJURYHIRUHVWV 5DLQIRUHVW
DIWHU 6WDO
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
Fig. 12. A: 3KRWRV\QWKHVLVYHUVXVOLJKWFXUYHLQDPLFURELDOPDWRI WKH1RUWK 6HDLVODQG RI0HOOXP *HUPDQ\ GDWDIURP9LOOEUDQGW $ ODUJH QXPEHU RI GHSWK LQWHJUDWHG PHDVXUHPHQWV RI SKRWRV\QWKHVLV UHFRUGHG DW GLIIHUHQW GD\V DQG GXULQJ GLIIHUHQW WLPHV RIWKHGD\DWWKH VDPHORFDWLRQLQWKHPDWZHUH XVHG WRSORW WKLVFXUYH 7KH FXUYHZDV ILWWHGE\OLQHDUUHJUHVVLRQ 3PPRO 2 / PLG , 5 1 3 B: 7\SLFDOSKRWRV\QWKHVLV YHUVXV OLJKWLQWHQVLW\FXUYH RIDFXOWXUHRIDF\DQREDFWHULXP
/- 6WDO SKRWRV\QWKHVLV $W KLJKHU OLJKW LQWHQVLW\ F\DQR EDFWHULD LQ WKH GHHSHU SDUWV RI WKH PDW ZLOO H[KLELW KLJKHU UDWHV RI SKRWRV\QWKHVLV 7KH GLXUQDO YDULDWLRQ RI SKRWRV\QWKHVLV RQDEULJKWFORXGOHVVGD\LQ -XO\ LQ D PDWXUH PDW RI M. chthonoplastes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scillatoria limosa DV WKH GRPLQDQW VSHFLHV DQG ZLWK RQO\ RI WKH ELRPDVV FRPSDUHG WR WKH PDWXUH PDW FKORURSK\OO FRQWHQW W\SLFDOO\ JP 6WDO HW DO 9LOOEUDQGW )LJ D 3KRWRV\QWKHVLV RQ D VXUIDFHEDVLV ZDV PXFK ORZHU ZKLFK LV RI FRXUVH GXH WR WKH PXFK ORZHU ELRPDVV DQG WKHUHIRUH WKH VXSSO\ RI ',& PD\ KDYH EHHQ VXIILFLHQW WKURXJKRXW WKH GD\ 3KRWRV\QWKHVLVRQDELRPDVV EDVLV ZDV DERXW WZLFH DV KLJK DV LQ WKH PDWXUH PDW ZLWK FRPSDUDEOH VXUIDFH LQFLGHQWOLJKWLQWHQVLW\ 7KLV LV DWWULEXWHG WRWKHPXFK VPDOOHU DWWHQXDWLRQ RI OLJKW ZKHQ ELRPDVV LV ORZ LH WKH LQGLYLGXDO F\DQREDFWHULDUHFHLYH PXFK PRUH OLJKW LQ WKH IUHVKO\ FRORQL]HG VHGLPHQW FRPSDUHG WR WKH PDWXUH PDW 7KH KLJK UDWHV RI SKRWRV\QWKHVLV RIWHQ REVHUYHG LQ PLFURELDO PDWV PD\ FDXVH VXFK KLJK OHYHOV RI VXSHUVDWXUDWLRQ WKDW R[\JHQ EXEEOHV DUH IRUPHG 7KH GHQVH RUJDQLF PDWUL[ LQ ZKLFK WKH PDW F\DQREDFWHULD DUH HPEHGGHG UHSUHVHQW D GLIIXVLRQEDUULHU WKDW OLPLWV JDV H[FKDQJH WR D FRQVLGHUDEOH H[WHQG 2[\JHQ JDV EXEEOHV WKDW HYHQWXDOO\ GHYHORS PD\ DOVR EH WUDSSHG LQ WKLV PDWUL[ :KHQ VXFK PDWV DUH WKHQ LQXQGDWHG WKH\ PD\ EHFRPH EXR\DQW DQG OLIW RII IURP WKH VHGLPHQW (URVLRQ RI PLFURELDO PDWV DV D UHVXOW RI WKLV SKHQRPHQRQ FDQEHUHJXODUO\REVHUYHG 3LHFHV RI PDW PD\ EH FDUULHG WR YHJHWDWHG DUHDV DQG EHFRPH
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Fig. 13. 7KHGDLO\OLJKWFXUYH VKDGHG DUHD DQG GHSWK LQWHJUDWHG SKRWRV\QWKHVLV RI A D IUHVKO\ FRORQL]HG VHGLPHQW ZLWK Oscillatoria limosa DV WKH GRPLQDQW F\DQREDFWHULXP DQG B D PDWXUH PDW RI Microcoleus chthonoplastes. %RWK PDWV ZHUH ORFDWHG RQ WLGDO VDQG IODWV RQ WKH 1RUWK 6HD LVODQG RI 0HOOXP *HUPDQ\'DWDIURP9LOOEUDQGWHWDO
GHVLFFDWHG ZKHQ WKH WLGH KDV JRQH 6XFK GHVLFFDWHG SLHFHV PD\ EH WUDQVSRUWHG E\ ZLQG RYHU ORQJ GLVWDQFHV 7KH ILUVW WLPH WKLV SKHQRPHQRQ ZDV UHSRUWHG ZDV LQ DQG EHFDPH NQRZQ DV ³0HWHRUSDSLHU´ EHFDXVH LW ZDV EHOLHYHG WKDW LW FDPH IURP VSDFH ,Q KLV SXEOLFDWLRQ ³8EHU GDV LP -DKUH LQ&XUODQG YRP+LPPHO JHIDOOHQH0HWHRUSDSLHU XQG XEHU GHVVHQ =XVDPPHQVHW]XQJ DXV &RQIHUYHQ XQG ,QIXVRULHQ´ (KUHQEHUJ LGHQWLILHG WKLV PHWHRUSDSHUDVGHVLFFDWHGPLFURELDOPDWV
& $QR[\JHQLF3KRWRV\QWKHVLV $QR[\JHQLF SKRWRV\QWKHVLV LQ PLFURELDO PDWV LV QRW WKHH[FOXVLYHWUDLWRISXUSOH RUJUHHQ VXOIXUEDFWHULD 6RPH VSHFLHV RI F\DQREDFWHULD DUH FDSDEOH RI D W\SLFDO EDFWHULDOLNH DQR[\JHQLF SKRWRV\QWKHVLV LQ ZKLFK RQO\ SKRWRV\VWHP , LV LQYROYHG $V ZLWK SKRWRWURSKLF VXOIXU EDFWHULD DQR[\JHQLF SKRWRV\QWKHVLVLQF\DQREDFWHULDGHSHQGVRQVXOILGHDV
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° 6 H2O 7KHHOHPHQWDOVXOIXUWKDW LVSURGXFHGLVGHSRVLWHG RXWVLGH WKH FHOO EXW LV RIWHQ IRXQG DWWDFKHG WR WKH RXWHUVKHDWKRI WKHF\DQREDFWHULXPDV ILQHO\ GLVSHUVHG SDUWLFOHV 7KH HOHPHQWDOVXOIXULV QRWIXUWKHUR[LGL]HG DVLVWKH FDVH LQ IRULQVWDQFHSXUSOHVXOIXUEDFWHULD ,Q WKH PDWIRUPLQJ Microcoleus chthonoplastes
/-6WDO
WKLRVXOIDWH ZDV IRXQG WR EH WKH SURGXFW RI VXOILGH R[LGDWLRQ 'H:LWDQG9DQ*HPHUGHQ 6 H2S + 3 H2O + 6 CO2 C6H12O6 + 3 H2S2O3 :KLOHWKHR[LGDWLRQRI VXOILGH WR VXOIXU\LHOGVRQO\ WZR HOHFWURQVWKHR[LGDWLRQWR WKLRVXOIDWH \LHOGVIRXU HOHFWURQVSHUVXOILGHR[LGL]HG7KXVWKHR[LGDWLRQRI VXOILGH WR WKLRVXOIDWH LQ M. chthonoplastes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amphigranulata LVFDSDEOHRI XVLQJHOHPHQWDO VXOIXU IRU DVVLPLODWRU\ SXUSRVHV &DVWHQKRO] DQG 8WNLOHQ ,Q WKH XQLFHOOXODU Anacystis nidulans ORZ UDWHV RI &2 IL[DWLRQ DUH VXSSRUWHG E\ WKH R[LGDWLRQ RI WKLRVXOIDWH 8WNLOHQ3HVFKHN 0LFURFROHXV chthonoplastes EHORQJV WRWKHJURXS WKDW SHUIRUPR[\JHQLFDQG DQR[\JHQLF SKRWRV\QWKHVLV FRQFXUUHQWO\ ,Q WKLV RUJDQLVP WKH JURZWK UDWH GHFUHDVHG H[SRQHQWLDOO\ ZLWK LQFUHDVLQJ FRQ FHQWUDWLRQV RI VXOILGH $W D FRQFHQWUDWLRQ RI P0 VXOILGH S+ JURZWK ZDV FRPSOHWHO\ LQKLELWHG 2[\JHQLF SKRWRV\QWKHVLV ZDV JUDGXDOO\LQKLELWHGZLWK LQFUHDVLQJ FRQFHQWUDWLRQV RI VXOILGH 7KH UHODWLYH
FRQWULEXWLRQ RI DQR[\JHQLF SKRWRV\QWKHVLV WR WRWDO SKRWRV\QWKHVLV ZDV ! DW FRQFHQWUDWLRQV RI VXOILGH H[FHHGLQJ P0 S+ 7KH LQKLELWLRQ RI JURZWK DW P0 RI VXOILGH LV SUREDEO\ FDXVHG E\ WKH FRPSHWH LQKLELWLRQ RI R[\JHQLFSKRWRV\QWKHVLV UDWKHU WKDQ DQR[\JHQLFSKRWRV\QWKHVLV,WZDVVKRZQWKDW M. chthonoplastes UHTXLUHV R[\JHQ IRU JURZWK :KHQ R[\JHQLFSKRWRV\QWKHVLVZDV LQKLELWHG E\ '&08 M. chthonoplastes ZDV QRW FDSDEOH RI VXOILGHGHSHQGHQW DQR[\JHQLF SKRWRWURSKLF JURZWK XQOHVV VRPH R[\JHQ ZDV SUHVHQW 7KHUHIRUH D VPDOO FRQWULEXWLRQ RI R[\JHQLFSKRWRV\QWKHVLVPD\ EH QHFHVVDU\ LQ RUGHUWR SURYLGH WKH HVVHQWLDO R[\JHQ 2[\JHQ PD\ EH UHTXLUHG IRU WKH R[LGDWLRQ RI IDWW\ DFLGV Anacystis halophytica, IRU LQVWDQFH SRVVHVVHV DQ R[\JHQ GHSHQGHQW GHVDWXUDWLRQ PHFKDQLVP 3DGDQ DQG &RKHQ 6HYHUDO F\DQREDFWHULD FRQWDLQ SRO\XQVDWXUDWHG IDWW\ DFLGV .HQ\RQ HW DO DQG 3DGDQ DQG &RKHQ VXJJHVWHG WKDW VXFK F\DQREDFWHULD PD\ EH LQFDSDEOH RIDQDHURELF JURZWK O. limnetica GRHV QRW FRQWDLQ SRO\XQVDWXUDWHG IDWW\ DFLGV ZKLFK PD\ H[SODLQ LWV FDSDFLW\ IRU DQDHURELF JURZWK 3DGDQ DQG &RKHQ M. chthonoplastes 6$* FRQWDLQVFRQVLGHUDEOHDPRXQWVRI OLQROHDWH OLQROHQDWH DQG WHWUDGHFDGLHQDWH UHJDUGOHVV RI ZKHWKHU WKH FXOWXUH ZDV JURZQ LQ WKH SUHVHQFH RU DEVHQFH RI VXOILGH 'H:LW HW DO 7KH DIILQLW\ RI M. chthonoplastes IRU VXOILGH LV H[WUHPHO\ORZ 7KH.P IRUVXOILGHKDV EHHQFDOFXODWHG DV0 DSSUR[LPDWHO\WKHFRQFHQWUDWLRQ DW ZKLFK JURZWK RI M. chthonoplastes, FHDVHV 'H:LWDQG 9DQ *HPHUGHQ 7KHVH DXWKRUV DQDO\]HG WKH GDWD RI -IOUJHQVHQ HW DO ZKR LQYHVWLJDWHG WKH WUDQVLWLRQ RI DQR[\JHQLF SKRWRV\QWKHVLV WR R[\JHQLF SKRWRV\QWKHVLV LQ D PDW RI 0 chthonoplastes. $ .P RI 0 ZDV FDOFXODWHGIRUVXOILGHR[LGDWLRQLQ WKLV PDW 7KLV DIILQLW\ LVFORVHWKHRQHFDOFXODWHGLQ FXOWXUH E\ 'H :LW DQG YDQ*HPHUGHQ 7KHVHDIILQLWLHV IRU VXOILGH DUH H[WUHPHO\ ORZ ZKHQ FRPSDUHG WR WKH YDOXH RI 0 IRUDQ DQR[\JHQLF SKRWRWURSKLF SXUSOH VXOIXU EDFWHULXP DV Thiocapsa roseopersicina, WKDW LV IUHTXHQWO\SUHVHQWLQ PLFURELDO PDWV 'H:LWDQG 9DQ *HPHUGHQ 7KH VLPLODULW\ RI WKH .P RI VXOILGH R[LGDWLRQHVWLPDWHG LQ DFXOWXUH WR WKDW HVWLPDWHG LQ D QDWXUDO PLFURELDO PDW RI Microcoleus chthonoplustes LQGLFDWHV WKDW WKLV RUJDQLVP ZDV SUREDEO\ UHVSRQVLEOH IRU WKH VXOILGH R[LGDWLRQ LQ WKH PLFURELDO PDW 7KLV ZDV DOVR VXJJHVWHG E\ -¡UJHQVHQ HW DO DOWKRXJK WKH\ FRQFOXGHG WKLV IURPWKHIDFWWKDWSXUSOH VXOIXU EDFWHULD FRQVWLWXWHG RQO\ D PLQRU IUDFWLRQ LQ WKDWPDW 7KH UHVXOWV RQ DQR[\JHQLF SKRWRV\QWKHVLV LQ D FXOWXUH RI 0 chthonoplastes REWDLQHG E\ 'H :LW
&KDSWHU &\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV DQG YDQ *HPHUGHQ FRQILUPHG WKRVH IRU D QDWXUDO PDW E\ -¡UJHQVHQ HW DO -¡UJHQVHQ HW DO IRXQG WKDW R[\JHQLF DQG DQR[\JHQLF SKRWRV\QWKHVLV ZHUH FDUULHG RXW FRQFXUUHQWO\ +RZHYHU DW KLJKHU FRQFHQWUDWLRQV RI VXOILGH R[\JHQLF SKRWRV\QWKHVLV ZDV LQVLJQLILFDQW 2[\JHQLF SKRWRV\QWKHVLV LQ WKLV PDW VWDUWHG ZKHQ WKH VXOILGH FRQFHQWUDWLRQ GHFUHDVHG WR DERXW P0 ZKLFK LV LQ DJUHHPHQW ZLWK WKH UHVXOWV REWDLQHG E\ 'H :LW DQG YDQ *HPHUGHQ 7KH H[SHULPHQWV RI -¡UJHQVHQ DQGFRZRUNHUVVKRZHGWKDW R[\JHQLF SKRWRV\QWKHVLV FRXOG RFFXU HYHQ ZKHQ WKH PLFURELDO PDW ZDV H[SRVHG WR P0 RI VXOILGH LQ WKH RYHUO\LQJ ZDWHU $Q R[\JHQSHDN ZDV VDQGZLFKHG EHWZHHQ OD\HUV RI VXOILGH 7KLV VDQGZLFKLQJ RI F\DQREDFWHULD LQ PLFURELDO PDWV LV RIWHQ REVHUYHG $SSDUHQWO\ M. chthonoplastes LV FDSDEOH RI UHVLVWLQJ KLJK FRQFHQWUDWLRQV RIVXOILGH DQG UHFRYHUV R[\JHQLF SKRWRV\QWKHVLV ,W LV DOVR FDSDEOH RI R[LGL]LQJ WKLV KLJK FRQFHQWUDWLRQ RI VXOILGH LQ WKHOLJKWEXWWKLV GRHV QRW PHDQ WKDW WKH RUJDQLVPLV LQGHHG JURZLQJ RU HYHQ IL[LQJ &2 7DNLQJ WRJHWKHU WKH H[WUHPHO\ ORZ DIILQLW\ RI M. chthonoplastes IRU VXOILGH WKH ORZ JURZWK UDWH ZLWK DQR[\JHQLF SKRWRV\QWKHVLV DQG WKH IDFW WKDWWKLV RUJDQLVP FDQQRW JURZ LQWKH DEVHQFH RI R[\JHQ WKH PDMRU IXQFWLRQ VHHPV WR EH WKH GHWR[LILFDWLRQ RI VXOILGH
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
H[FKDQJHEHWZHHQWKH PHGLXP DQG WKH VLWH RI IL[DWLRQ LV UDSLG 'HV 0DUDLV DQG &DQILHOG LQYHVWLJDWHG WKH FDUERQ LVRWRSH GLVFULPLQDWLRQ LQ WZR PLFURELDO PDWV LQ *XHUUHUR 1HJUR %DMD &DOLIRUQLD 0H[LFR 7KH &PDW LQ WKHVH PDWV ZDV QRW YHU\ QHJDWLYH ,Q WKH PDW RI Lyngbya aestuarii WKLV YDOXH FRUUHVSRQGHG ZLWK WKHIUDFWLRQDWLRQIDFWRU 7KLV ORZ YDOXH ZDV HYLGHQWO\ DWWULEXWHG WR WKH FORVHG UHVHUYRLU EHKDYLRXU RI WKH V\VWHP 7KH ',& WKDW ZDV SURGXFHG E\ WKH PDW GXULQJ WKH QLJKW SRVVHVVHG WR VDPH QHJDWLYH YDOXH DQG WKHUHIRUH QR FKDQJHV LQ WKH &PDW ZHUH H[SHFWHG LQ WKH L. aestuarii PDW ,Q WKH PDW RI M. chthonoplastes SKRWRV\QWKHVLV GLG QRW GLVFULPLQDWH EHWZHHQ WKH OLJKWHU DQG KHDYLHU LVRWRSHV $W SUHVHQW D FRQFOXVLYH H[SODQDWLRQ IRU WKH QHJDWLYH YDOXH RI &PD[ LQ WKH M. chthonoplastes PDW LV QRW DYDLODEOH'HV0DUDLVDQG &DQILHOG 3URFHVVHV VXFK DVH[FUHWLRQIHUPHQWDWLRQDQG UHVSLUDWLRQ GR QRW FKDQJH LVRWRSLF GLVFULPLQDWLRQ 'LDJHQHVLV RI RUJDQLF PDWWHU GRHV QRW DOWHU LWV LVRWRSLF FRPSRVLWLRQ 'HV 0DUDLV HW DO DQG WKH &',& LV VLPLODU DV &PDW %DXHU HW DO ,WLV NQRZQ WKDW WKH µSRQG ¶ PDWV RI M. chthonoplastes DUH PRUH RU OHVV LQ µVWHDG\ VWDWH¶ LH PRVW RI WKH RUJDQLF PDWWHU WKDW LV SURGXFHG E\ SKRWRV\QWKHVLV LV PLQHUDOL]HG LQ WKH PDW ,W LV OLNHO\ WKDW SKRWRV\QWKHVLV VFDYHQJHV YHU\ HIILFLHQWO\ WKH ',& WKDW LV SURGXFHG LQ WKH PDW WKHUHE\ OLPLWLQJ QHW LVRWRSH IUDFWLRQDWLRQ ,Q IRVVLO 3URWHUR]RLF VWURPDWROLWHV & LV PXFK PRUH QHJDWLYH WKDQ LQ SUHVHQW GD\ PLFURELDO PDWV DQG VWURPDWROLWHV 7KLV PD\ UHIOHFW WKH KLJKHU OHYHOV RI ',& LQ WKH 3UHFDPEULDQ FRPSDUHG WR WRGD\¶V FRQFHQWUDWLRQV .HPSHDQG .D]PLHUF]DN D RUDPRUH QHJDWLYH &',&
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nabaena cylindrica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±2± ±'±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nabaena WKDW ZHUH VFUHHQHG DFFXPXODWHGVXFURVHLQUHVSRQVHWRRVPRWLFVWUHVV $ KDELWDWUHODWLRQVKLSLV VXJJHVWHGDPRQJVSHFLHVRIWKH XQLFHOOXODU Synechococcus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icrocoleus chthonoplastes DQG Oscillatoria limosa DFFXPXODWHG JOXFRV\OJO\FHURO DQG WUHKDORVH
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scillatoria VS FRQWDLQV WUHKDORVH ZKLOH WKH W\SLFDO RUJDQLVP LQ HVWDEOLVKHG PLFURELDOPDWV Microcoleus chthonoplastes, FRQWDLQV JOXFRV\OJO\FHURO )RU WKH VDPH UHDVRQ K\SHUVDOLQH VSHFLHV FRQWDLQ VXFURVH LQ DGGLWLRQ WR EHWDLQH 6LQFH EHWDLQH LV D QLWURJHQFRQWDLQLQJ FRPSRXQG QLWURJHQ GHILFLHQF\ PD\DOVROHDGWRWKHDFFXPXODWLRQRIQRQQLWURJHQRXV VXFURVHDVVHFRQGDU\RVPRO\WHV 7USHUDQG*DOLQVNL 2VPRWLFGRZQVKRFNH[HUWHGRQEHWDLQHFRQWDLQLQJ Aphanothece halophytica UHVXOWVLQWKHUHOHDVHRIWKLV RVPRO\WH LQWR WKH HQYLURQPHQW 5HHG DQG 6WHZDUW 7KLVPD\KDYHLPSRUWDQWFRQVHTXHQFHV IRU DQ HFRV\VWHP VXFK DV D PLFURELDO PDW EHFDXVH LW PD\ DOORZ FKHPRWURSKLF EDFWHULD WKDW FDQ QRW V\QWKHVL]H EHWDLQHWRWDNHLWXSIURPWKHHQYLURQPHQW5HHG DQG 6WHZDUW 0RUHRYHU EHWDLQH PD\ VHUYH DV VXEVWUDWHIRUVXOIDWHUHGXFLQJEDFWHULDDQGWKHSURGXFW RILWVPHWDEROLVPWULPHWK\ODPLQH70$ LVNQRZQDV D VRFDOOHG QRQFRPSHWLWLYH VXEVWUDWH IRU PHWKDQR JHQLFEDFWHULD+HLMWKXLVHQDQG+DQVHQ 2VPRWLFGRZQVKRFNLQRivularia atra UHVXOWHGLQ D FRUUHVSRQGLQJ GHFUHDVH RI WKH RVPRWLFXP WUHKDORVH EXWRQO\ ZDVUHFRYHUHGIURPWKHPHGLXPDQGWKH UHVW ZDV DSSDUHQWO\ PHWDEROL]HG RU FRQYHUWHG WR JO\FRJHQ 5HHG DQG 6WHZDUW 7KH JOXFRV\O JO\FHURO DFFXPXODWLQJ Synechocystis 3&& DQG WKH VXFURVHFRQWDLQLQJ Synechococcus 3&& UHOHDVHG RIWKHLUFDUERK\GUDWHV DQG RYHU RI WKHLU DPLQR DFLGV DIWHU K\SRRVPRWLF VKRFN 5HHG HW DO E +RZHYHU LQ VRPH RWKHU F\DQREDFWHULD
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icrocoleus chthonoplastes PD\ SURGXFH'063DV D VHFRQGDU\ RVPRO\WH DQGFRXOGEHWKHVRXUFHRI '06 LQ PLFURELDO PDWV +RZHYHU 9DQ %HUJHLMN DQG 6WDO IRXQG D FRUUHODWLRQ EHWZHHQ WKH QXPEHU RI GLDWRPV LQ WKHVH PDWV DQG WKH DPRXQW RI '06 WKDW HYROYHG IURP LW 6RPH EHQWKLF GLDWRPV DFFXPXODWH ODUJH DPRXQWV RI '063 DV RVPRWLFXP
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
/- 6WDO 6WDO DQG 0RH]HODDU UHYLHZHGIHUPHQWDWLRQLQ F\DQREDFWHULD 7DEOH OLVWV F\DQREDFWHULD FDSDEOH RI IHUPHQWDWLRQ 7KH SKHQRPHQRQ ZDV ILUVW GLVFRYHUHG LQ Oscillatoria limnetica 2UHQ DQG 6KLOR ZKLFK RFFXUV LQ WKH VXOILGHULFK K\SROLPQLRQ RI 6RODU /DNH 6LQDL DQG LV W\SLFDOO\ DGDSWHG WR DQDHURELF JURZWK ,Q WKH GDUN WKLV RUJDQLVP IHUPHQWV JO\FRJHQ WR ODFWDWH 6LQFH QR RWKHU IHUPHQWDWLRQ SURGXFWV ZHUH IRXQG LW ZDV DVVXPHG WKDW WKH KRPRODFWLF DFLG SDWKZD\ ZDV XVHG LQ WKLV RUJDQLVP ,Q WKH QRQ KHWHURF\VWRXV GLD]RWURSKLF PDWEXLOGLQJ Oscillatoria limosa, KHWHURODFWLF DFLG IHUPHQWDWLRQ ZDV IRXQG +H\HU HW DO 7KLV RUJDQLVP SURGXFHG HTXLPRODU DPRXQWV RI ODFWDWH DQG HWKDQRO IURP JO\FRJHQ ,Q DGGLWLRQ LW LV FDSDEOH RI KRPRDFHWLF IHUPHQWDWLRQ IRU ZKLFK LWV RVPRSURWHFWDQW WUHKDORVH ZDV XVHG DV VXEVWUDWH 7UHKDORVH ZDV GHJUDGHG WR DSSUR[LPDWHO\ DFHWDWH DQG VRPH K\GURJHQ DQG &2 7KH RFFXUUHQFH RI KRPRDFHWDWH IHUPHQWDWLRQ LQ F\DQREDFWHULD LV UHPDUNDEOH VLQFH LW IXUWKHU RQO\ RFFXUV LQ D JURXS RI VSHFLDOL]HG DQDHURELF EDFWHULD WKH DFHWRJHQLF EDFWHULD +RPRDFHWDWH IHUPHQWDWLRQ LV HQHUJHWLFDOO\ HIILFLHQW 0RUH UHFHQWO\ WKH RFFXUUHQFH KRPRDFHWDWH IHUPHQWDWLRQ KDV EHHQ SURSRVHG LQ D QXPEHU RI RWKHU F\DQREDFWHULD 'H 3KLOLSSLV HW DO 7KH GHJUDGDWLRQ RI WKH RVPRSURWHFWDQW LQ limosa ZDV DQRWKHU XQH[SHFWHG SKHQRPHQRQ 2I FRXUVHWUHKDORVHUHSUHVHQWV D ODUJHDPRXQWRI HQHUJ\ ZKLFK PD\ EHLPSRUWDQWIRUWKHRUJDQLVPWRXVHXQGHU D VLWXDWLRQ RIVHYHUH VWDUYDWLRQ 7KH TXHVWLRQ RI KRZ WKH RUJDQLVP FRPSHQVDWHV IRU WKH ORVV RI FRPSDWLEOH VROXWH KDV QRW EHHQ DQVZHUHG EXW LW KDV EHHQ VXJJHVWHG WKDW WKLV PD\ EH WKURXJK D WHPSRUDU\ DFFXPXODWLRQ RI LQRUJDQLF LRQV VXFK DV . 6WDO DQG 0RH]HODDU $OVR WKH PDW EXLOGLQJ F\DQREDFWHULXP Microcoleus chthonoplastes KDV EHHQ VKRZQ WR IHUPHQW SDUW RI LWV RVPRSURWHFWDQW 0RH]HODDU HW DO M. chthonoplastes DFFXPXODWHVWKHKHWHURVLGHJOXFRV\OJO\FHURO ZKLFK LV RQO\ GHJUDGHGLQFXOWXUHVWKDWFRQWDLQ ORZ DPRXQWV RI JO\FRJHQ 8QOLNH LQ O. limosa, M. chthonoplastes SRVVHVVHV RQO\ RQH IHUPHQWDWLRQ SDWKZD\ *O\FRJHQ DQG WKH JOXFRVH SDUW RI JOXFRV\O JO\FHURO DUH IHUPHQWHG YLD D PL[HG DFLG IHUPHQWDWLRQ UHVXOWLQJ LQ WKH IRUPDWLRQ RI IRUPDWH DFHWDWH HWKDQRO ODFWDWH + DQG VRPH&2 7KHUH LV D JUHDWGLYHUVLW\RI IHUPHQWDWLRQ SDWKZD\V LQ F\DQREDFWHULD ,Q VRPH FDVHV WKH SDWKZD\V KDYH EHHQ HOXFLGDWHG E\ WKH GHPRQVWUDWLRQ RI WKH HQ]\PHV LQYROYHG ,Q O. limosa WKH NH\ HQ]\PHV RI WKH KRPRDFHWDWH SDWKZD\KDYHEHHQ GHPRQVWUDWHG DV ZHOO +H\HU HW DO ,Q WKH PDMRULW\ RI SDWKZD\V WKH
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Table 2. &\DQREDFWHULDFDSDEOHRIIHUPHQWDWLRQ
25*$1,60
675$,1 25,*,1
)(50(17$7,21 352'8&76
6\PELRQW IURP Azolla caroliniana 6\PELRQW IURP Azolla caroliniana 6\PELRQW IURP Azolla filiculoides 3&&,QVW3DVWHXU
+RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH 0L[HGDFLG
Microcoleus chthonoplastes
0LFURELDOPDW
0L[HGDFLG
Microcystis aeruginosa Nostoc VS&F Nostoc VS $O Nostoc VS (IO Nostoc VS0$& Nostoc VS 0PO Nostoc VS0 Nostoc VS*P Nostoc VS 7 Nostoc VS %DOL Oscillatoria limnetica Oscillatoria limosa
3&&,QVW3DVWHXU 6\PELRQWIURP Cycas circinalis 6\PELRQW IURP Anthoceros laevis 6\PELRQW IURP Encephalartos ferox 6\PELRQW IURP Macrozamia lucida 6\PELRQW IURP Macrozamia moorei 6\PELRQWIURP Macrozamia VS 6\PELRQW IURP Gunnera manicata 3DGG\ILHOG 3DGG\ILHOG +\SROLPQLRQ 6RODU /DNH 0LFURELDOPDW
Oscillatoria sp.
0LFURELDOPDW
0L[HGDFLG +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRDFHWDWH +RPRODFWDWH +HWHURODFWDWH +RPRDFHWDWH QRWNQRZQ
Oscillatoria terebriformis Spirulina platensis
+RWVSULQJPLFURELDOPDW QRWNQRZQ
+RPRODFWDWH" 0L[HGDFLG
Spirulina minosa
QRWNQRZQ
QRWNQRZQ
Anabaena azollae $D/ Anabaena azollae $D1 Anabaena azollae $D6 Anabaena siamensis $VO Cyanothece
3$7+:$<
3DGG\ ILHOG
DFHWDWHODFWDWH&2 + DFHWDWHODFWDWH&2 + DFHWDWHODFWDWH&2+ DFHWDWH &2 + + HWKDQROODFWDWH IRUPDWHDFHWDWH +HWKDQROODFWDWH IRUPDWHDFHWDWH + HWKDQRO DFHWDWH DFHWDWHODFWDWH&2 + DFHWDWHODFWDWH&2 + DFHWDWHODFWDWH&2+ DFHWDWHODFWDWH &2 + DFHWDWH ODFWDWH &2 + DFHWDWH&2 + DFHWDWHODFWDWH DFHWDWHIRUPDWH&2+ DFHWDWH&2 + ODFWDWH ODFWDWHHWKDQRODFHWDWH ODFWDWHHWKDQRODFHWDWH IRUPDWH " +HWKDQRODFHWDWH IRUPDWHODFWDWH ODFWDWHDFHWDWH
FRPSRXQGVLQEUDFNHWVDUHSURGXFHGLQPLQRUTXDQWLWLHV)URP6WDODQG0RH]HODDU
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scillatoria limnetica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limosa LV FDSDEOH RI PDLQWDLQLQJ D FRQVLGHUDEOH UDWH RI QLWURJHQ IL[DWLRQ XQGHU DQDHURELF FRQGLWRQV LQ WKH GDUN 6WDO DQG +H\HU
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
Fig. 14. &DSVXODUDQGVOLPHH[WUDSRO\PHULFVXEVWDQFHV(36 DQG WKHIRUPDWLRQRIDPLFURELDOPDW $IWHU'HFKR
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
/- 6WDO
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miliania KX[OH\L ZRXOG SUHGRPLQDQWO\ VHUYHWKHSURGXFWLRQRI &2
2 HCO3
Ca2+
CaCO3
CO2 H2O
7KHVRFDOOHG µZKLWLQJV¶ FORXGVRI DUDJRQLWH QHHGOHV WKDWRIWHQRFFXU LQ WURSLFDOODJRRQVDQGWKDWKDYH EHHQ FRQVLGHUHG DV LQRUJDQLF SUHFLSLWDWHV PD\ EH SURGXFHG E\ WKH SKRWRV\QWKHWLF DFWLYLW\ RI GHQVH FRPPXQLWLHV RI SLFRSODQNWRQ 5REELQV DQG %ODFNZHOGHU &DOFLILFDWLRQ LQ PLFURELDO PDWV PD\ VHUYH DV D PHFKDQLVP RI SURGXFLQJ &2 'XH WR WKH GHQVH SKRWRWURSKLF ELRPDVV DQG KLJK UDWHV RI SKRWRV\QWKHVLV DQG WKH DONDOLQH FRQGLWLRQV LW LV OLNHO\
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
Ca2+ &2
CaCO3
,Q QRUPDO VHDZDWHU WKH FRQFHQWUDWLRQ RI FDOFLXP LRQ LV DERXW 0 (KUOLFK :KHQ WKH FRQFHQWUDWLRQ RI FDOFLXP LRQ LV DVVXPHG WR EH FRQVWDQW XQGHU FHUWDLQ FRQGLWLRQV WKHQ WKH FDUERQDWH FRQFHQWUDWLRQ GHWHUPLQHV FDOFLILFDWLRQ &2 UHDFWV ZLWK ZDWHU WR IRUP ELFDUERQDWH DQG WKLV GLVVRFLDWHV DFFRUGLQJWRWKHIROORZLQJUHYHUVLEOHUHDFWLRQV
&2 +2 HCOJ’
H+ HCOJ’ H+
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
&KDSWHU &\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV XVXDOO\ LQYROYHV &2 IL[DWLRQ LW KDV RIWHQ EHHQ FRQVLGHUHG LPSRUWDQW IRU FDOFLILFDWLRQ *ROXELF .UXPEHLQDQG*LHOH 3HQWHFRVW 7KH IL[DWLRQ RI &2 IURP D ELFDUERQDWHVROXWLRQZLOO UHVXOW LQ DQ LQFUHDVHLQFDUERQDWHLRQ
2 HCO3-
CO2 + H2O + CO32-
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³5HGILHOG´ VWRLFKLRPHWU\ RI &13 RI LV R[LGL]HG E\ 2 12 DQG 62 DFFRUGLQJ WR WKH IROORZLQJ UHDFWLRQV %RXGUHDXDQG&DQILHOG
CH2O NH 3 )16H3PO4 + 138O2 122 H2O + 106 CO2 + 16 NO3- + 16 H+ + H3PO4
CH2O NH3 HPO + 94.4NO384.8 H2O 13.6 CO2 92.4 HCO3 55.2 N2 H3PO4 CH2O NH3 16H3PO4 53SO42106 HCO3- + 53 H2S + 16 NH3 + H3PO4 7KHIRUPDWLRQRI&2 DQGWKH DFLGLILFDWLRQ RIWKH PHGLXP FRXOG UHVXOW LQ WKH GLVVROXWLRQ RI FDOFLXP FDUERQDWH UDWKHU WKDQ FDXVH LWV SUHFLSLWDWLRQ $QDHURELF UHVSLUDWLRQ UHVXOWV LQ WKH IRUPDWLRQ RI ELFDUERQDWHDQGFRXOGJLYHULVHWRVXSHUVDWXUDWLRQRI FDOFLXP FDUERQDWH 7KH SUHFLSLWDWLRQ RI VXOILGH DV LURQ VXOILGH DFWV DV D S+ EXIIHU 7KH HIIHFWV RIWKH VHTXHQWLDO R[LGDWLRQ RI RUJDQLF PDWWHU E\ WKH WKUHH HOHFWURQ DFFHSWRUV R[\JHQ QLWUDWH DQG VXOIDWH RQ SRUHZDWHU S+ DQG FDOFLXP FDUERQDWH VDWXUDWLRQ DUH FRPSOH[ DQG GHSHQG RQ WKH SUHYDLOLQJ FRQGLWLRQV %RXGUHDXDQG&DQILHOG ,Q WKH PLFURELDO PDWV RI6RODU /DNH 6LQDL LW KDV EHHQ VKRZQ WKDW VXOIDWH UHGXFWLRQ DQG &D&2 IRUPDWLRQZHUHVWRLFKLRPHWULFDOO\UHODWHG DQG RUJDQLF FDUERQ ZDV WUDQVIRUPHG LQWR D QXPEHU RI GLIIHUHQW FDUERQDWH PLQHUDOV -¡UJHQVHQ DQG &RKHQ .UXPEHLQ DQG &RKHQ .UXPEHLQ HW DO +RZHYHUZKHWKHUVXOIDWHUHGXFWLRQLQPLFURELDOPDWV LQ UHDOLW\ UHVXOWV LQ FDOFLXP FDUERQDWH SUHFLSLWDWLRQ GHSHQGVODUJHO\RQDYDULHW\RIFRQGLWLRQVWKDWSUHYDLO LQ WKHVH PLFURELDO PDWV 0RVW LPSRUWDQW DUH WKH GHYHORSPHQW RI DONDOLQH FRQGLWLRQV WKH UHPRYDO RI H[FHVV &2 RU WKH SUHVHQFH RI D VXLWDEOH EXIIHU (KUOLFK ,Q WKH DEVHQFH RI LURQ VXOIDWH UHGXFWLRQ SURGXFHV HTXDO DPRXQWV RI+¶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pIDUJH HW DO D E RU LQ WKH %DKDPDV 5HLG DQG %URZQH (YHQ LI WKH HDUO\ RFHDQV ZHUH PRUH DONDOLQH WKH PDULQH HQYLURQPHQW WRGD\ LV VWLOO VXSHUVDWXUDWHG ZLWK FDOFLXP FDUERQDWH )XUWKHUPRUH LQ PLFURELDO PDWV VHYHUDO ELRORJLFDO SURFHVVHV SUHGRPLQDWH ZKLFK SUHVXPDEO\ LQFUHDVH WKH FRQFHQWUDWLRQ RI FDUERQDWH LRQ ZKLFK WKHRUHWLFDOO\ VKRXOG OHDG WR FDOFLXP FDUERQDWH SUHFLSLWDWLRQ $V D UHVXOW RI DFWLYHSKRWRV\QWKHVLVDQG &2 IL[DWLRQ LQ WKH WRS OD\HU RI F\DQREDFWHULDO PDWV WKH S+ LQ WKHVH PDWV
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miliania huxleyi %RUPDQ HW DO ,Q PLFURELDO PDWV LW LV K\SRWKHVL]HG WKDW H[WUDFHOOXODU SRO\PHULF VXEVWDQFHV (36 ZKLFK DUH PDLQO\ FRPSRVHG RI SRO\VDFFKDULGHV VHUYH DV DJHQWV WKDW LQKLELW FDOFLILFDWLRQ (36 SURGXFHG E\ F\DQREDFWHULD DUH RIWHQ ULFK LQ XURQLF DFLGV DQG FRQWDLQ RWKHU DFLGLF JURXSV %HUWRFFKL HW DO 'HFKR 0DQ\ PLFURELDO PDWV DUH
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Fig. 15. 6LPSOLILHGPRGHO RIWKHSRVVLEOH LQWHUDFWLRQV RIFKDUJHG H[WUDFHOOXODU SRO\PHULF VXEVWDQFHV ZLWK FDOFLXP FDUERQDWH A: QXFOHDWLRQRIFDOFLXP DQGFDUERQDWH LRQV B: ,QKLELWLRQRI QXFOHDWLRQ RI SRO\DQLRQ C: ,QKLELWLRQ RI FU\VWDO JURZWK E\ DVVRFLDWLRQ RI D FU\VWDOOL]DWLRQ QXFOHXV ZLWK D SRO\DQLRQ D: &DOFLXPFDUERQDWH FU\VWDO ERXQG WR D OD\HU RIFKDUJHG SRO\PHUV 7KH JURZWK RIWKH FU\VWDO PD\ EH DUUHVWHG 7KH FKDUJHG SRO\PHU PD\GHWHUPLQHWKHHYHQWXDOVKDSHRIWKHFDOFLXPFDUERQDWHFU\VWDO $IWHU:HVWEURHNHWDO
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
0XFLODJH (36 LV RIWHQ SURGXFHG E\ F\DQREDFWHULD DV DQ RYHUIORZ PHWDEROLVP ZKHQ WKH\ JURZ XQGHU QXWULHQW OLPLWDWLRQ 7KLV LV SDUWLFXODUO\ WKH FDVH ZLWK QLWURJHQ DV WKH OLPLWLQJ IDFWRU D VLWXDWLRQ WKDW LV QRUPDOO\ RFFXUULQJ LQ WKH PDULQH HQYLURQPHQW ,Q PLFURELDO PDWV ZKHUH H[WUHPH GHQVH FRPPXQLWLHV RI F\DQREDFWHULD DUH SUHVHQW WKHUH LV D KLJK GHPDQG IRU QLWURJHQ ZKLOH WKHUH LV RIWHQ D VKRUWDJH RI WKLV LPSRUWDQW QXWULHQW 7KHUHIRUH PDQ\ PLFURELDO PDWV DUH GLD]RWURSKLF LH WKH F\DQREDFWHULD WKDW EXLOG WKHVH PDWV IL[ DWPRVSKHULF QLWURJHQ +RZHYHU PRVW GLD]RWURSKLF PDWV FRQVLVW RI QRQKHWHURF\VWRXV F\DQREDFWHULD $V DUJXHG LQ WKH VHFWLRQ RQ QLWURJHQ IL[DWLRQWKHVH F\DQREDFWHULD DUH QRWHIILFLHQW QLWURJHQ IL[HUV EHFDXVH WKH SURFHVV LV VHULRXVO\ KLQGHUHG E\ R[\JHQ LQ WKHVH RUJDQLVPV ,W LV OLNHO\ WKDW WKHVH F\DQREDFWHULD DUH LQ IDFW VWLOO QLWURJHQ OLPLWHG &\DQREDFWHULD WKDW JURZ XQGHU QLWURJHQ OLPLWHG FRQGLWLRQV WHQG WR SURGXFH D ORW RI PXFLODJH 2UWHJD &DOYR DQG 6WDO 3HUKDSV 3UHFDPEULDQ PLFURELDO PDWV DV ZHOO DV PRGHUQ FDOFLI\LQJ PDWV GR QRW JURZ QLWURJHQOLPLWHG DQG WKHUHIRUH SURGXFH OHVV PXFLODJH 6RPH RI WKHVH PDWV PD\ IRU RWKHU UHDVRQV UHFHLYH VXIILFLHQW FRPELQHG QLWURJHQ IRU JURZWK 2WKHU PLFURELDO PDWV DUH FRPSRVHG RI KHWHURF\VWRXV F\DQREDFWHULD ZKLFK DUH H[WUHPHO\ ZHOO DGDSWHG WR JURZ GLD]RWURSKLFDOO\ DQG DUH VHOGRP OLNHO\ WR H[SHULHQFH QLWURJHQ OLPLWDWLRQ XQGHU QRUPDO FRQGLWLRQV 2QH H[DPSOH IURP IUHVKZDWHU HQYLURQPHQWV LV WKH 5LYXODULDFHDH 7KLV JURXS RI KHWHURF\VWRXV F\DQREDFWHULD SURGXFHV H[WHQW FDOFLXP FDUERQDWH IRUPDWLRQV :KLWWRQ 7KH PDULQH Calothrix EHORQJV WR WKLV WD[RQRPLF JURXS DQG LQ VRPH H[FHSWLRQDOFDVHVDOVRIRUPVPLFURELDOPDWV $ PRGHO IRU FDOFLILFDWLRQ DQG WKH GHYHORSPHQW RI VWURPDWROLWHV LQ WKH ([XPD &D\V %DKDPDV LV SUHVHQWHG LQ )LJ 6XEWLGDO DQG LQWHUWLGDO VWURPDWROLWHV WKDW FDQ EH IRXQG LQ WKH ([XPD &D\V %DKDPDV DUH FKDUDFWHUL]HG E\ PDWV RI WKH F\DQREDFWHULXP Schizothrix VS 7KH PRGHO IRU FDOFLILFDWLRQ LQ WKHVH PDWV LV EDVHG RQ D QXPEHU RI REVHUYDWLRQV DQG DVVXPSWLRQV 7ZR W\SHV RI PDWV RI Schizothrix FDQ EH GLVWLQJXLVKHG /LWKLI\LQJ PLFURELDO PDWV RI Schizothrix DUH XVXDOO\ FKDUDFWHUL]HG E\ ORZ UDWLRV RI SKRWRV\QWKHVLV WR UHVSLUDWLRQ ZKLOH WKH RSSRVLWH LV WUXH LQ QRQOLWKLI\LQJ PDWV 3LQFNQH\ HW DO 0RUHRYHU FDOFLXP FDUERQDWH ZDV QRW DVVRFLDWHG ZLWK WKH F\DQREDFWHULD EXW ZLWK KHWHURWURSKLF EDFWHULD&KDIHW] DQG %XF]\QVNL &KDIHW] ,W ZDV IXUWKHU DVVXPHG WKDW H[WUDFHOOXODU SRO\PHULF VXEVWDQFHV (36 PD\ LQWHUIHUH E\ ELQGLQJ FDOFLXP DQG PDJQHVLXP LRQV
Fig. 16. 6LPSOLILHG VFKHPH RI WKH GHYHORSPHQW RI OLWKLILHG PLFULWLFOD\HUV LQ%DKDPDVVWURPDWROLWHV 6HHWH[W IRUH[SODQDWLRQ 7KLV K\SRWKHVLV ZDV GHYHORSHG E\ 3 5HLG - 3LQFNQH\ +: 3DHUO $: 'HFKR DQG /- 6WDO GXULQJ D UHVHDUFK FUXLVH LQ WKH %DKDPDVLQ
WKXV ORFDOO\ LQKLELWLQJ FDUERQDWH SUHFLSLWDWLRQ %RUPDQHWDO :HVWEURHNHWDO ([XPD &D\V VWURPDWROLWHV DUH IRUPHG DW KLJK HQHUJ\ VLWHV ,QWHUWLGDO VWURPDWROLWHV FDQ EH IRXQG RQ WKH $WODQWLF 2FHDQ FRDVW DQG DUH H[SRVHG WR KLJK ZDYH HQHUJ\ 6XEWLGDO VWURPDWROLWHV DUH DOPRVW H[FOXVLYHO\ HQFRXQWHUHG LQ FKDQQHOV ZLWK KLJK FXUUHQWV 7KH F\DQREDFWHULXP Schizothrix VS LV D ILODPHQWRXV RUJDQLVP FRPSRVHG RI WKLQ RIWHQ OHVV
/- 6WDO WKDQ PZLGH WULFKRPHVZKLFKDUHHQYHORSHGE\D WKLQ ULJLG SRO\VDFFKDULGH VKHDWK 7KLV RUJDQLVP LV FDSDEOH RI FRORQL]LQJ D VROLG VXEVWUDWH %HFDXVH JUD]LQJ SUHVVXUH ZLOO EH ORZ LQ WKHVH KLJKHQHUJ\ DUHDV D FRPPXQLW\ RI Schizothrix PD\ GHYHORS 8QGHU FRQGLWLRQV RIORZ VHGLPHQWDWLRQ UDWH D PDW RI Schizorhrix DQG DVVRFLDWHG PLFURRUJDQLVPV ZLOO GHYHORS )LJ D 7KHVH PDWV DUH ULJLG DQG WLJKWO\ DVVRFLDWHG ZLWK WKH XQGHUO\LQJ VXEVWUDWH 7KH F\DQREDFWHULD ZLOO JURZ DQG SURGXFH VKHDWK PDWHULDO DQGSRVVLEO\VRPHPXFXV,WLV DVVXPHGWKDWWKLV(36 ZLOO ELQG &D RU WKDW XURQLF DFLGV SUHYHQW IXUWKHU JURZWKRIFU\VWDOOL]DWLRQQXFOHL%RUPDQHWDO 'XULQJ D SHULRG RI VHGLPHQWDWLRQ Schizorhrix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chizothrix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
&KDSWHU &\DQREDFWHULDO 0DWVDQG6WURPDWROLWHV
DQG RWKHU RUJDQLVPV DQG LV FDSDEOH RILPPRELOL]LQJ FDOFLXP9HUUHFFKLDHWDO 7KLV PRGHO H[SODLQV WKDWin situ FDOFLXPFDUERQDWH SUHFLSLWDWLRQ DQG OLWKLILFDWLRQ RIWKH PDWLV FRQWUROOHG E\ ELRORJ\ ,W LV LQGLUHFWO\ DVVRFLDWHG ZLWK WKH F\DQREDFWHULD EXW GHJUDGDWLRQ RI RUJDQLF PDWWHU QRWDEO\(36 E\KHWHURWURSKLFEDFWHULDLVUHTXLUHG IRU WKLV SURFHVV $OWHUQDWLQJ SHULRGV ZLWK KLJK DQG ORZ UDWHV RI VHGLPHQWDWLRQ DUH UHVSRQVLEOH IRU WKH IRUPDWLRQ RI WKH ODPLQDWHG VWUXFWXUH RI WKH OLWKLILHG PLFURELDO PDWV ZKLFK FRXOG WKHUHIRUH EH WHUPHG VWURPDWROLWHV
9,, 1LWURJHQ0HWDEROLVPDQG1LWURJHQ )L[DWLRQ
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
% 7KH1LWURJHQ&\FOHLQ0LFURELDO0DWV 1LWURJHQ RFFXUV LQ GLIIHUHQW FKHPLFDO R[LGDWLRQ VWDWHV YDU\LQJ IURP LWV PRVW UHGXFHG IRUP DPPRQLD 1+ WR K\GUR[\ODPLQH 1+2+ GLQLWURJHQ 1 QLWURXV R[LGH 12 QLWULF R[LGH 12 QLWULWH 12 WR LWV PRVW
Fig. 17. 7KH QLWURJHQ F\FOH LQ D F\DQREDFWHULDO PDW &\DQREDFWHULDWDNHXSDQG DVVLPLODWHDPPRQLXPLQWR DPLQRDFLGV $$ ZKLFK DUH XVHG IRU SURWHLQ V\QWKHVLV RU FDQ EH VWRUHG DV F\DQRSK\FLQ $PLQR DFLGV DQG DPPRQLD PD\ OHDN RXW WKH FHOOV DQGEHR[LGL]HGWRQLWUDWHE\QLWULI\LQJEDFWHULD ,QWKHDQR[LFSDUW RI WKH PDW QLWUDWH LV FRQYHUWHG WR GLQLWURJHQ E\ GLQLWULI\LQJ EDFWHULD 1LWUDWH FDQ EH WDNHQ XS E\ WKH F\DQREDFWHULD DQG DVVLPLODWHG 1LWURJHQIL[LQJ F\DQREDFWHULD UHGXFH 1 WR DPPRQLXP
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
/- 6WDO
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
&1LWURJHQDVH ,Q DOO QLWURJHQIL[LQJ RUJDQLVPV WKH HQ]\PH FRPSOH[ QLWURJHQDVH LV SUHVHQW 7KLV HQ]\PH LV YHU\ VLPLODU LQ DOO RUJDQLVPV WKDW FRQWDLQ LW 7KH FRPSOH[ LV FRPSRVHG RI WZR HQ]\PHV GLQLWURJHQDVH UHGXFWDVH ZKLFK LVDGLPHU RI LGHQWLFDOVXEXQLWVDQG DOVR WHUPHG WKH LURQSURWHLQ DQG GLQLWURJHQDVH D WHWUDPHU FRPSRVHG RI WZR GLIIHUHQW VXEXQLWV 'LQLWURJHQDVH LV DOVR NQRZQ DV WKH PRO\EGHQXPLURQ SURWHLQ+RZDUGDQG 5HHV 1LWURJHQDVHFDWDO\]HVWKHIROORZLQJ UHDFWLRQ N2 + 8e + 8H+ + 16 MgATP 2 NH3 + H2 + 16MgADP + 16 3i 5HGXFHG IHUUHGR[LQ LV WKH HOHFWURQ GRQRU RI QLWURJHQDVH 7KH HTXDWLRQ VKRZQ DERYH PDNHV FOHDU WKDW WKH IL[DWLRQ RI QLWURJHQ LV DW WKH H[SHQVH RI FRQVLGHUDEOH DPRXQW RI HQHUJ\ DQG ORZ SRWHQWLDO HOHFWURQV 7KLV KLJK HQHUJ\ GHPDQG RI QLWURJHQDVH SUHVHQWV RIWHQ D SUREOHP IRU GLD]RWURSKLF RUJDQLVPV H[FHSW IRU F\DQREDFWHULD ZKR SURGXFH UHGXFHG
IHUUHGR[LQ DQG FRQYHUW OLJKW HQHUJ\ LQWR FKHPLFDO HQHUJ\ GXULQJ SKRWRV\QWKHVLV +RZHYHU DOO QLWURJHQDVHV DUH H[WUHPHO\ VHQVLWLYH WR R[\JHQ DQG WKHUHIRUH GLD]RWURSKLF RUJDQLVPV PXVW SURYLGH DQ DQDHURELF HQYLURQPHQW LQ RUGHU WR EH DEOH WR IL[ QLWURJHQ &\DQREDFWHULD DV R[\JHQLF SKRWRWURSKLF DQG SULQFLSDOO\ DHURELF RUJDQLVPV QHHG VSHFLDO DGDSWDWLRQV
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alothrix KDV EHHQ IRXQG DV WKH GRPLQDQW RUJDQLVP LQ PLFURELDO PDWV LQ WKHWLGDO DUHD RI WKH 3DFLILF FRDVW LQ %DMD &DOLIRUQLD 6XU 0H[LFR Calothrix LV DOVR NQRZQ IURP D YDULHW\ RI RWKHU PDULQH DQG EUDFNLVK KDELWDWV VXFK DV WKH VSUD\ ]RQH
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
Table 3. 7\SHVDQGFKDUDFWHULVWLFV RIQLWURJHQIL[LQJF\DQREDFWHULD
+HWHURF\VWRXV&\DQREDFWHULD H[FOXVLYHO\ ILODPHQWRXV VSHFLHV WKDWGLIIHUHQWLDWH VSHFLDO FHOOV KHWHURF\VWV VWUDWHJ\ VSDWLDO VHSDUDWLRQRI1 IL[DWLRQDQGR[\JHQLFSKRWRV\QWKHVLV DQGSURWHFWLRQRIQLWURJHQDVHLQWKHKHWHURF\VW GLD]RWURSKLF JURZWK XQGHU IXOO\ DHURELF FRQGLWLRQV H[DPSOHV Anabaena, Nostoc, Aphanizomenon, Nodularia, Calothrix, Scytonema RFFXUUHQFHZDWHUEORRPVIUHVKZDWHUODNHVDQGEUDFNLVKVHDV SDGG\ILHOGVPLFURELDOPDWVV\PELRWLFZLWKDYDULHW\RI GLIIHUHQW RUJDQLVPV
Type I
$QDHURELF 1)L[LQJ1RQ+HWHURF\VWRXV &\DQREDFWHULD ILODPHQWRXVDQGXQLFHOOXODUVSHFLHV VWUDWHJ\DYRLGDQFHRIR[\JHQ LQGXFWLRQDQGPDLQWHQDQFHRIQLWURJHQDVHRQO\XQGHUDQR[LDRUORZR[\JHQ VXOILGHPD\EHQHFHVVDU\LQRUGHUWRLQKLELW R[\JHQLF SKRWRV\QWKHVLV H[DPSOHV Plectonema boryanum, Oscillatoria limnetica, Synechococcus VSPDQ\RWKHUF\DQREDFWHULD
Type II
RFFXUUHQFHPDQ\GLIIHUHQWHQYLURQPHQWVSDUWLFXODUO\LQPLFURELDOPDWV
Type III $HURELF1)L[LQJ1RQ+HWHURF\VWRXV &\DQREDFWHULD
ILODPHQWRXVDQGXQLFHOOXODUVSHFLHV
VWUDWHJ\QRWSUHFLVHO\NQRZQSRVVLEO\ WHPSRUDOVHSDUDWLRQRI1IL[DWLRQDQGR[\JHQLFSKRWRV\QWKHVLVLQFRQFHUWZLWK RWKHUR[\JHQSURWHFWLRQPHFKDQLVPV
GLD]RWURSKLFJURZWKSRVVLEOHXQGHUIXOO\DHURELFFRQGLWLRQV
H[DPSOHV Gloeothece, Oscillatoria, Trichodesmium, Lyngbya, Microcoleus
RFFXUUHQFHWURSLFDORFHDQ7ULFKRGHVPLXP FDUERQDWHFDYHZDOOVDQGSDGG\ILHOGV*ORHRWKHFH PLFURELDOPDWV Oscillatoria, Lyngbya, Microcoleus DIWHU 6WDO
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
QLWURJHQDVH LV LQFRPSOHWH $OWKRXJK WKH VSHFLHV WKDW SRVVHVV WKLV FDSDELOLW\ DUH VWLOO UHODWLYHO\ UDUH WKH QXPEHUV DUH LQFUHDVLQJ DW VWHDG\ SDFH ([DPSOHV FDQ EH IRXQG LQ WHUUHVWULDO HQYLURQPHQWV VXFK DV FDYH ZDOOV DQG SDGG\ ILHOGV DQG LQ WKH PDULQHHQYLURQPHQW )UHVKZDWHU ODNHV DSSDUHQWO\ GR QRW KDUERXU DHURELF QLWURJHQIL[LQJ QRQKHWHURF\VWRXV F\DQREDFWHULD ,Q WKH RFHDQ WKH SODQNWRQLF FRORQ\IRUPLQJ Trichodesmium VSS Oscillatoria VSS LV NQRZQ DV DQ HIILFLHQW GLD]RWURSKLF JURZLQJ QRQKHWHURF\VWRXV F\DQREDFWHULXP ,Q PLFURELDO PDWV DHURELF QLWURJHQ IL[LQJ QRQKHWHURF\VWRXV F\DQREDFWHULD DUH UHSRUWHG WR EHORQJ SUHGRPLQDQWO\ WR WKH PRUSKRORJLFDOO\ FORVHO\ UHODWHGJHQHUDOscillatoria DQG Lyngbya. 7KH ILUVW UHSRUW RI D FXOWXUH RI D ILODPHQWRXV QRQ KHWHURF\VWRXV DHURELF QLWURJHQIL[LQJ F\DQREDFWHULXP ZDV E\ 3HDUVRQ HW DO 7KLV RUJDQLVP ZDV RULJLQDOO\ LGHQWLILHG DV Microcoleus chthonoplastes EXW ODWHU UHQDPHG DV Symploca VS 5 5LSSND SHUV FRPP Symploca LV DOVRPRUSKRORJLFDOO\UHODWHG WR Oscillatoria, DQG ZDV LVRODWHG IURP D WLGDO PLFURELDO PDW 3HDUVRQ HW DO 0DOLQDQG 3HDUVRQ ,W KDV EHHQSURSRVHGWKDWWKH VWUDWHJ\RI DHURELF QRQ KHWHURF\VWRXV F\DQREDFWHULD LQ DQDORJ\ ZLWK WKH KHWHURF\VWRXV VSHFLHV LV WHPSRUDO VHSDUDWLRQ RI WKH LQFRPSDWLEOH SURFHVVHV RI SKRWRV\QWKHVLV DQG QLWURJHQ IL[DWLRQ 7KH ODWWHU ZRXOG WKDQ RFFXU GXULQJ WKH GDUN0XOOLQHDX[HW DO 6WDO DQG .UXPEHLQ +RZHYHU QRW DOO VSHFLHV LQ WKLV JURXS IROORZ WKLV VWUDWHJ\ Trichodesmium VSS IL[ QLWURJHQ GXULQJ WKH GD\ &DSRQH HW DO 0RUHRYHU DOO VSHFLHV WKDW KDYH EHHQ FXOWXUHG DUH FDSDEOH RI JURZLQJ GLD]RWURSKLFDOO\ XQGHU FRQWLQXRXV OLJKW DQG LQ WKH XQLFHOOXODU Gloeothece VS FXOWXUH FRQGLWLRQV FDQ EH FKRVHQ XQGHU ZKLFK QLWURJHQ IL[DWLRQ RFFXUV GXULQJ WKH OLJKW SHULRG RI D OLJKW GDUN F\FOH 2UWHJD&DOYR DQG 6WDO
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loeothece DQG Oscillatoria. :KHUHDVGXULQJWKHGD\ PLFURELDO PDWV RIWHQ EHFRPH H[WUHPHO\ VXSHUVDWXUDWHG ZLWK R[\JHQ EHFDXVHWKH GLIIXVLRQ RI WKLV JDV LV OLPLWHG DW QLJKW WKH\ PD\ WXUQ DQR[LF ZLWKLQ PLQXWHV 6WDO 7KH PLFURELDO FRPPXQLW\ LQFOXGLQJ WKH F\DQREDFWHULD FRQVXPHV R[\JHQLQWKHGDUNE\ UHVSLUDWLRQ2EYLRXVO\DQR[LF
&KDSWHU &\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV
Fig. 18. )LYH W\SLFDO SDWWHUQV RI GDLO\ YDULDWLRQV RI QLWURJHQ IL[DWLRQ DFHW\OHQH UHGXFWLRQ DFWLYLW\ $5$ UHODWLYH XQLWV LQ PLFURELDO PDWV A: PDW RI WKH KHWHURF\VWRXV F\DQREDFWHULXP Calothrix VS LQ %DMD &DOLIRUQLD 0H[LFR GDWD IURP 6WDO HW DO B: PDW RIWKH XQLFHOOXODU F\DQREDFWHULXP Gloeothece VS RQ WKHZDOO RIDFDUERQDWH FDYH GDWDIURP*ULIILWKV HW DO C: PL[HGPDWRIWKHQRQKHWHURF\VWRXVF\DQREDFWHULD Microcoleus chthonoplastes DQG Oscillatoria limosa IURP D WLGDO IODW RQ WKH 1RUWK 6HDLVODQGRI0HOOXPGDWDIURP9LOOEUDQGWHWDO D: PDWGRPLQDWHG E\ Oscillatoria limosa ORFDWLRQDV C GDWDIURP 9LOOEUDQGW HW DO ( PDW RI WKH QRQKHWHURF\VWRXV F\DQREDFWHULXP /\QJE\D DHVWXDULL ORFDWLRQ DV $ 6WDO XQSXEOLVKHG
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scillatoria limosa 6WDO DQG +H\HU ,Q PLFURELDO PDWV LQ ZKLFK WKLV F\DQREDFWHULXP RFFXUUHG GDLO\ SDWWHUQV RI QLWURJHQ IL[DWLRQZHUHIRXQGLQZKLFKWKLVDFWLYLW\ZDVORZEXW WRWDOO\ FRQILQHG WR WKH GDUN 9LOOEUDQGW HW DO +RZHYHU RWKHU SDWWHUQV ZHUH DOVR REVHUYHG DW GLIIHUHQW WLPHV LQ WKH VDPH PDWV ZLWK WKH VDPH RUJDQLVP )RU LQVWDQFH LW FRXOG RIWHQ EH VHHQ WKDW QLWURJHQDVHDFWLYLW\SHDNHGDURXQGVXQVHWDQG VXQULVH )LJ F 7KLV ZDV FRQILUPHG E\ H[SHULPHQWV ZLWK O. limosa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limosa LV RIWHQ WKH SLRQHHU F\DQREDFWHULXP6WDOHWDO 7KLVLVPRVWOLNHO\ EHFDXVH RI LWV FDSDFLW\ WR JURZ GLD]RWURSKLFDOO\ ,Q WKLV SLRQHHU VWDWH ELRPDVV LV ORZ DQG WKHUHIRUH VR LV WKH R[\JHQ GHPDQG LQ WKH GDUN 6XFK VHGLPHQWV QRUPDOO\ GR QRW WXUQ DQR[LF +RZHYHU GXULQJ WKH
OLJKW WKH\ PD\ DFFXPXODWH R[\JHQ XS WR VHYHUDO IROG VDWXUDWLRQ 9LOOEUDQGW HW DO 1LWURJHQ IL[DWLRQ LQ VXFK V\VWHPV LV W\SLFDOO\ FRQILQHG WR WKH QLJKW SHDNLQJ DW VXQULVH ZKHQ OLJKW EHFRPHV DYDLODEOH EXW DW R[\JHQ OHYHOV ZHOO EHORZ DLUVDWXUDWLRQ )LJ G ,Q RWKHU V\VWHPV VXFK DV LQ PDWV RI WKH XQLFHOOXODU QLWURJHQIL[LQJ Gloeothece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a KLJKHVWVSHFLILF QLWURJHQDVH DFWLYLW\ ZDV SUHVHQW LQ WKH ORZHVW OD\HU RI WKH F\DQREDFWHULDO PDW PP )LJ &\DQREDFWHULDO ELRPDVV ZDV KLJKHVW LQ WKH WRS OD\HU GHFUHDVLQJ JUDGXDOO\ XQWLO DERXW PP GHSWK 7KXV LW LV OLNHO\ WKDW D VSDWLDO VHSDUDWLRQ RIQLWURJHQ IL[DWLRQ DQG R[\JHQLFSKRWRV\QWKHVLVKDG RFFXUUHG LQWKLVPDW 7KHWRSOD\HU FDUULHV RXWR[\JHQLFSKRWRV\QWKHVLV DQG &2 IL[DWLRQ ZKLOH QLWURJHQ LV IL[HG LQ WKH ORZHU OD\HUV
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scillatoria limnetica LV FDSDEOH RI QLWURJHQ IL[DWLRQ ZKHQ FDUU\LQJ RXW DQR[\JHQLF SKRWRV\QWKHVLV VXOILGHGHSHQGHQW 9LOOEUDQGW DQG 6WDO LQYHVWLJDWHG WKH HIIHFW RI VXOILGH RQ QLWURJHQ IL[DWLRQ LQ F\DQREDFWHULDO PDWV DQG RQ FXOWXUHV RI F\DQREDFWHULD LVRODWHG IURP WKHVH PDWV 7KH\ FRPSDUHG D PDW GRPLQDWHG E\ D KHWHURF\VWRXV F\DQREDFWHULXP Anabaena ZLWK DQRWKHU GRPLQDWHG E\ QRQKHWHURF\VWRXV ILODPHQWRXV RUJDQLVPV Oscillatoria DQG Phormidium 6XOILGH LQKLELWHG QLWURJHQDVH DFWLYLW\ LQ WKH PDW RI Anabaena EXW JUHDWO\ VWLPXODWHG LW LQ WKH PDW RI WKH QRQ KHWHURF\VWRXV F\DQREDFWHULD)LJ %RWK OLJKW DQG GDUN QLWURJHQDVH DFWLYLW\ ZDV LQKLELWHG E\ VXOILGH LQ
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV
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nabaena VS LVRODWHG IURPWKLV PDW WKH HIIHFW RI VXOILGH VWURQJO\ GHSHQGHG RQ WKH S+ $W S+ D WRWDO VXOILGH FRQFHQWUDWLRQ RI P0 KDG QR HIIHFW RQ QLWURJHQ IL[DWLRQ EXW DW S+ WKLV FRQFHQWUDWLRQDOPRVWFRPSOHWHO\LQKLELWHGQLWURJHQDVH DFWLYLW\ 7KLV VKRZV WKDW WKH HIIHFW RI VXOILGH LV WKURXJK WKH JDVHRXV VSHFLHV +6 7KLV JDV ZLOO SDVVLYHO\ GLIIXVH LQWR WKH FHOO %HFDXVH WKH S+ LQ WKHVH PDWV LV XVXDOO\ KLJK YHU\ OLWWOH +6 ZLOO EH SUHVHQW HYHQZKHQWKHWRWDOFRQFHQWUDWLRQRIVXOILGH LVKLJK
$5$QPRO&+ K J&KO Fig. 19. 9HUWLFDO GLVWULEXWLRQ RI FKORURSK\OO D A SRWHQWLDO QLWURJHQDVHDFWLYLW\DFHW\OHQHUHGXFWLRQ$5$ B DQGVSHFLILF FKORURSK\OODEDVHG$5$C LQDPLFURELDOPDWGDWDIURP6WDOHW DO
WKH PDW RI Anabaena VS EXW ZKHQ '&08 ZDV DGGHG LQ RUGHU WR LQKLELW R[\JHQLF SKRWRV\QWKHVLV YLUWXDOO\ QR HIIHFW RI VXOILGH RQ QLWURJHQ IL[DWLRQ ZDV VHHQ LQ WKLV PDW 7KHUHIRUH WKH HIIHFW RI VXOILGH ZDV PDLQO\ WKURXJK WKH LQKLELWLRQ RI R[\JHQLF SKRWRV\QWKHVLV DQG
Fig. 20. (IIHFW RI VXOILGH RQ QLWURJHQDVH DFWLYLW\ DFHW\OHQH UHGXFWLRQ$5$ LQA DPDWRI WKHKHWHURF\VWRXVF\DQREDFWHULXP $QDEDHQD VS ODJRRQ $WODQWLF FRDVW RI)UDQFH DQG B DPDW RI QRQKHWHURF\VWRXV F\DQREDFWHULD Oscillatoria VS DQG Phormidium VS ODJRRQ 0HGLWHUUDQHDQ FRDVW RI )UDQFH GDWD IURP9LOOEUDQGWDQG6WDO
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hormidium LW ZDV VKRZQ WKDW VXOILGH WRWDO FRQFHQWUDWLRQXS WR P0 KDG QR HIIHFWRQQLWURJHQDVHDFWLYLW\ZKHQ WKLV ZDV LQGXFHG DQDHURELFDOO\ ZLWK '&08 7KHUHIRUHLW ZDV FRQFOXGHGWKDWVXOILGHGLGQRWDFWDVDQ HOHFWURQ GRQRU WR QLWURJHQDVHDQG WKDW WKH VWLPXODWRU\ HIIHFW REVHUYHG LQ WKH PDW ZDV PRVW SUREDEO\ GXH WR WKH VFDYHQJLQJ RI HQYLURQPHQWDO R[\JHQ 6XOILGH YHU\ HIILFLHQWO\ LQGXFHG QLWURJHQDVH LQ Phormidium DQG RWKHU QRQKHWHURF\VWRXVF\DQREDFWHULDZLWK DQDHURELF QLWURJHQDVH $ERXW P0WRWDOVXOILGHZDV VXIILFLHQW IRU IXOO LQGXFWLRQRIQLWURJHQDVH+RZHYHULQFRQWUDVW ZLWK ZKDW ZDV VHHQ ZLWK WKH KHWHURF\VWRXV F\DQREDFWHULXP DQG WR ZKDW ZDV H[SHFWHG LW DSSHDUHG WKDW LQGXFWLRQ RI QLWURJHQDVH ZLWK VXOILGH ZDV RSWLPDO DW KLJK S+ ZKLFK PHDQV WKDW WKH LRQV +6 DQGRU 6 ZHUH PRUH HIILFLHQW WKDQ WKHJDV+6 7KLV VXJJHVWHG WKDW Phormidium FRXOGDFWLYHO\WDNH XS VXOILGH LRQ 7KXV WKHXSWDNHRI VXOILGHLRQPD\ EH HVVHQWLDO WR DOORZ GLD]RWURSKLF JURZWK LQ WKHVH RUJDQLVPV
+ 2[\JHQ3URWHFWLRQRI1LWURJHQDVHLQ 0LFURELDO0DWV %HFDXVH F\DQREDFWHULDO PDWV RIWHQ FRQWDLQ D KLJK GHQVLW\ RI ELRPDVV DQG KDYH ORZ UDWHV RI PROHFXODU
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scillatoria, Lyngbya, Microcoleus, Gloeothece, Cyanothece DQG Synechococcus %HUJPDQ HW DO $PRQJ WKH GLIIHUHQW PHFKDQLVPV WKDW KDYH EHHQ SURSRVHG IRU R[\JHQ SURWHFWLRQ RI QLWURJHQDVH WKH XSWDNH DQG UHGXFWLRQ RIR[\JHQ VHHPV WR EH WKHPRVWSURPLVLQJ
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV 6XFK V\VWHPV PD\ DFW LQ FRQFHUW ZLWK HQ]\PHV WKDW UHPRYH R[\JHQUDGLFDOV ,Q 7DEOH WKH HIIHFWV RIGLIIHUHQW WUHDWPHQWV RI D GLD]RWURSKLF PLFURELDO PDW FRPSRVHG RI Oscillatoria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
LW LV SRVVLEOHWKDWRWKHUPHWDEROLFSURFHVVHVPD\ KDYH EHHQ UHVSRQVLEOHIRUWKHGHFUHDVHRIR[\JHQWKLV PD\ DOVRKDYHEHHQWKHUHVXOWRI WKHLQFUHDVHGQLWURJHQDVH DFWLYLW\LQWKHILUVWSODFH 2QHSRVVLELOLW\ WKDW PXVW QRZ EH LQYHVWLJDWHG LV WKH FDSDFLW\ RI QLWURJHQDVH LQ WKHVHF\DQREDFWHULDWR UHGXFH R[\JHQ DXWRSURWHFWLRQ %HUJPDQ HW DO 7KLVFDXVHVWKHUHGXFWLRQRI2 WR +2 ZKLFK FDQ EHIXUWKHUUHGXFHGE\ SHUR[LGDVHV
Table 4. (IIHFW RI GLIIHUHQW WUHDWPHQWV RI D QLWURJHQIL[LQJ PLFURELDO PDW RI Oscillatoria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
7UHDWPHQW 5HIHUHQFH +LJKVDOLQLW\ 3KRVSKDWHIHUWLOL]DWLRQ 7LGDOPRYHPHQW
1LWURJHQDVHDFWLYLW\ QPRO &+ FP K
Fig. 21. 9HUWLFDO SURILOHV RI R[\JHQ LQ GLD]RWURSKLF PDWV RI Oscillatoria VSVXEMHFWHGWRGLIIHUHQWWUHDWPHQWV6HH7DEOH
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aestuarii DQG FRYHUVWKHORZHU DUHDV RI WKH WLGDO IODW 2Q WKH XSSHU WLGDO IODW DQG RQ VOLJKWO\ HOHYDWHG VSRWV D SXVWXODU PDW GHYHORSV ZKLFK LV FRPSRVHG RI WKH KHWHURF\VWRXV F\DQREDFWHULXP Calothrix. %RWK PDWV IL[ QLWURJHQ EXW VKRZGLVWLQFW GLIIHUHQFHV LQ WKHLU GDLO\ QLWURJHQDVH SDWWHUQV Calothrix IL[HV SUHGRPLQDQWO\ GXULQJWKHGD\ ZKLOH QLWURJHQDVH DFWLYLW\ LQ WKH PDWV RI / aestuarii LV FRQILQHG WR WKH QLJKW 'XH WR WKHLU ORFDWLRQVRQ WKH WLGDO IODW WKH PDWV RI L. aestuarii DUH FRYHUHG PRUH RIWHQ DQG GXULQJORQJHUSHULRGVRI WLPH DW KLJK WLGH WKDQ WKH PDWV RI Calothrix. 'XULQJLQXQGDWLRQRI WKH PDWV RI L. aestuarii GLIIXVLRQLVOLPLWHG 7KLVFDXVHV R[\JHQ VXSHUVDWXUDWLRQ GXULQJ WKH SHULRG RI SKRWRV\QWKHVLV DQG DQR[LFFRQGLWLRQVDWQLJKW 7KHVH DQR[LF FRQGLWLRQV DOVR DOORZ WKH GHYHORSPHQW RI D FRPPXQLW\RIVXOIDWHUHGXFLQJEDFWHULD 7KLV PDWKDV D YHU\ GHQVH ELRPDVV DQG LV FKDUDFWHUL]HG E\ VWHHS JUDGLHQWV RI R[\JHQ DQG VXOILGHW\SLFDOIRUPLFURELDO PDWV 'XHWRWKLVGHQVHPDW VWUXFWXUHWKHJUDGLHQWVRI
/- 6WDO R[\JHQ DQG VXOILGH H[LVW UHJDUGOHVV ZKHWKHU WKH PDW LVLQXQGDWHGRU QRW 7KH VLWXDWLRQLQWKH &DORWKUL[ PDW LV WRWDOO\ GLIIHUHQW 7KLV SXVWXODU PDW KDV D SRURXV VWUXFWXUH 'XH WR WKLV VWUXFWXUH LQ WKLV PDW WKHUH LV D IUHH H[FKDQJH RI R[\JHQ ZLWK WKH DWPRVSKHUH DQG R[\JHQ VXSHU VDWXUDWLRQ RU DQR[LF FRQGLWLRQV DUH QRW XVXDOO\ WKHFDVH ,QH[FHSWLRQDO FDVHVZKHQ WKH PDW LV LQXQGDWHG IRU D SURORQJHG SHULRG RI WLPH DQR[LF FRQGLWLRQV RU R[\JHQ VXSHU VDWXUDWLRQ PD\ RFFXU EXW QRUPDOO\ WKH PDW ZLOO EH LQXQGDWHG IRU VKRUW SHULRGV RU QRW DW DOO 6WDO HW DO K\SRWKHVL]HG WKDW KHWHURF\VWRXV F\DQREDFWHULD ZRXOG QRW EH DEOH WR PDLQWDLQ WKHPVHOYHV LQ DQ HQYLURQPHQW LQ ZKLFK HLWKHU GDUN DQR[LF FRQGLWLRQV RU KLJK FRQFHQWUDWLRQV RI VXOILGH RFFXU &\DQREDFWHULD LQFDSDEOH RI IHUPHQWDWLRQ ZLOO GLH ZLWKLQ KRXUV RI GDUN DQR[LF FRQGLWLRQV 6WDO DQG 0RH]HODDU +RZHYHU WKHUH LV QR UHDVRQ ZK\ KHWHURF\VWV VKRXOG EH LQFDSDEOH RI IHUPHQWDWLRQ DQG WKLV KDV EHHQ GHPRQVWUDWHG LQ D QXPEHU RI V\PELRWLF Nostoc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alothrix DUH RULHQWDWHG YHUWLFDOO\ ZLWK WKH WHUPLQDO KHWHURF\VWV VLWXDWHG DZD\ IURP WKH VXUIDFH 7KHVH ILODPHQWV GR QRW JOLGH IUHHO\ 7KH VDPHLVWKHFDVHIRUDQRWKHU PDW EXLOGLQJ KHWHURF\VWRXV F\DQREDFWHULXP Scytonema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
/-6WDO
Fig. 22. $VLPSOLILHGVFKHPHVKRZLQJWKHUROHRIF\DQREDFWHULDLQWKHF\FOHRIVXOIXULQDPLFURELDOPDW 65% &6% DQG 36% DUHVXOIDWH UHGXFLQJEDFWHULDFRORUOHVVVXOIXUEDFWHULDDQGSKRWRV\QWKHWLFVXOIXUEDFWHULDUHVSHFWLYHO\ 6HHWH[WIRUIXUWKHUH[SODQDWLRQ
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scillatoria limnetica WKDW PD\ EH FDSDEOH RI WUXH VXOIXU UHVSLUDWLRQ2UHQDQG 6KLOR +RZHYHU IRU WKH PDMRULW\ RI F\DQREDFWHULDVXOIXU VHUYHV DV DQ HOHFWURQ VLQN GXULQJ IHUPHQWDWLRQ 2WKHU PLFURRUJDQLVPV LQ PLFURELDO PDWV WKDW FRQWULEXWH WR WKH SURGXFWLRQ RI VXOILGH LQFOXGH IHUPHQWDWLYH DQG GLVVLPLODWRU\ VXOIXU UHGXFLQJ EDFWHULD DQG VXOIXU WKLRVXOIDWH DQG VXOILWH GLVSURSRUWLRQDWLQJ EDFWHULD -¡UJHQVHQ
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amphigranulata R[\JHQLF SKRWRV\QWKHVLV UHFRYHUHG DIWHU LQLWLDO LQKLELWLRQ E\ PRGHUDWH FRQFHQWUDWLRQV RI VXOILGH 7KLV UHFRYHU\ RFFXUUHG DW KLJK OLJKW LQWHQVLWLHVDQGGHSHQGHGRQde novo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nacystis nidulans DQG Plectonema boryunum LQ ZKLFK &2 IL[DWLRQ ZDV LQKLELWHG DW DQG 0 &RKHQ HW DO ,Q *URXS F\DQREDFWHULD DUH UHSUHVHQWHG WKDW DUH LQFDSDEOH RI DQR[\JHQLF
Table 5. *URXSV RI F\DQREDFWHULD ZLWK GLIIHUHQW W\SHV RI DGDSWDWLRQWRVXOILGH *URXS
6XOILGHVHQVLWLYH R[\JHQLF SKRWRV\QWKHVLV RQO\
*URXS
6XOILGHUHVLVWDQW R[\JHQLF SKRWRV\QWKHVLV RQO\
*URXS
6XOILGHLQVHQVLWLYH R[\JHQLF SKRWRV\QWKHVLV FRQFXUUHQW ZLWK VXOILGH GHSHQGHQW DQR[\JHQLF SKRWRV\QWKHVLV
6XOILGHVHQVLWLYH R[\JHQLF SKRWRV\QWKHVLV UHSODFHG E\ VXOILGH GHSHQGHQW DQR[\JHQLF SKRWRV\QWKHVLV $IWHU&RKHQHWDO DQG6WDO
*URXS
SKRWRV\QWKHVLV EXW WKDW UHVLVW FRQVLGHUDEOH OHYHOV RI VXOILGH 2[\JHQLF SKRWRV\QWKHVLV LQ WKHVH RUJDQLVPV LV RIWHQ VWLPXODWHG DW PRGHUDWH P0 VXOILGH FRQFHQWUDWLRQ 7KLV W\SH RI DGDSWDWLRQ LV W\SLFDO IRU PDULQH PLFURELDO PDWV ZLWK IOXFWXDWLQJ VXOILGH FRQFHQWUDWLRQV 7KH PDWIRUPLQJ DQG GLD]RWURSKLF F\DQREDFWHULXP Oscillatoria limosa LV D W\SLFDO H[DPSOH RI WKLV JURXS 6WDO $OVR *URXS F\DQREDFWHULDDUHW\SLFDOO\IRXQG LQPDULQHPLFURELDO PDWV 7KHVH F\DQREDFWHULD DUH FKDUDFWHUL]HG E\ R[\JHQLF SKRWRV\QWKHVLV VXOILGHLQVHQVLWLYH FRQFXUUHQW ZLWK VXOILGHGHSHQGHQW DQR[\JHQLF SKRWRV\QWKHVLV 7KH FRVPRSROLWDQ PDWIRUPLQJ F\DQREDFWHULXP Microcoleus chthonoplastes EHORQJV WR WKLV JURXS 'H :LW DQG 9DQ *HPHUGHQ 2[\JHQLFSKRWRV\QWKHVLV LQ*URXSF\DQREDFWHULDLV DV VHQVLWLYH WR VXOILGH DV LQ WKRVH F\DQREDFWHULD EHORQJLQJ WR *URXS 7KH GLIIHUHQFH LV LQ WKHLU FDSDFLW\ IRU FDUU\LQJ RXW VXOILGHGHSHQGHQW DQR[\JHQLF SKRWRV\QWKHVLV 7KH VXOILGH WROHUDQFH RI WKLV JURXS RIF\DQREDFWHULD YDULHV FRQVLGHUDEO\ IURP OHVV WKDQ P0 WR P0 Oscillatoria limnetica LV WKH EHVWVWXGLHG F\DQREDFWHULXP EHORQJLQJ WR WKDW JURXS 3KRWRV\VWHP ,, LQ WKLV F\DQREDFWHULXP LV VZLWFKHG RIIZKHQ H[SRVHG WR P0 RIVXOILGH $QR[\JHQLF SKRWRV\QWKHVLV LV LQGXFHG LQ D SURFHVV UHTXLULQJSURWHLQ V\QWKHVLV O. limnetica WROHUDWHV XS WR P0 VXOILGH EXW DQR[\JHQLF SKRWRV\QWKHVLV LV JUDGXDOO\ LQKLELWHG DW FRQFHQWUDWLRQV H[FHHGLQJ P0 ,Q PLFURELDO PDWV PRVW RIWKH VXOILGH LV SUHVHQWDV µDFLGYRODWLOH VXOILGH¶ $96 ZKLFK LV PRVWO\ LQ WKH IRUP RIIHUURXV VXOILGH )H6 )LJ ,Q WKLV IRUP VXOILGHLVYLUWXDOO\LQVROXEOH 2QO\IUHHVXOILGHPD\EH WR[LF )UHH GLVVROYHG VXOILGH RFFXUV DV K\GURJHQ VXOILGHRUVXOILGHLRQVLQDS+GHSHQGHQWHTXLOLEULXP
H 2S
HS
S
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¡UJHQVHQ DQG &RKHQ +RZHYHU 9LVVFKHU PHDVXUHG YHU\ KLJK FRQFHQWUDWLRQV RI SRO\VXOILGHV LQ D F\DQREDFWHULDO PDW LQGLFDWLQJ WKDW WKLV FRPSRXQG PD\ EH PRUH FRPPRQ WKDQ SUHYLRXVO\ DVVXPHG :KLOH RQ WKH RQH KDQG SRO\VXOILGHV DUH DQ RUGHU RI PDJQLWXGH PRUH WR[LF IRU PRVW RUJDQLVPV WKDQ VXOILGH RQ WKH RWKHU KDQG WKH\ PD\ VHUYH DV WKH IRUP RI HOHPHQWDO VXOIXU WKDW LVWUDQVSRUWHGLQFHOOV6WHXGHOHW DO
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hiobacillus ferrooxidans /HGXF DQG )HUURQL 8QGHU QHXWUDO FRQGLWLRQV IHUURXV LURQ LV UDSLGO\ R[LGLVHG E\ R[\JHQ KRZHYHU Gallionella ferruginea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e3+ + H2S
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
2 Fe2+ + 2 H2S 2 Fe3+ + 3 H2S
2 Fe2+ + S° + 2 H+ 2 FeS + 4H+ FeS + S° + 6H+
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¶V LV DERXW 7RJHWKHU LURQ R[LGHV DQG VLOLFD PD\ PDNH XS WR RI WKH ZHLJKWRI %,) ,URQ R[LGHV DQG VLOLFD FKHUW RFFXU LQ DOWHUQDWLQJ OD\HUV 7KH FKHUW\ EDQGHG LURQ IRUPDWLRQ RI +DPHUVOH\ %DVLQ $XVWUDOLD LV RQH RI WKH ODUJHVW LQ WKH ZRUOG DQG LV FKDUDFWHUL]HG E\ VWUDWLILFDWLRQ DW GLIIHUHQW VFDOHV $WWKHPLOOLPHWHU VFDOH PLFUREDQGV RI LURQ PLQHUDOV DUH UHFRJQL]HG VHSDUDWHG DW WKH
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¶V DWPRVSKHUH ZKLFK VWDUWHG 0\ DJR 2QH PHFKDQLVP IRU %,) IRUPDWLRQ PD\ EHWKHFKHPLFDO R[LGDWLRQRIIHUURXV LURQZLWK R[\JHQ HYROYHG E\ R[\JHQLF SKRWRV\QWKHVLV PRVW OLNHO\ E\ F\DQREDFWHULD 7KH KXJH DPRXQWV RI IHUURXV LURQ LQ WKH HDUWK¶V FUXVW ZRXOG DFW OLNH D EXIIHU DQG SUHYHQW WKH R[\JHQDWLRQ RI WKH DWPRVSKHUH XQWLO PRVW RI WKH LURQ ZDV R[\JHQDWHG $OWKRXJK OHVVDEXQGDQWWKHIDFW WKDW %,)¶V DUH DOVR NQRZQ IURP WKH PLG $UFKHDQ 0\ PLJKW LQGLFDWH RWKHU PHFKDQLVPV *HRORJLFDO HYLGHQFH LV DYDLODEOH WKDW LQGLFDWHV WKDW XQWLO 0\DJRWKHHDUWK¶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¶V DWPRVSKHUH LW LV LQWHUHVWLQJ WKDW VRPH PDWIRUPLQJ F\DQREDFWHULD VXFK DV 0LFURFROHXV FKWKRQRSODVWHV DFFXPXODWH ODUJH DPRXQWV RI LURQ DW WKH RXWHU SRO\VDFFKDULGH VKHDWK 6WDO ,URQ LV ERXQG WR QHJDWLYHO\ FKDUJHG SRO\VDFFKDULGHV SDUWLFXODUO\ WKURXJK WKH SUHVHQFH RI XURQLF DFLGV %HQGHU HW DO
/- 6WDO
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chthonoplastes LVDOVRFDSDEOHRI UHGXFLQJIHUULF LURQ 6WDO
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
2 FePO 4 3
2 FeS 2 H 3PO4 S º
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
/- 6WDO
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
PXFXV FDOFLILFDWLRQ WKHUHIRUH QRW EHLQJ LQKLELWHG )XWXUH UHVHDUFK VKRXOG LQYHVWLJDWH WKH QLWURJHQ VWDWH RI PRGHUQ FDOFLI\LQJ PLFURELDO PDWV DQG WKH UROH RI (36 DV DQWLFDOFLILFDWLRQ DJHQWV LQ QRQOLWKLI\LQJ PDWV
$FNQRZOHGJHPHQWV , ZRXOG OLNHWR WKDQN'U3 :HVWEURHNZKR GUDZP\ DWWHQWLRQWR WKHFRQFHSWRI µDQWLFDOFLILFDWLRQ¶ DQG'UV $:'HFKR+:3DHUO-3LQFNQH\DQG35HLGIRU WKH GLVFXVVLRQV GXULQJ D UHVHDUFK FUXLVH LQ WKH %DKDPDV LQ WKDW OHG WR WKH IRUPXODWLRQ RI D PRGHO IRU FDOFLILFDWLRQ LQ WKH ([XPD &D\V VWURPDWROLWHV , DP DOVR JUDWHIXO WR 'U - 0LGGHOEXUJ IRUKLVFRQVWUXFWLYHFRPPHQWV
5HIHUHQFHV $OEHUWDQR 3 DQG .RYDFLN / /LJKW DQG WHPSHUDWXUH UHVSRQVHVRIWHUUHVWULDOVFLDSKLORXV VWUDLQVRILeptolyngbya VS LQ FURVVJUDGLHQWFXOWXUHV$OJRO6WXG $OOHQ00 DQG 6PLWK$- 1LWURJHQ FKORURVLV LQ EOXHJUHHQ DOJDH $UFK 0LNURELRO $UWXV 11 6RPHUYLOOH 6& DQG 6RPHUYLOOH &5 7KH ELRFKHPLVWU\DQGFHOOELRORJ\RISKRWRUHVSLUDWLRQ &5&&ULW5HY 3ODQW6FL $ZUDPLN60 $QFLHQWVWURPDWROLWHVDQGPLFURELDOPDWV ,Q &RKHQ Y, &DVWHQKRO]5: DQG +DOYRUVRQ +2 HGV 0LFURELDO 0DWV6WURPDWROLWHVSS $ODQ5/LVV,QF1HZ
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miliania huxleyi. (XU-%LRFKHP %RUPDQ $+ 'H-RQJ (: 7KLHUU\ 5 :HVWEURHN3 DQG %RVFK / &RFFROLWKDVVRFLDWHG SRO\VDFFKDULGHV IURP FHOOV RI Emiliania huxleyi +DSWRSK\FHDH -3K\FRO %RXGUHDX %3 DQG &DQILHOG '( $ FRPSDULVRQ RI FORVHG DQGRSHQV\VWHPPRGHOV IRUSRUHZDWHUS+DQGFDOFLWHVDWXUDWLRQ VWDWH*HRFKLP&RVPRFKLP$FWD %URFN 7' (IIHFW RI ZDWHU SRWHQWLDO RQ D Microcoleus &\DQRSK\FHDH IURPDGHVHUWFUXVW-3K\FRO %URFN 7' 7KHUPRSKLOLF 0LFURRUJDQLVPV DQG /LIHDW +LJK 7HPSHUDWXUHV6SULQJHU9HUODJ1HZ
&DVWHQKRO] 5: 7KH HIIHFW RI VXOILGH RQ WKH EOXH JUHHQ DOJDHRIKRW VSULQJV , 1HZ=HDODQG DQG ,FHODQG - 3K\FRO &DVWHQKRO] 5: 7KH HIIHFW RI VXOILGH RQ WKH EOXHJUHHQ DOJDH RIKRW VSULQJV ,,
'H 3KLOLSSLV 5 0DUJKHUL 0& DQG 9LQFHQ]LQL 0 )HUPHQWDWLRQ LQ V\PELRWLF DQG IUHHOLYLQJ F\DQREDFWHULD $UFK +\GURELRO 'H :LQGHU % 0DWWKLMV +&3 DQG 0XU /5 D 7KH UROH RI ZDWHUUHWDLQLQJVXEVWUDWDRQWKHSKRWRV\QWKHWLFUHVSRQVHRIthree GURXJKW WROHUDQW SKRWRWURSKLF PLFURRUJDQLVPV LVRODWHG IURP D WHUUHVWULDO KDELWDW $UFK 0LFURELRO 'H :LQGHU % 3OXLV - 'H 5HXV / DQG 0XU /5 E &KDUDFWHUL]DWLRQ RI D F\DQREDFWHULDO DOJDO FUXVW LQ WKH FRDVWDO GXQHV RIWKH 1HWKHUODQGV ,Q &RKHQ < DQG 5RVHQEHUJ ( HGV 0LFURELDO 0DWV 3K\VLRORJLFDO (FRORJ\ RI %HQWKLF 0LFURELDO &RPPXQLWLHVSS$60:DVKLQJWRQ 'H:LQGHU%6WDO/-DQG0XU/5 Crinalium epipsammum VS QRY D ILODPHQWRXV F\DQREDFWHULXP ZLWK WULFKRPHV FRPSRVHG RI HOOLSWLFDO FHOOV DQG FRQWDLQLQJ SRO\EHWD JOXFDQFHOOXORVH -*HQ0LFURELRO 'H :LW 5 9DQ %RHNHO :+0 DQG 9DQ *HPHUGHQ + *URZWK RI WKH F\DQREDFWHULXP Microcoleus chthonoplastes RQ VXOILGH)(06 0LFURELRO(FRO 'H :LW 5 DQG 9DQ *HPHUGHQ + 2[LGDWLRQ RI VXOILGH WR WKLRVXOIDWH E\ Microcoleus chthonoplastes. )(06 0LFURELRO (FRO 'H :LW 5 DQG 9DQ *HPHUGHQ + ,QWHUDFWLRQV EHWZHHQ SKRWRWURSKLF EDFWHULD LQ VHGLPHQW HFRV\VWHPV +\GURELRO %XOO 'HFKR $: 0LFURELDO H[RSRO\PHU VHFUHWLRQV LQ RFHDQ HQYLURQPHQWV WKHLUUROHV LQ IRRG ZHEV DQG PDULQH SURFHVVHV Oceanogr 0DU%LRO$QQ5HY 'HFKR $: 0ROHFXODUVFDOH HYHQWV LQIOXHQFLQJ WKH PDFURVFDOH FRKHVLYHQHVV RI H[RSRO\PHUV ,Q .UXPEHLQ :( 3DWHUVRQ'0DQG6WDO/-HGV %LRVWDELOL]DWLRQRI6HGLPHQWVSS %,69HUODJ2OGHQEXUJ 'HFKR $: DQG 0RULDUW\ '-: %DFWHULDO H[RSRO\PHU XWLOL]DWLRQ E\ D KDUSDFWLFRLG FRSHSRG $ PHWKRGRORJ\ DQG UHVXOWV/LPQRO2FHDQRJU 'pIDUJH & 7ULFKHW- DQG &RXWH $ D 2Q WKH DSSHDUDQFH RI F\DQREDFWHULDO FDOFLILFDWLRQ LQ PRGHP VWURPDWROLWHV 6HG *HRO 'pIDUJH& 7ULFKHW- 0DXULQ$DQG+XFKHU0 E .RSDUDLQ 3RO\QHVLDQ DWROOV HDUO\ VWDJHV RI IRUPDWLRQ RI FDOFDUHRXV VWURPDWROLWHV 6HG*HRO 'HV 0DUDLV '- DQG &DQILHOG '( 7KH FDUERQ LVRWRSH ELRJHRFKHPLVWU\ RIPLFURELDOPDWV ,Q 6WDO /- DQG &DXPHWWH 3 HGV 0LFURELDO 0DWV 6WUXFWXUH 'HYHORSPHQW DQG (QYLURQPHQWDO 6LJQLILFDQFH SS 6SULQJHU 9HUODJ +HLGHOEHUJ 'HV0DUDLV'-'¶$PHOLR()DUPHU-'-¡UJHQVHQ%%3DOPLVDQR $& DQG 3LHUVRQ %. &DVH VWXG\ RI D PRGHP PLFURELDO PDWEXLOGLQJFRPPXQLW\ WKH VXEPHUJHG F\DQREDFWHULDOPDWV RI *XHUUHUR 1HJUR %DMD &DOLIRUQLD 6XU 0H[LFR ,Q 6FKRSI -: DQG .OHLQ & HGV 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS &DPEULGJH 8QLYHUVLW\ 3UHVV1HZ
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|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ynechococcus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scillatoria VSS XQGHU DXWRWURSKLF DQG SKRWRKHWHURWURSKLF FRQGLWLRQV-*HQ0LFURELRO *DUFLD3LFKHO) DQG %HERXW %0 3HQHWUDWLRQ RIXOWUDYLROHW UDGLDWLRQ LQWR VKDOORZ ZDWHU VHGLPHQWV +LJK H[SRVXUH IRU SKRWRV\QWKHWLFFRPPXQLWLHV0DU(FRO3URJ6HU *DUFLD3LFKHO ) DQG &DVWHQKRO] 5: &RPSDUDWLYH DQR[\JHQLF SKRWRV\QWKHWLF FDSDFLW\ LQ VWUDLQV RI D WKHUPRSKLOLFF\DQREDFWHULXP$UFK0LFURELRO
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icrocoleus chthonoplastes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lGHU '3 E 3KRWRVHQVRU\ EHKDYLRU LQ SURFDU\RWHV 0LFURELRO5HY +DGHU '3 6LJQDO SHUFHSWLRQ DQG DPSOLILFDWLRQ LQ SKRWRUHVSRQVHVRIF\DQREDFWHULD%LRSK\V&KHP +DOIHQ /1 DQG &DVWHQKRO] 5: *OLGLQJ PRWLOLW\ LQ WKH EOXHJUHHQDOJDOscillatoria princeps. -3K\FRO +DOOEHFN / DQG 3HGHUVHQ . $XWRWURSKLF DQG PL[RWURSKLF JURZWK of Gallionella ferruginea. - *HQ 0LFURELRO +HLMWKXLMVHQ -+)* DQG +DQVHQ 7$ %HWDLQH IHUPHQWDWLRQ DQGR[LGDWLRQE\ PDULQH Desulfuromonas VWUDLQV $SSO (QYLURQ 0LFURELRO +H\HU + 6WDO /- DQG .UXPEHLQ :( 6LPXOWDQHRXV KHWHURODFWLF DQG DFHWDWH IHUPHQWDWLRQ LQ WKH PDULQH F\DQREDFWHULXP Oscillatoria limosa LQFXEDWHG DQDHURELFDOO\ LQ WKHGDUN$UFK0LFURELRO +ROWRQ 5: DQG )UHHPDQ $: 6RPH WKHRUHWLFDO DQG H[SHULPHQWDO FRQVLGHUDWLRQV RI WKH JOLGLQJ PRYHPHQW RI EOXH JUHHQDOJDH $P-%RW +RXFKLQV-3 7KHSK\VLRORJ\DQGELRFKHPLVWU\RIK\GURJHQ PHWDEROLVP LQ F\DQREDFWHULD %LRFKLP %LRSK\V $FWD +RZDUG -% DQG 5HHV '& 1LWURJHQDVH D QXFOHRWLGH GHSHQGHQWPROHFXODUVZLWFK$QQ5HY%LRFKHP
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alothrix scopulorum - 0DU%LRO $VV 8. -¡UJHQVHQ %% $ WKLRVXOIDWH VKXQW LQ WKH VXOIXU F\FOH RI PDULQHVHGLPHQWV6FLHQFH -¡UJHQVHQ %% DQG &RKHQ < 6RODU /DNH 6LQDL 7KH VXOIXU F\FOH RI WKH EHQWKLF F\DQREDFWHULDO PDWV /LPQRO 2FHDQRJU -¡UJHQVHQ%%&RKHQ<DQG'HV0DUDLV'- 3KRWRV\QWKHWLF DFWLRQ VSHFWUD DQG DGDSWDWLRQ WR VSHFWUDO OLJKW GLVWULEXWLRQ LQ D EHQWKLF F\DQREDFWHULDO PDW $SSO (QYLURQ 0LFURELRO -¡UJHQVHQ%% &RKHQ<DQG5HYVEHFK13 7UDQVLWLRQIURP DQR[\JHQLF WR R[\JHQLF SKRWRV\QWKHVLV LQ D Microcoleus chthonoplastes F\DQREDFWHULDO PDW $SSO (QYLURQ 0LFURELRO -¡UJHQVHQ %% DQG 'HV 0DUDLV '- 2SWLFDO SURSHUWLHV RI EHQWKLF SKRWRV\QWKHWLF FRPPXQLWLHV )LEHURSWLF VWXGLHV RI F\DQREDFWHULDOPDWV/LPQRO2FHDQRJU -¡UJHQVHQ %% DQG 1HOVRQ '& %DFWHULDO ]RQDWLRQ SKRWRV\QWKHVLV DQG VSHFWUDO OLJKW GLVWULEXWLRQ LQ KRW VSULQJ PLFURELDOPDWVRI,FHODQG0LFURE(FRO -¡UJHQVHQ %% 5HYVEHFK 13 %ODFNEXUQ 7+ DQG&RKHQ < 'LXUQDO F\FOH RI R[\JHQ DQG VXOILGH PLFURJUDGLHQWV DQG PLFURELDOSKRWRV\QWKHVLVLQDF\DQREDFWHULDOPDWVHGLPHQW $SSO (QYLURQ 0LFURELRO -¡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
/-6WDO /HKPDQQ0 DQG:REHU* $FFXPXODWLRQ PRELOL]DWLRQ DQG WXPRYHU RI JO\FRJHQ LQ WKH EOXHJUHHQ EDFWHULXP Anacystis nidulans. $UFK0LFURELRO /RJDQ%: &U\SWR]RRQDQGDVVRFLDWHVWURPDWROLWHVIURPWKH UHFHQWRI6KDUN%D\:HVWHP$XVWUDOLD-*HRO /RUHQ] 0* DQG :DFNHUQDJHO : 1DWXUDO JHQHWLF WUDQVIRUPDWLRQRIPseudomonas sfutzen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nabaena cylindrica DQG Synechocystis - *HQ 0LFURELRO 0DFNHUUDV $+
&KDSWHU &\DQREDFWHULDO 0DWVDQG 6WURPDWROLWHV 0LOOLPDQ -' 3URGXFWLRQ DQG DFFXPXODWLRQ RI FDOFLXP FDUERQDWH LQ WKH RFHDQ EXGJHW RI D QRQVWHDG\ VWDWH *OREDO %LRJHRFKHP&\FOHV 0L]LRUNR +0 DQG /RULPHU *+ 5LEXORVHELSKRVSKDWH FDUER[\ODVHR[\JHQDVH$QQ5HY%LRFKHP 0RH]HODDU 5 %LMYDQN 60 DQG 6WDO /- )HUPHQWDWLRQ DQG VXOIXU UHGXFWLRQ LQ WKH PDWEXLOGLQJ F\DQREDFWHULXP 0LFURFROHXV chthonoplastes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yngbya DHVWXDULL $SSO (QYLURQ 0LFURELRO 3DOLQVND .$ /LHVDFN : 5KLHO ( DQG .UXPEHLQ :( 3KHQRW\SH YDULDELOLW\ RI LGHQWLFDO JHQRW\SHV WKH QHHG IRU D
FRPELQHG DSSURDFK LQ F\DQREDFWHULDO WD[RQRP\ GHPRQVWUDWHG RQ0HULVPRSHGLDOLNHLVRODWHV$UFK0LFURELRO 3DQNUDW]+6 DQG%RZHQ&& &\WRORJ\RIEOXHJUHHQDOJDH ,7KHFHOOVRISymploca muscorum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lyngbyaceus *RPRQW - *HQ 0LFURELRO 3HQWHFRVW $ *URZWK DQG FDOFLILFDWLRQ RI WKH F\DQREDFWHULXP Homeothrix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
%LRJHRFKHPLVWU\ DQG *HRPLFURELRORJ\ 0HWKRGV0HWDOVDQG $VVHVVPHQWSS $QQ $UERU 6FLHQFH $QQ $UERU 3RWWV 0 2FDPSR)ULHGPDQQ 5 %RZPDQ 0$ DQG 7R]XQ % Chroococcus 6 DQG Chroococcus 1 F\DQREDFWHULD PRUSKRORJLFDO ELRFKHPLFDO DQG JHQHWLF FKDUDFWHUL]DWLRQ DQG HIIHFWV RI ZDWHU VWUHVV RQ XOWUD VWUXFWXUH $UFK 0LFURELRO 3ULQJVKHLP (* &\DQRSK\FHHQ6WXGLHQ $UFK 0LNURELRO 5DEHQVWHLQ $ 5HWKPHLHU - DQG )LVFKHU 8 6XOSKLWH DV LQWHUPHGLDWH VXOSKXU FRPSRXQG LQDQDHURELFVXOSKLGHR[LGDWLRQ WR WKLRVXOSKDWHE\ PDULQH F\DQREDFWHULD=HLWVFKU1DWXUIRUVFK& 5DPVLQJ 1% DQG3UXIHUW%HERXW/ 0RWLOLW\RIMicrocoleus chthonoplastes VXEMHFWHG WR GLIIHUHQWOLJKWLQWHQVLWLHVTXDQWLILHG E\ GLJLWDO LPDJH DQDO\VLV ,Q 6WDO /- DQG &DXPHWWH 3 HGV 0LFURELDO 0DWV 6WUXFWXUH 'HYHORSPHQW DQG (QYLURQPHQWDO 6LJQLILFDQFH SS 6SULQJHU9HUODJ+HLGHOEHUJ 5HHG 5+ %RURZLW]ND /- 0DFND\ 0$ &KXGHN -$ )RVWHU 5 :DUU 6&5 0RRUH '- DQG 6WHZDUW:'3D 2UJDQLFVROXWH DFFXPXODWLRQ LQ RVPRWLFDOO\ VWUHVVHG F\DQREDFWHULD )(06 0LFURELRO 5HY 5HHG 5+ DQG 6WHZDUW :'3 3K\VLRORJLFDO UHVSRQVHV RI Rivularia atra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nabaena cylindrica. 3K\VLRO 3ODQW 5HQVWURP.HOOQHU(DQG %HUJPDQ% *O\FRODWHPHWDEROLVP LQF\DQREDFWHULD ,9 8SWDNH JURZWK DQG PHWDEROLFSDWKZD\V 3K\VLRO 3ODQW 5HYVEHFK 13 -¡UJHQVHQ%%%ODFNEXUQ7+ DQG&RKHQ< 0LFURHOHFWURGH VWXGLHV RI WKH SKRWRV\QWKHVLV DQG 2 +6 DQG S+ SURILOHV RI D PLFURELDO PDW /LPQRO 2FHDQRJU 5LFKDUGVRQ // DQG &DVWHQKRO] 5: D 'LHO YHUWLFDO PRYHPHQWV RI WKHF\DQREDFWHULXPOscillaroria terebrifomis LQ DVXOILGHULFK KRWVSULQJPLFURELDOPDW$SSO(QYLURQ0LFURELRO 5LFKDUGVRQ // DQG &DVWHQKRO] 5: E (QKDQFHGVXUYLYDORI WKHF\DQREDFWHULXPOscillatoria terebriformis LQGDUNQHVVXQGHU DQDHURELF FRQGLWLRQV $SSO(QYLURQ0LFURELRO 5LFKDUGVRQ // DQG&DVWHQKRO]5: &KHPRNLQHWLFPRWLOLW\ UHVSRQVHV RI WKH F\DQREDFWHULXP Oscillatoria terebriformis. $SSO (QYLURQ 0LFURELRO 5LSSND5'HUXHOOHV- :DWHUEXU\ -% +HUGPDQ0DQG6WDQLHU5< *HQHULF DVVLJQPHQWVVWUDLQ KLVWRULHV DQG SURSHUWLHV RI SXUHFXOWXUHVRIF\DQREDFWHULD - *HQ 0LFURELRO
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’Leary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hloroflexusOLNH EDFWHULD LQ KRW VSULQJ PLFURELDO PDWV $SS (QYLURQ 0LFURELRO 6FKDX / %HQVRQ $$ %DVVKDP -$DQG&DOYLQ 0 7KHSDWK RI FDUERQ LQ SKRWRV\QWKHVLV ;, 7KH UROH RI JO\FROLF DFLG 3K\VLRO 3ODQW 6FKDXE %(0 DQG 9DQ *HPHUGHQ + 6XOIXU EDFWHULD LQ VHGLPHQWV RI WZR FRDVWDO HFRV\VWHPV WKH %DVVLQ G $UFDFKRQ DQG WKH(WDQJGX3UHYRVW)UDQFH+\GURELRORJLD 6FKPHWWHUHU * &\DQREDFWHULDO UHVSLUDWLRQ ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU$FDGHPLF3XEOLVKHUV'RUGUHFKW 6FKRSI -: HG (DUWK V(DUOLHVW%LRVSKHUH,WV2ULJLQDQG (YROXWLRQ SS 3ULQFHWRQ 8QLYHUVLW\ 3UHVV 3ULQFHWRQ 1HZ -HUVH\ 6FKRSI -: DQG .OHLQ & 7KH 3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS&DPEULGJH8QLYHUVLW\ 3UHVV 1HZ
&KDSWHU &\DQREDFWHULDO 0DWV DQG 6WURPDWROLWHV 6WDO /- 0LNURELHOOH0DWWHQ ,Q0H\HU5HLO/$DQG.|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scillatoria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hromatium vinosum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
7DOERW00% %DWH*& DQG &DPSEHOO (( $ UHYLHZ RIWKH HFRORJ\ RI VXUI]RQH GLDWRPV ZLWK VSHFLDO UHIHUHQFH WR Anulus australis. 2FHDQRJU0DU%LRO$QQX5HY 7HDVH % -LLUJHQV 8- *ROHFNL -5 +HLQULFK 85 5LSSND 5 DQG :HFNHVVHU - +QH6WUXFWXUDO DQG &KHPLFDO $QDO\VHV RQ ,QQHU DQG 2XWHU 6KHDWK RI WKH &\DQREDFWHULXP Gloeothece VS 3&&$QWRQLH9DQ/HHXZHQKRHN,QW-*HQ0RO0LFURELRO 7USHU +* DQG *DOLQVNL ($ &RPSDWLEOH VROXWHV LQ KDORSKLOLFSKRWRWURSKLFSURFDU\RWHV ,Q &RKHQ<DQG5RVHQEHUJ ( HGV 0LFURELDO 0DWV 3K\VLRORJLFDO (FRORJ\ RI %HQWKLF 0LFURELDO&RPPXQLWLHVSS $60:DVKLQJWRQ 7XUQHU 60 0DOLQ * /LVV 36 +DUERXU '6 DQG +ROOLJDQ 30 7KH VHDVRQDO YDULDWLRQ RI GLPHWK\O VXOILGH DQG GLPHWK\OVXOIRQLRSURSLRQDWH FRQFHQWUDWLRQV LQ QHDUVKRUH ZDWHUV /LPQRO2FHDQRJU 7XUSLQ '+ 0LOOHU $* DQG &DQYLQ '7 &DUER[\VRPH FRQWHQWRISynechococcus leopoliensis &\DQRSK\WD LQUHVSRQVH WRLQRUJDQLF FDUERQ -3K\FRO 8WNLOHQ +& 7KLRVXOSKDWH DV HOHFWURQ GRQRU LQ WKH EOXH JUHHQDOJDAnacystis nidulans. -*HQ0LFURELRO 9DQ %HUJHLMN 6$ DQG 6WDO /- 7KH UROH RI R[\JHQLF SKRWRWURSKLF PLFURRUJDQLVPV LQ SURGXFWLRQ DQG FRQYHUVLRQ RI GLPHWK\OVXOIRQLRSURSLRQDWH DQG GLPHWK\OVXOILGH LQ PLFURELDO PDWV,Q.LHQH539LVVFKHU37.HOOHU0'DQG.LUVW*2HGV %LRORJLFDO DQG (QYLURQPHQWDO &KHPLVWU\ RI'063 DQG 5HODWHG 6XOIRQLXP&RPSRXQGVSS 3OHQXP3UHVV1HZ
:DOWHU05%XLFN5DQG'XQORS-65 6WURPDWROLWHV 0\U ROG IURP WKH 1RUWK 3ROH DUHD :HVWHUQ $XVWUDOLD 1DWXUH :DOWHU 05 *URW]LQJHU -3 DQG 6FKRSI -: 3URWHUR]RLF VWURPDWROLWHV ,Q 6FKRSI-: DQG .OHLQ &HGV 7KH3URWHUR]RLF %LRVSKHUH $ 0XOWLGLVFLSOLQDU\ 6WXG\ SS &DPEULGJH 8QLYHUVLW\3UHVV1HZ
/-6WDO
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
6XPPDU\ &\DQREDFWHULD DUH D WD[RQRPLFDOO\ GLYHUVH SURGXFWLYH DQG ELRJHRFKHPLFDOO\ LPSRUWDQW FRPSRQHQW RI RFHDQLF FRDVWDO DQG HVWXDULQH SK\WRSODQNWRQ FRPPXQLWLHV 5HFHQW PROHFXODU DQG DQDO\WLFDO FKDUDFWHUL]DWLRQ RI WKHVH FRPPXQLWLHV WRJHWKHU ZLWK D JURZLQJ QXPEHU RI FUXLVHEDVHG DQG UHPRWHVHQVLQJ VXUYH\V RIWKH ZRUOG¶V RFHDQ VXUIDFH ZDWHUV LQGLFDWH WKDW F\DQREDFWHULDSOD\ D KLJKO\ VLJQLILFDQWUROH LQ PDULQH FDUERQ DQG QLWURJHQ F\FOLQJ 9DVW VHJPHQWV RI WKH ZRUOG¶V RFHDQ VXUIDFH ZDWHUV DUH QLWURJHQ GHSOHWH DQG SODQNWRQLF 1IL[LQJ F\DQREDFWHULD FDQ FRPSHWH HIIHFWLYHO\ LQ WKLV ODUJH QLFKH 6XUIDFH EORRPV RI WKH ILODPHQWRXV GLD]RWURSKLF JHQHUD Trichodesmium DQG Richelia HQGRV\PELRQW RI WKHGLDWRP Rhizosolenia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ynechococcus, Synechocystis DQG Prochlorococcus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hitton and M. Potts (eds), The Ecology of Cyanobacteria, © Kluwer Academic Publishers. Printed in the Netherlands.
+ : 3DHUO
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ynechococcus, Prochlorococcus DQG Synechocystis. 7KHVH IRUPV DUH UHVSRQVLEOH IRU D ODUJH IUDFWLRQ WR ! RIWKH
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richodesmium; &DUSHQWHU D E Nodularia, Aphanizomenon; +XEHU .RQRQHQ HW DO 'LD]RWURSKLF F\DQREDFWHULD SOD\ NH\ UROHV LQ PXWXDOLVWLF DQG V\PELRWLF DVVRFLDWLRQV ZLWK PLFURDOJDH HJ Rhizosolenia-Richelia: 9LOODUHDO PDFURDOJDH HJ ZLWK WKH VLSKRQRXV JUHHQ VHDZHHG Codium DQG WKH SODQNWRQLF SKDHRSK\WH Sargassum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
&KDSWHU 0DULQH3ODQNWRQ
IRU WKLV WD[RQRPLFDOO\ SK\VLRORJLFDOO\ HFRORJLFDOO\GLYHUVHJURXSRIGLD]RWURSKV
DQG
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richodesmium, "tricho" EHLQJ GHULYHG IURP WKH *UHHN IRU KDLU DQG "desmos" *UHHN IRU FKDLQ )RJJ 3ODWHV D E ,QKLVDFFRXQWRI WKH YR\DJHV RI WKH %HDJOH 'DUZLQ SURYLGHG GHWDLOHG GHVFULSWLRQV RI ODUJH DQG ZLGHO\GLVWULEXWHG EORRPV RI Trichodesmium LQ EHFDOPHG SHODJLF DQG QHDUVKRUH VRXWKZHVWHUQ $WODQWLF 2FHDQ ZDWHUV RII 6RXWK $PHULFD 6LPLODU UHSRUWV RI VHD VDZGXVW DWWULEXWHG WR VXUIDFH EORRPV RI Trichodesmium, ZHUH
Table 1. 0RUSKRORJLFDO JURXSV DQG UHVSUHVHQWDWLYH JHQHUD RI PDULQH SODQNWRQLF F\DQREDFWHULD FKDUDFWHULVWLFVRIJHQHUDDUHOLVWHG
*52836
8QLFHOOXODU VROLWDU\
*(1(5$
Synechococcus Prochlorococcus Synechocystis Aphanothece
FRORQLDO
Merismopedia
1 ),;$7,21
1IL[LQJ HFRORJLFDO DQG WURSKLF
&+$5$&7(5,67,&6
0RVW
6ROLWDU\ FRPPRQ LQ ROLJRWURSKLFRFHDQVIRUP GHHS FKORURSK\OO PD[LPD FDQIRUPEORRPV LQ K\SHUVDOLQH ZDWHUV 6ROLWDU\ EORRP IRUPLQJ LQ K\SHUVDOLQH ODJRRQVED\V 6RPH 6ROLWDU\SLFRSODQNWRQLFROLJRKDOLQH " 3UHIHUV QXWULHQW HQULFKHG ZDWHUV
)LODPHQWRXV QRQKHWHURF\VWRXV
KHWHURF\VWRXV
Lyngbya Oscillatoria
6RPH 6RPH
$JJUHJDWHV RIWHQ DVVRFLDWHG ZLWK SURWLVWV PHWD]RDQV $ IHZ EORRPV UHSRUWHG VRPH DVVRFLDWHG ZLWK SODQNWRQLF PDFURDOJDHDQGKLJKHUSODQWV
Phormidium Spirulina Trichodesmium
6ROLWDU\ DQG DVVRFLDWHG ZLWK RWKHU SURWLVWV 6ROLWDU\K\SHUVDOLQHDQG QXWULHQW HQULFKHGZDWHUV $JJUHJDUHVIRUPEORRPV LQ WURSLFDO DQG VXEWURSLFDO ZDWHUV DVVRFLDWHG ZLWK SURWLVWVLQYHUWHEUDWHJUD]HUV
Anabaena Aphanizomenon Nodularia Richelia
6ROLWDU\ LQ ODJRRQV ED\V $JJUHJDWHGEORRPIRUPHU LQ QXWULHQW HQULFKHG ZDWHUV %ORRP IRUPHU LQ FRDVWDOQXWULHQWHQULFKHGZDWHUV (QGRV\PELRQW LQ WKHGLDWRPV Rhizosolenia DQG Hemiaulus LQ WURSLFDO DQG VXEWURSLFDOZDWHUV
+ : 3DHUO
D
E
F )LJ 1. ([DPSOHVRI SODQNWRQLFF\DQREDFWHULDYLHZHGZLWKOLJKWPLFURVFRS\EULJKWILHOGDQGSKDVHFRQWUDVW 7XIWVKDSHDJJUHJDWHV RI WKH QRQKHWHURF\VWRXV1IL[LQJRFHDQLFEORRPIRUPHUTrichodesmium thiebautii DUHVKRZQLQ3ODWHE D Synechocystis, DFRPPRQFRFFRLGHVWXDULQHFRDVWDOIRUPIRXQGDVVROLWDU\FHOOV RUDJJUHJDWHV DVVKRZQ 6DPSOHIURP%RJXH6RXQG 1RUWK&DUROLQDDIXOOVDOLQLW\SSW ROLJRWURSKLFFRDVWDOVRXQG E $JJUHJDWHGTrichodesmium thiebautii WULFKRPHVIURPEORRPLQWKH*XOI6WUHDPZHVWHUQ1RUWK $WODQWLF2FHDQ DSSUR[LPDWHO\ NP VRXWKHDVWRI%HDXIRUW1RUWK&DUROLQD c +HWHURF\VWRXVFRDVWDOHVWXDULQHEORRPIRUPHUNodularia: QRWHWKLFNZDOOHGKHWHURF\VWVDQGEDFWHULDOHSLSK\WHV
&KDSWHU 0DULQH3ODQNWRQ ORJJHG E\ &DSWDLQ - &RRN DQG KLV QDWXUDOLVW FROOHDJXH - %DQNV GXULQJ FUXLVHV RI +06 (QGHDYRXU LQ WKH 6RXWK 3DFLILF GXULQJ %HDJOHKROH 0DQ\ RI WKHVH REVHUYDWLRQV GRFXPHQW RFFXUUHQFHV LQ ZDWHUV ZHOODZD\ IURP ODQG PDVVHV DQG UHIOHFW WKH SUHVHQFH RI F\DQREDFWHULDO EORRPV LQ ROLJRWURSKLF RFHDQ UHJLRQV :KLOH QRW UHDOL]HG DW WKH WLPH LW LV QRZ ZHOO HVWDEOLVKHG WKDW Trichodesmium DQG RWKHU IUHTXHQWO\ RFFXUULQJ KHWHURF\VWRXV F\DQREDFWHULD LQ EORRPV VXFK DV Richelia, RU DV DQ HQGRV\PELRQW LQ WKH SHODJLF GLDWRPV Rhizosolenia DQG Hemiaulus, IL[ QLWURJHQ DQG DUH VLJQLILFDQWFRQWULEXWRUV WR RFHDQLF 1 EXGJHWV &DUSHQWHU E 0DJXH HW DO 0DUWLQH] HW DO &DUSHQWHU DQG 5RPDQV ,Q FRDVWDO ZDWHUVWKDWXQGHUJRHXWURSKLFDWLRQ VXFK DV WKH %DOWLF WKH KHWHURF\VWRXV GLD]RWURSKLF JHQHUD Nodularia DQG Aphanizomenon FDQ FRQWULEXWH DV PXFK DV RI WKHELRORJLFDOO\DYDLODEOH QHZ 1 6¡UHQVVRQ DQG 6DKOVWHQ :XOII ,Q DGGLWLRQ WR WKHLU ELRJHRFKHPLFDO VLJQLILFDQFH PDMRU EORRP WD[D VXFK DV VXUIDFHGZHOOLQJ Nodularia EORRPV LQ WKH %DOWLF 6HD FDQ DOWHU VSHFWUDO TXDOLWLHV LQFUHDVH OLJKW DEVRUSWLRQ DQG WKXV FRQWULEXWH WR WKH KHDWLQJ RI VXUIDFH ZDWHUV .DKUX HW DO 7KLFNZDOOHG SRRUO\SLJPHQWHG KHWHURF\VWV DUH D GLDJQRVWLF IHDWXUH RI WKH ILODPHQWRXV KHWHURF\VWRXV )LJ JHQHUD OLVWHG LQ 7DEOH +HWHURF\VWV DUH GLIIHUHQWLDWHG 2 GHSOHWH FHOOV ZKLFK KDUERU WKH 2 ODELOH 1IL[LQJ HQ]\PH FRPSOH[ RI QLWURJHQDVH 'RQ]H HW DO :RON +HWHURF\VWVRIIHUD FOHDU DGYDQWDJH LQ WKH DHURELF 1 GHSOHWH VXUIDFH ZDWHUV ZKLFK FKDUDFWHUL]H PXFK RIWKH ZRUOG V RFHDQV ,Q DGGLWLRQ KHWHURF\VWV RIWHQ VHOHFWLYHO\ DWWUDFW KHWHURWURSKLF 2UHVSLULQJ EDFWHULDO HSLSK\WHV ZKLFK IDFLOLWDWH UHPRYDO RI 2 LQ WKHLU YLFLQLW\ 3DHUO DQG .HOODU 2QH SDUWLFXODUO\ VWULNLQJ DVSHFW RI PDULQH SODQNWRQLF KHWHURF\VWRXV F\DQREDFWHULD LV WKHLU FRQILQHPHQW WR VSHFLILF JHRJUDSKLF UHJLRQV )UHH IORDWLQJ JHQHUD VXFK DV Nodularia DQG Aphanizomenon FDQ EH IRXQG RIWHQ DV EORRPV LQ FHUWDLQHVWXDULQH DQG FRDVWDO HQYLURQPHQWV LQFOXGLQJ SHULRGLFDOO\VWUDWLILHG UHJLRQV RI WKH QXWULHQW HQULFKHG %DOWLF 6HD DQG SK\VLFDOO\ LVRODWHG HVWXDULQH DQG ODJRRQ V\VWHPV ZKHUH WKHUH LV OLPLWHG H[FKDQJH ZLWK WKH RFHDQ .RQRQHQ HW DO 7KH EHVW GRFXPHQWHG H[DPSOH LV WKH VWUDWLILHG K\SHUVDOLQH 3HHO+DUYH\ HVWXDU\ LQ :HVWHUQ $XVWUDOLD +XEHU 6RPH FRPPRQ IHDWXUHVHPHUJHZKHQ RQHFRQVLGHUV WKH UHDVRQV IRU WKLV UDWKHU H[FOXVLYH FRQILQHPHQW RI
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ichelia ZKLFK LQKDELWV WKH ODUJH SHQQDWH GLDWRPV Rhizosolenia DQG Hemiaulus LQ WURSLFDO DQG VXEWURSLFDO 1 GHSOHWH ROLJRWURSKLF ZDWHUV 7D\ORU 9LOODUHDO 5HODWLYHO\ KLJK UDWHV RI 1 IL[DWLRQDUHDWWULEXWHGWR WKHVH DVVRFLDWLRQV 0DJXH HW DO 0DUWLQH] HW DO HVSHFLDOO\ LQ ODUJH VXUIDFH PDWV JURZLQJ LQ FDOP VHDV 'XULQJ VXFK WLPHV WKH Richelia-Rhizosolenia V\PELRVLV PD\ FRH[LVW ZLWK WKH QRQKHWHURF\VWRXV 1 IL[HU Trichodesmium 9LOODUHDO /RZ OHYHOV RI WXUEXOHQFH VPDOOVFDOH VKHDU DQG VWUHVV ODUJH VFDOH YHUWLFDO PL[LQJ IDYRU VXUIDFH DFFXPXODWLRQV RI DFWLYHO\ JURZLQJ EORRPV RI ERWK DVVHPEODJHV &DUSHQWHU 3DHUO HW DO 7KH ZD\V LQ ZKLFK 1 IL[DWLRQ LV FDUULHG RXW E\ WKHVH PRUSKRORJLFDOO\GLVWLQFW GLD]RWURSKV PD\ EH TXLWH GLIIHUHQW KRZHYHU DQG DUHWKH VXEMHFW RI FRQVLGHUDEOH UHVHDUFK 7KH IDFW WKDW Richelia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ichelia WRRN UHIXJH LQ WKH PLFUR HQYLURQPHQW SURYLGHG E\ WKH ODUJH GLDWRP Rhizosolenia. Rhizosolenia EHQHILWV IURP KDYLQJ DFFHVVWRELRORJLFDOO\ DYDLODEOH1LQZKDWDUH VRPH RI WKH PRVW1GHILFLHQW ZDWHUV RQHDUWK ,Q DGGLWLRQ WKH KRVW GLDWRPSURYLGHV EXR\DQF\ DQG WKHUHE\ PDLQWDLQV WKH HQGRV\PELRQW LQ WKH LOOXPLQDWHG HXSKRWLF ]RQH
+ : 3DHUO
ZKHUH UDGLDQW HQHUJ\ FDQ VXSSRUW LWV QLWURJHQIL[LQJ DFWLYLWLHV ,QWHUHVWLQJO\ WKH RWKHU LQKDELWDQWRIWKHVH QXWULHQW GHSOHWH ZDWHUV Trichodesmium, DSSHDUV WR HPSOR\ D YHU\GLIIHUHQW\HWHIIHFWLYHVWUDWHJ\IRUPHHWLQJLWV1 DQG&UHTXLUHPHQWV 5DWKHUWKDQIRUPLQJKHWHURF\VWV WKLVXQGLIIHUHQWLDWHGGLD]RWURSKIL[HV&2 DQG1DQG HYROYHV 2 VLPXOWDQHRXVO\GXULQJ GD\OLJKW DIHDWQRW VKDUHG ZLWK RWKHU QRQKHWHURF\VWRXV ILODPHQWRXV GLD]RWURSKLF F\DQREDFWHULD VXFK DV Lyngbya DQG Oscillatoria. 7KH ODWWHUVWUDLQVIL[PRVWRIWKHLU1 DW QLJKWZKHQ 2 HYROXWLRQ FHDVHV 6WDO DQG .UXPEHLQ *DOORQ DQG 6WDO 6HYHUDO PHFKDQLVPV ZHUH SURSRVHG WR DFFRXQW IRU WKLV XQLTXH DELOLW\ RI Trichodesmium. 0RVW PHFKDQLVPV LQYROYH WKH QHHG WR H[FOXGH 2 IURP QLWURJHQDVH RQ HLWKHU LQWUD RU LQWHUFHOOXODUOHYHOV 7KHVHPHFKDQLVPVDUHGLVFXVVHG LQ6HFWLRQ,,,&
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
Fig. 2. 3LFRSODQNWRQLF P F\DQREDFWHULD FRQFHQWUDWHG E\ PLFURILOWUDWLRQIURPWKH6DUJDVVR6HD 7KHFHOOVZHUHLQFXEDWHGIRU K ZLWK WKH WHWUD]ROLXP VDOW LRGRQLWURWHWUD]ROLXP ,17 GDUN FHOOXODU LQFOXVLRQV DUH FRORUHG IRUPD]DQ FU\VWDOV WKDW IRUP XSRQ UHGXFWLRQ ,17LVFRPPRQO\XVHGDVDQLQGLFDWRURIPHWDEROLFDOO\ DFWLYHLHJURZLQJ FHOOV
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ynechococcus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
Table 2. $FKURQRORJ\RIPDMRUGLVFRYHULHVWHFKQLFDODQGFRQFHSWXDOFRQWULEXWLRQVWRWKHXQGHUVWDQGLQJRIWKHUROHV F\DQREDFWHULDSOD\LQ PDULQHSODQNWRQLFFRPPXQLW\VWUXFWXUHDQGIXQFWLRQ
'$7(
&2175,%87,21
5()(5(1&(
PLG WRODWH V
)LODPHQWRXV EORRPIRUPLQJ WD[D GHVFULEHG WKURXJKRXW WHPSHUDWHWURSLFDODQGVXEWURSLFDORFHDQV
'DUZLQ .DUVWHQV 2VWHQIHOG 6FKPLGW
HDUO\ V V
1DQRSODQNWRQGLVFRYHUHGLQWKHVHD 1 IL[DWLRQDVVRFLDWHGZLWKILODPHQWRXVQRQKHWHURF\VWRXV DQGKHWHURF\VWRXVEORRPIRUPLQJWD[D
/RKPDQ 6RXUQLD 'XJGDOHHWDO *RHULQJHWDO
ODWH V
$FHW\OHQHUHGXFWLRQDQG 1 DVVLPLODWLRQ DVVD\V HQDEOH ODUJHVFDOHDVVHVVPHQWVRIRFHDQLF1 IL[DWLRQLWLV VLJQLILFDQWLQSODFHV $SSOLFDWLRQ RIIOXRUHVFHQFHPLFURVFRS\LQELRORJLFDO RFHDQRJUDSK\
7D\ORUHWDO 0DJXHHWDO &DUSHQWHU 0DUX\DPD
ODWH V
3LFRSODQNWRQLFWD[DIRXQGWREHXELTXLWRXV DQGDEXQGDQW
HDUO\ V
8VHRIIORZF\WRPHWU\FRXSOHGWR IOXRUHVFHQFHGHWHFWLRQ DOORZV VSDWLDOFKDUDFWHULVDWLRQDQGTXDQWLWDWLRQRI SLFRSODQNWRQLF EDFWHULD 0ROHFXODUFKDUDFWHULVDWLRQRIWD[DDQGIXQFWLRQDO JHQHV IDFLOLWDWHVFRPPXQLW\VWUXFWXUDODQGIXQFWLRQDODQDO\VLV
-RKQVRQ 6LHEXUWK :DWHUEXU\ HWDO 2OVRQHWDO &KLVKROPHWDO
V
ODWHV HDUO\ V
*LRYDQQRQLHWDO :\PDQHWDO =HKUHWDO
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nabaena, Aphanizomenon, Microcystis, Nodularia DQG Oscillatoria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icrocystis), FRDVWDO ODJRRQV DQG HPED\PHQWV LQ &RORPELD Anabaena, Aphanizomenon DQG VKDOORZ FRDVWDO ODJRRQV ZKLFK GUDLQ DUDEOH ODQGV LQ 6SDLQ 3RUWXJDO DQG 1HZ =HDODQG Anabaena, Microcystis, Nodularia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
&KDSWHU 0DULQH3ODQNWRQ
Table 3. 3K\VLDOFKHPLFDO IDFWRUV SRWHQWLDOO\ FRQWUROOLQJ F\DQREDFWHULDO EORRPV
)$&7256 Physical )OXVKLQJ$OWHUHG ZDWHU UHVLGHQFH WLPH
,03$&76 3RWHQWLDOUHPRYDOPHFKDQLVPIRUEORRPVLIIOXVKLQJH[FHHGVJURZWKUDWHVRI EORRPWD[D
/DUJH VFDOH YHUWLFDO PL[LQJ
&RXQWHUDFWVQHDUVXUIDFHDFFXPXODWLRQVRIEXR\DQWEORRPSRSXODWLRQV 0D\ HQKDQFHFRPSHWLWLRQIRUOLJKWDQGQXWULHQWVZLWKQRQERX\DQWHXNDU\RWLFWD[D
6PDOO VFDOH WXUEXOHQFH VKHDU
0D\GLVUXSWILODPHQWVFRORQLHVDJJUHJDWHVDQGDVVRFLDWLRQVZLWKRWKHU PLFURIORUDDQGPLFURPDFURIDXQD
6KDGLQJ UHGXFHG VXUIDFH LUUDGLDQFH
&DQDOWHUSK\WRSODQNWRQFRPPXQLW\FRPSRVLWLRQDQGFDQQHJDWLYHO\DIIHFW F\DQREDFWHULDOVXUIDFHEORRPWD[D
7HPSHUDWXUH
*HQHUDOO\WHPSHUDWXUHVLQH[FHVVRI&DFFRPSDQLHGE\VWUDWLILFDWLRQDQG KLJKQXWULHQWORDGLQJFDQSURPRWHFHUWDLQEORRPV
Chemical 1XWULHQW1DQG3 LQSXWV
/RQJWHUPPRQWKV\HDUV UHGXFWLRQVLQERWK1DQG3LQSXWVLQHVWXDULHVPD\ UHGXFHF\DQREDFWHULDOEORRPSRWHQWLDOV /RZ1WR3ORDGLQJUDWLRV DFRPSDQLHGE\KLJK3PD\HQKDQFHEORRPSRWHQWLDOVRIGLD]RWURSKLFWD[D
6DOLQLW\
6DOLQLW\LQH[FHVVRIDIHZSSWDV1D&O FDQEHDQHIIHFWLYHEDUULHUWR GHYHORSPHQWDQGSHUVLVWHQFHRIIUHVKZDWHUQXLVDQFHVSHFLHV 2WKHUVSHFLHV WROHUDWHVDOLQLW\DQGK\SHUVDOLQLW\
7UDFH PHWDOV
8QGHUKLJK1DQG3ORDGLQJFRQGLWLRQVRUZKHQ1IL[LQJWD[DSUHYDLO UHVWULFWHGDYDLODELOLW\RI)HPD\FRQWUROJURZWK 6RPHF\DQREDFWHULDDUHDEOH WRFRPSHWHIRUORZOHYHOVRI)H 1RFRQYLQFLQJHYLGHQFHIRURWKHUWUDFHPHWDO OLPLWDWLRQV
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
FRQVLGHUDWLRQ RI VDOLQLW\ DV D IDFWRU ZKLFK UHJXODWHV HVWXDULQH 1 IL[DWLRQ ,I DOORFKWKRQRXV SRSXODWLRQV ZHUH UHVSRQVLEOH IRU VHHGLQJ GLD]RWURSKV LQ HVWXDULHV VDOLQLW\ PLJKW UHSUHVHQW D FRQVLGHUDEOH EDUULHUWRWKHLUHVWDEOLVKPHQW :LWK UHVSHFW WR LRQLF VWUHQJWK FHUWDLQ EORRPIRUPLQJ F\DQREDFWHULD PD\ EH KLJKO\ VHQVLWLYH WR D VOLJKWFKDQJHLQ VDOLQLW\ ZKHQ WKH\ HQWHUHVWXDULQH ZDWHUV 3DHUO HW DO ,Q DGGLWLRQ +RZDUWK DQG &ROH DQG +RZDUWK HW DO SURSRVHG WKDW UHODWLYHO\ KLJK FRQFHQWUDWLRQV RI VXOIDWH 62 D VWUXFWXUDO DQDORJXH RI PRO\EGDWH 0R2 DQG SUHVHQW LQ VHDZDWHU DQG VDOLQH ODNHV PLJKW FRPSHWLWLYHO\ LQKLELW 1 IL[DWLRQ :KLOH FRPSHWLWLYH LQKLELWLRQ RI 0R2 XSWDNH E\ WKH 62 FRQFHQWUDWLRQV SUHVHQW LQ VHDZDWHU LV RSHUDWLYH &ROH HW DO WKH 1 IL[LQJ SRWHQWLDOV RI PDULQH GLD]RWURSKV DSSHDU XQDIIHFWHG E\ WKLV FRPSHWLWLRQ 3DXOVHQ HW DO 0RVW OLNHO\ WKH VPDOO FHOOXODU 0R UHTXLUHPHQWV IRU 1 IL[DWLRQ DUH PHW WKRXJK UHGXFHGEXWVXIILFLHQWXSWDNHDQG VWRUDJH (VWXDULQH DQG RFHDQLF ZDWHUV DUH a priori QRW EDUULHUV WR WKH HVWDEOLVKPHQW DQG SUROLIHUDWLRQ RI F\DQREDFWHULDO GLD]RWURSKV 1LWURJHQ GHILFLHQW HVWXDULHV DQG FRDVWDO ODJRRQ HQYLURQPHQWV ZRUOGZLGH VXSSRUW EHQWKLF QRQ KHWHURF\VWRXV DQG KHWHURF\VWRXV WD[D WKDW VKRZ UHODWLYHO\ KLJK 1 IL[DWLRQ UDWHV 2FHDQLF FRUDO UHHI VKHOI DQG FRDVWDO PDQJURYH KDELWDWV DUH RIWHQ ULFKO\ HQGRZHG ZLWK GLYHUVH QRQ KHWHURF\VWRXV HJ Oscillatoria, Lyngbya, Microcoleus) DQG KHWHURF\VWRXV HJ Scytonema, Nostoc, Anabaena, Cylindrospermum, Calothrix) JHQHUD )UpP\ 3RWWV :KLWWRQDQG 3RWWV 3RWWV DQG :KLWWRQ D E ,Q DGGLWLRQ QXPHURXV GLD]RWURSKLF EDFWHULD KDYH EHHQ LVRODWHG IURP PDULQH VHGLPHQWV VXVSHQGHG DJJUHJDWHV DQG VXVSHQGHG SODQWV DQG DQLPDOV 3DHUO 7KLV PDNHV LW GLIILFXOW WR LQYRNH HLWKHU VDOLQLW\ DW\SLFDO LRQLF UDWLRV RU SDXFLW\ RI JHQHWLFSRWHQWLDO DV EDUULHUV per se WR WKH GHYHORSPHQW DQG SUROLIHUDWLRQ RI GLD]RWURSKV ,W LV VWULNLQJ KRZHYHU WKDW YLUWXDOO\ DOO H[DPSOHV FLWHG DERYH DUH DOO EHQWKLF RU RWKHUZLVH DWWDFKHG SRSXODWLRQV $ QRWDEOH H[FHSWLRQ WR WKH SUHSRQGHUDQFH RI EHQWKLF 1IL[LQJ F\DQREDFWHULD LV WKH SODQNWRQLF DJJUHJDWHG ILODPHQWRXV JHQXV Trichodesmium. 7KLV VXUIDFH EORRPIRUPLQJ QRQKHWHURF\VWRXV GLD]RWURSK LV ZLGHO\ GLVWULEXWHG LQ ROLJRWURSKLF WURSLFDO DQG VXEWURSLFDO SHODJLF ZDWHUV DQG LV FRQVLGHUHG D PDMRU FRQWULEXWRU WR RFHDQLF 1 DQG & EXGJHWV &DUSHQWHU DQG 5RPDQV .DUO HW DO &DSRQH HW DO /LNH VRPH IUHVKZDWHUEORRP IRUPHUV VXFK DV
+ : 3DHUO Anabaena DQG Aphanizomenon, Trichodesmium FHOOV FRQWDLQ JDV YHVLFOHV ZKLFK LPSDUW EXR\DQF\ DQG HQDEOH WKLV JHQXV WR RFFXS\ D QLFKH LQ 1 GHILFLHQW ROLJRWURSKLF VXUIDFHZDWHUV 1RQGLD]RWURSKLF F\DQREDFWHULDO JHQHUD DUH EURDGO\ GLVWULEXWHG WKURXJKRXW HVWXDULQH FRDVWDO DQG SHODJLF ZDWHUV &RPPRQ SODQNWRQLF JHQHUD LQFOXGH FRFFRLG HJ Synechococcus, Synechocystis, Merismopedia DQG D YDULHW\ RI QRQKHWHURF\VWRXV ILODPHQWRXV (Oscillatoria, 3KRUPLGLXP IRUPV %HFDXVH WKH\ DUH JHQHUDOO\ VPDOO DQG GR QRW IRUP VXUIDFH EORRPV FRFFRLG F\DQREDFWHULD DUH OHVV FRQVSLFXRXV WKDQ WKHLU IUHVKZDWHU QXLVDQFH FRXQWHUSDUWV DQG WKH\ ZHUH JHQHUDOO\ RYHUORRNHG DQG XQGHUHVWLPDWHG DV FRQWULEXWRUV WR SK\WRSODQNWRQ ELRPDVV DQG SURGXFWLYLW\ LQ WKHVH ZDWHUV 'HWDLOHG H[DPLQDWLRQV XVLQJ HSLIOXRUHVFHQFH PLFURVFRS\ 'DYLV DQG 6LHEXUWK /L ,WWXULDJD DQG 0LWFKHOO IORZ F\WRPHWU\ FRXSOHG WR IOXRUHVFHQFH GHWHFWLRQ 2OVRQ HW DO KLJK SHUIRUPDQFH OLTXLG FKURPDWRJUDSK\ +3/& DQG GLDJQRVWLF SKRWRSLJPHQW FDURWHQRLGV DQDO\VHV 3LQFNQH\ HW DO OHG WR D JUHDWHU DSSUHFLDWLRQ RI WKH SUHYDOHQFH DQG LPSRUWDQFH RI WKHVH WD[D 5HFHQW VWXGLHV LQ WKH HXWURSKLF 1HXVH 5LYHU HVWXDU\ 1RUWK &DUROLQD 86$ H[HPSOLILHG WKH JURZLQJ DZDUHQHVV RI WKH LPSRUWDQFH RI QRQEORRP WD[D LQ HVWXDULQH DQG FRDVWDO ZDWHUV 3LQFNQH\ HW DO 3ULRU WR WKH ODWH V FRFFRLG DQG ILODPHQWRXV F\DQREDFWHULD ZHUH ODUJHO\ WKRXJKW WR EH FRQILQHG WR WKH XSSHU IUHVKZDWHU FRPSRQHQWV RI WKLV HVWXDULQH V\VWHP 7KH LQWURGXFWLRQ RI +3/& DV D TXDOLWDWLYH DQG TXDQWLWDWLYH DQDO\WLFDO WRRO LQ WKH ODWH V LQFUHDVHG RXU FRQFHSWXDO DQG IXQFWLRQDO DSSUHFLDWLRQ RI WKHVHJURXSVLQ WKHORZHU PRUH VDOLQH VHJPHQWV RI WKH HVWXDU\ &\DQREDFWHULD OLNH PDQ\ LPSRUWDQW HXNDU\RWLF SK\WRSODQNWRQ JURXSV SRVVHVV XQLTXH DFFHVVRU\ SKRWRSLJPHQWV LQFOXGLQJ WKH FDURWHQRLGV ]HD[DQWKLQ P\[R[DQWKRSK\OO HFK LQHQRQH DSKDQ[DQWKLQ DQG RVFLOOD[DQWKLQ 5RZDQ 0LOOLH HW DO 5RXWLQH PRQLWRULQJ DQG DQDO\VHV RI FKORURSK\OOV a, b DQG c DQG FDURWHQRLGV LQGLFDWHGWKDWF\DQREDFWHULDZHUHDKLJKO\ VLJQLILFDQW DQG DW WLPHV D GRPLQDQW FRPSRQHQW RI WKH SK\WRSODQNWRQ FRPPXQLW\ 3ODWH 3LQFNQH\ HW DO $QDO\WLFDO WHFKQLTXHV ZHUH GHYHORSHG IXUWKHU WR LQFOXGH UDGLRGHWHFWLRQ DQG TXDQWLWDWLRQ RI SKRWRSLJPHQWV WKDW LQFRUSRUDWHG & GXULQJ PHDVXUHPHQWV RI SULPDU\ SURGXFWLYLW\ 5HGDOMH *HRULFNH DQG :HOVKPH\HU D E 8VLQJ WKLV DSSURDFK WKH UDWH RI V\QWKHVLV RI
&KDSWHU 0DULQH 3ODQNWRQ SKRWRSLJPHQWV ZDV XVHG DV DQ LQGLFDWRU RI JURXS VSHFLILF JURZWK UDWHV 3LQFNQH\ HW DO 7KHVH VWXGLHV SRLQWHG WR F\DQREDFWHULD DV G\QDPLF DQG TXDQWLWDWLYHO\ LPSRUWDQW FRQWULEXWRUV WR FRPPXQLW\ SURGXFWLYLW\ DQG FRPSRVLWLRQ 3ODWHV F G H 1XWULHQW GULYHQ HXWURSKLFDWLRQ RI WKLV DQG RWKHU HVWXDULQH DQG FRDVWDO ZDWHUV PD\ EH HQKDQFLQJ WKH H[SDQVLRQ RI F\DQREDFWHULDO SURGXFWLRQ DQG GRPLQDQFH 3DHUO E 7KH SDVW IRXU \HDUV RI DQDO\VHV LQ WKH 1HXVH 5LYHU HVWXDU\ LQGLFDWH D WUHQG RI LQFUHDVLQJ ELRPDVV DQG GRPLQDQFH E\ F\DQREDFWHULD 2Q D VHDVRQDO EDVLV WKLV WUHQG LV PRVW QRWLFHDEOH GXULQJ VXPPHU PRQWKV ZKHQ HOHYDWHG ZDWHU WHPSHUDWXUHV HQKDQFH F\DQREDFWHULDO JURZWK 0LFURVFRSLF REVHUYDWLRQV FRQGXFWHG RYHUWKHSDVWGHFDGHDSSHDUWR FRQILUPWKLV WUHQG 7KH UHDGLO\ GHWHFWDEOH ILODPHQWRXV QRQ KHWHURF\VWRXV JHQHUD Oscillatoria DQG Phormidium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
% 1XWULWLRQDO&RQVWUDLQWV :LWK UHVSHFW WR QXWULHQW VXSSO\ UDWHV FRQFHQWUDWLRQV DQG UDWLRV QXPHURXV HVWXDULHV VKRXOG EH DEOH WR VXSSRUW F\DQREDFWHULD LQFOXGLQJ KHWHURF\VWRXV 1 IL[LQJ JHQHUD 7KH ODWWHU KRZHYHU DUH FRQILQHG WR SHULRGLFDOO\ VWUDWLILHG ZDWHUV HJ Nodularia RUPD\ RFFXU DV HQGRV\PELRQWV LQ GLDWRPV LH RhizosoleniaRichelia ,QWHUHVWLQJO\HQGRV\PELRWLFKHWHURF\VWRXV F\DQREDFWHULD FDQ EH IRXQG HYHQ LQ ZHOO PL[HG XOWUD ROLJRWURSKLFVXUIDFHZDWHUV VXFK DV WKH6DUJDVVR 6HD 7KHUHIRUH LW VHHPV XQOLNHO\ WKDW ZLGHVSUHDG LQRUJDQLF QXWULHQW 3 )H 0R RWKHU WUDFH PHWDOV GHILFLHQFLHV UHDGLO\ H[SODLQ WKH XQILOOHG QLFKH SKHQRPHQRQ LQ HVWXDULHV 6SHFLILF QXWULHQW GHILFLHQFLHV PD\ KRZHYHU VHOHFWLYHO\ UHVWULFWWKHDFWLYLWLHVDQG JHRJUDSKLFH[WHQW RI ERWK GLD]RWURSKLF DQG QRQGLD]RWURSKLF
F\DQREDFWHULD 2I WKRVH LQRUJDQLF QXWULHQWV H[FOXGLQJ QLWURJHQ ZKLFK PD\ FRQWURO RFHDQLF SK\WRSODQNWRQ SULPDU\ SURGXFWLYLW\ DQG JURZWK JHRFKHPLVWV RIWHQ PHQWLRQHG 3 'RUHPXV 6PLWK 7KLV YLHZ ZDV EDVHG RQ WKH DVVXPSWLRQ WKDWELRORJLFDO 1 IL[DWLRQ ZDV FDSDEOH RI VDWLVI\LQJ RFHDQLF 1 UHTXLUHPHQWV RYHU ORQJ WLPH VFDOHV LH PLOOLRQV RI \HDUV )RU ELRORJLFDOO\ UHOHYDQW WLPH VFDOHV LH GD\V WR \HDUV WKHUH LV OLWWOH H[SHULPHQWDO HYLGHQFHWR VXSSRUWWKLV DVVXPSWLRQ DQG RQ WKHZKROH WKH RFHDQV UHPDLQ 1 OLPLWHG &DUSHQWHU DQG &DSRQH )XUWKHUPRUH RFHDQLF1 IL[DWLRQ LV FRQWUROOHG E\ D FRPSOH[ LQWHUSOD\ RI SK\VLFDO FKHPLFDO DQG ELRWLF IDFWRUV JRLQJ IDU EH\RQG PHUH 3 VXSSO\ DORQH3DHUO ,PSRUWDQWIDFWRUVLQFOXGH WXUEXOHQFH 2 WHQVLRQ RUJDQLF PDWWHU VXSSO\ WUDFH PHWDO SDUWLFXODUO\ )H DYDLODELOLW\ HVWDEOLVKPHQW RI V\PELRVHV HWF 3KRVSKRUXV VXSSO\ UDWHV KDYH KRZHYHU EHHQ VKRZQ WR FRQWURO SURGXFWLRQ DQG 1 IL[DWLRQ LQ VRPH ORFDOL]HG KLJKO\ SURGXFWLYH EHQWKLF KDELWDWV LQFOXGLQJ UHHIV VHDJUDVV EHGV PLFURELDO PDWV FDUERQDWH VHGLPHQWV DQG PDQJURYH FRPPXQLWLHV 3DHUO HW DO &DSRQH ,Q DGGLWLRQ KDELWDWV ZKLFK UHFHLYH HOHYDWHG OHYHOV RI DQWKURSRJHQLFDOO\ JHQHUDWHG QXWULHQWV HJ 1 ULFK VHZDJH RXWIDOOV DJULFXOWXUDO DQG XUEDQ UXQRII DWPRVSKHULF GHSRVLWLRQ FDQ DW WLPHV H[KLELW 3 OLPLWDWLRQ6PLWK ,Q UHJDUG WR QXWULHQW UHTXLUHPHQWV IRU JURZWK FRDVWDO DQG SHODJLF ZDWHUV PD\ DW WLPHV KDYH KLJK 1 DQG 3 FRQFHQWUDWLRQV RWKHU QXWULHQWV VXFK DV VLOLFRQ 6L PD\ WKXV FRQWURO SURGXFWLRQ *OLEHUW HW DO :KLOH 6L OLPLWDWLRQ PD\ DSSO\ WR HXNDU\RWLF SK\WRSODQNWRQ GLDWRPV VLOLFRIODJHOODWHV F\DQR EDFWHULD H[KLELW YHU\ ORZ UHTXLUHPHQWV IRU WKLV QXWULHQW DQG KHQFH DUH QRW DIIHFWHG E\ LWV UHODWLYH VFDUFLW\ ,Q FRQWUDVW WKH DYDLODELOLW\ RI VSHFLILF WUDFH PHWDOV NQRZQ WR VHUYH DV FRIDFWRUV IRU NH\ DVVLPLODWRU\ HQ]\PHV 58%,6&2 QLWURJHQDVH QLWUDWH DQG QLWULWHUHGXFWDVH DQG IRUWKH V\QWKHVLV RIFHOOXODU FRQVWLWXHQWV PD\ EH UHVWULFWHG LQ ZDWHUV ZKHUH 1 DQG 3 DUH LQ H[FHVV,URQKDVUHFHLYHGPXFK FRQVLGHUDWLRQ LQ SDUW EHFDXVH LW LV HVVHQWLDO IRU JURZWK DQG LWV ELRORJLFDO DYDLODELOLW\ LV UHVWULFWHG GXH WR FKHPLFDO SUHFLSLWDWLRQ FKHODWLRQ DQG RWKHU QDWXUDO ELQGLQJORVV SURFHVVHV ZKLFK RFFXU LQ VHDZDWHU %UXODQG HW DO 0XFK RI WKH UHFHQW IRFXV RQ )H OLPLWDWLRQ LQ WKH RFHDQV ZDV RQ WKH H[FHVV 1 DQG 3 UHJLRQV RI WKH ROLJRWURSKLF RFHDQ LQFOXGLQJ WKH 1RUWK 3DFLILF J\UH PLG$WODQWLF DQG 6RXWKHUQ DQG $QWDUFWLF 2FHDQV 0DUWLQ HW DO ZDWHUV ZKHUH
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
+ : 3DHUO
7UHDWPHQWVFRQFHQWUDWLRQVLQ0 Fig.3. (IIHFWV RI LURQ HLWKHU DV )H&O RU ('7$FKHODWHG )H&O HTXLPRODU DQGPRO\EGHQXPDV1D0R2 RQQLWURJHQDVHDFWLYLW\ 1$ DQG FKORURSK\OO a RI Trichodesmium thiebautii. $IUHVKO\ FROOHFWHG 1RUWK &DUROLQD : $WODQWLF *XOI6WUHDP T. thiebautii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richodesmium. ,WLVWKHUHIRUH FRQFOXGHG WKDW LQFUHDVLQJ)H DYDLODELOLW\ HLWKHUYLDHQULFKPHQW RU E\ VXSSO\LQJ ('7$ VWLPXODWHG 1 IL[DWLRQ DQG JURZWK RI WKLV F\DQREDFWHULDOEORRPIRUPHU
&KDSWHU 0DULQH 3ODQNWRQ ,Q D UHODWHG K\SRWKHVLV 5XHWHU HW DO VXJJHVWHG WKDW E\ IRUPLQJ ODUJH EXR\DQW DJJUHJDWHV QHDU WKH ZDWHU V VXUIDFH Trichodesmium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nabaena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
&7KH2[\JHQ3UREOHP $V R[\JHQLF SKRWRWURSKV F\DQREDFWHULD DUH IDFHG ZLWK D IRUPLGDEOH HFRSK\VLRORJLFDO SUREOHP PROHFXODUR[\JHQ 2 HYROYHGGXULQJSKRWRV\QWKHVLV LV D SRWHQW LQKLELWRU RI QLWURJHQDVH WKH HQ]\PH FRPSOH[ ZKLFK PHGLDWHV 1 IL[DWLRQ
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scillatoria, Lyngbya DQG Trichodesmium, DQG FRFFRLG IRUPV LQFOXGLQJ Synechococcus DQG Gloeothece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ynechococcus,
+ : 3DHUO Synechocystis, Erythrospira DQG Gloeothece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scillatoria DQG Lyngbya 6WDO DQG .UXPEHLQ 6WDO DQG %HUJPDQ 3DHUO HW DO &XUULQ HW DO 0DULQH F\DQREDFWHULDO PDW FRPPXQLWLHV GRPLQDWHGE\WKHVHIRUPVRIWHQVKRZSURQRXQFHGGLHO SDWWHUQV RI SKRWRV\QWKHVLV GXULQJ WKH GD\ DQG 1 IL[DWLRQPD[LPDDWQLJKW%HERXWHWDO 3DHUOHW DO 9LOOEUDQGW HW DO :KHQ JURZQ HLWKHUXQGHUORZWXUEXOHQFHFRQGLWLRQV OLNHSODQNWRQ RUDVEHQWKLFPDWVWKHFRVPRSROLWDQVSHFLHVLyngbya aestuarii VKRZV RSWLPL]DWLRQ RI1 IL[DWLRQ DW QLJKW EXW DOVR ORZ OHYHOV RI 1 IL[DWLRQ GXULQJ WKH GD\ 3DHUO HW DO ,Q WKLV GLD]RWURSK WZR GLVWLQFW VWUDWHJLHV DSSHDU WR RSHUDWH 7KH QRFWXUQDO RSWLPL]DWLRQ VWUDWHJ\ LV VLPLODU WR WKDW GHVFULEHG IRU Oscillatoria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
&KDSWHU 0DULQH3ODQNWRQ
Fig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
,PPXQRJROG ODEHOLQJ RI Lyngbya aestuarii VXJJHVWV WKDW ERWK GLVWDO DQG SUR[LPDO FHOOV FRQWDLQ WKH 1IL[LQJ HQ]\PH FRPSOH[ QLWURJHQDVH DQG PD\ LPSO\ WKDW 1 IL[DWLRQ FDQ RFFXU LQ ERWK UHJLRQV 3DHUO HW DO 3UHVXPDEO\ ZLWK WKH DEVHQFH RI 2HYROYLQJ SKRWRV\QWKHVLV LQ FHUWDLQ GLVWDO FHOOV L.
aestuarii VKRXOG EH DEOH WR IL[ 1 DOEHLW LQ OLPLWHG DPRXQWV GXULQJ GD\OLJKW 7KLV VSDWLDOSDUWLWLRQLQJ RI 1 DQG &2 IL[DWLRQLV QRW FRQVLVWHQWO\REVHUYHG LQ L. aestuarii; LW DSSHDUV GHSHQGHQW RQ JURZWK SKDVH DQG GLHOSHULRGLFLW\ Tnchodesmium VKRZV 7KH EORRPIRUPLQJ FRQWHPSRUDQHRXV GD\WLPH PD[LPD RI 1 IL[DWLRQ &2 IL[DWLRQ DQG 2 HYROXWLRQ &DUSHQWHU D E &DSRQH HW DO 7KLV SURSHUW\ GLVWLQJXLVKHV Trichodesmium IURP DOO RWKHU PDULQH &\DQREDFWHULDO GLD]RWURSKV QRQKHWHURF\VWRXV &DSRQH HW DO GHPRQVWUDWHG WKDW HYHQ ZKHQ VXSSOLHG ZLWK OLJKW ZHOO LQWR WKH KRXUV RI QLJKW 1 IL[DWLRQ EXW QRW &2 IL[DWLRQ FHDVHG 7KHVH REVHUYDWLRQV VXJJHVW D VWURQJ GD\WLPH GHSHQGHQFH DQG D GLHO UK\WKP LQ LWV GLD]RWURSKLF EHKDYLRU 0ROHFXODU VWXGLHV LQGLFDWH WKDW WKH QLWURJHQDVH HQ]\PH FRPSOH[ RI Trichodesmium LV KLJKO\ FRQVHUYHG DQG VLPLODU WR WKDW IRXQG LQ RWKHU GLD]RWURSKV =HKU HW DO &DSRQH HW DO )XUWKHUPRUH WKH QLWURJHQDVH RI Trichodesmium LV 2 VHQVLWLYHOLNHWKHHQ]\PHV RI RWKHUGLD]RWURSKV +RZ WKHQ GRHV Trichodesmium IL[ 1 LQ WKH SUHVHQFH RI R[\JHQLF SKRWRV\QWKHVLV" ,Q LWV RFHDQLF KDELWDW Trichodesmium RFFXUV DV EXR\DQW PDFURVFRSLF D IHZ PP LQ OHQJWK UDGLDO SXII RUIXVLIRUPWXIW DJJUHJDWHV3ODWHV F G H :KHQ FDOP VHDV SUHYDLO DJJUHJDWHV FDQ IUHTXHQWO\ DFFXPXODWH LQ VXUIDFH ZDWHUV LPSDUWLQJ EURZQLVK \HOORZ VOLFNV ZKLFK FDQ H[WHQG NLORPHWHUV RU PRUH )LHOG VWXGLHV LQGLFDWHG WKDW Trichodesmium IL[HG 1 DW PD[LPXP UDWHV ZKHQ LW ZDV SUHVHQW DV VXUIDFHGZHOOLQJ DJJUHJDWHV &DUSHQWHU 3DHUO DQG %HERXW $JJUHJDWHVKDYHFRPSDFWGHQVH GDUNSLJPHQWHG FHQWUDO FRUHV ZKLOH WKHLU SHULSKHUDO UHJLRQV DSSHDU GLIILVH DQG KDLUOLNH )LJ 7KH FRQWUDVW EHWZHHQ FRUH DQG SHULSKHUDO UHJLRQV LV HYLGHQW ZKHQ YLHZLQJ VXVSHQGHG DJJUHJDWHV XQGHU QDWXUDO FRQGLWLRQV LH LQ WKH XSSHU UHJLRQ RI WKH ZDWHU FROXPQ 7KH GLIIHUHQFH LQ FRPSDFWLRQ DQG UHVXOWDQWFKDQJH LQ RSWLFDO GHQVLW\ EHWZHHQ LQQHU DQG RXWHU UHJLRQV RI Trichodesmium FRORQLHV LV VXJJHVWLYH RI PRUSKRORJLFDO DQG IXQFWLRQDO GLIIHUHQWLDWLRQ )RJJ K\SRWKHVL]HG WKDW &2 DQG 1 IL[DWLRQ PD\ EH SDUWLWLRQHG +H VXJJHVWHG WKDW FDUERQ GLR[LGH IL[DWLRQ ZDV FRQILQHG WR H[WHUQDO UHJLRQV VXEMHFW WR KLJK LUUDGLDQFH DQG 1 IL[DWLRQ ZDV FRQILQHG WR GHQVH VHOIVKDGHG LQWHUQDO UHJLRQV ,I LW LV DVVXPHG WKDW SKRWRV\QWKDWH DQG UHFHQWO\ V\QWKHVL]HG RUJDQLF 1 FRPSRXQGV DUH H[FKDQJHG EHWZHHQ LQQHU DQG RXWHU UHJLRQV RI DJJUHJDWHV WKHQ
Fig. 5. 0RUSKRORJ\ RI Trichodesmium thiebautii DJJUHJDWHV YLHZHG ZLWK VFDQQLQJ HOHFWURQ PLFURVFRS\ VKRZLQJ SXII DQG WXIW VKDSHG FRORQLHV FRPPRQO\ HQFRXQWHUHG LQ RFHDQLF QHDUVXUIDFH EORRP SRSXODWLRQV 6DPSOHV ZHUH REWDLQHG IURP WKH :HVWHUQ $WODQWLF 2FHDQ QHDU 6DQ 6DOYDGRU ,VODQG 7KH %DKDPDV
WKLV PD\ VXJJHVW D PHFKDQLVP E\ ZKLFK &2 DQG 1 FRXOG EH IL[HG VLPXOWDQHRXVO\ )RJJ &DUSHQWHU DQG 3ULFH JHQHUDWHG HYLGHQFH LQ VXSSRUW RI )RJJ V K\SRWKHVLV E\ XVLQJ PLFURDXWRUDGLRJUDSK\ WR VKRZ WKDW H[WHUQDO UHJLRQV RI LQGLYLGXDOWULFKRPHV ZHUH SKRWRV\QWKHWLFDOO\ PXFK PRUH DFWLYH WKDQ LQWHUQDO UHJLRQV 7KH\ DOVR VKRZHG D VWURQJ SRVLWLYH FRUUHODWLRQ EHWZHHQ FDOP VHD VWDWH IDFLOLWDWLQJ DJJUHJDWLRQ DQG UHODWLYHO\ KLJK UDWHV RI 1 IL[DWLRQ LQ &DULEEHDQ 6HD Trichodesmium SRSXODWLRQV %U\FHVRQ DQG )D\ DOVR VKRZHG FORVH UHODWLRQVKLSV EHWZHHQ DJJUHJDWH VKDSH VL]H DQG 1 IL[DWLRQ SRWHQWLDOV LQ Trichodesmium VDPSOHG RII WKH HDVW FRDVW RI $IULFD $Q H[DPLQDWLRQ RI WHWUD]ROLXP G\H 77& UHGXFWLRQ VKRZHG WKDW KLJKO\ UHGXFHG 2 GHSOHWH ]RQHV ZHUH SUHVHQW LQ FRUH UHJLRQV RI DFWLYHO\ 1 IL[LQJ ZHVWHUQ $WODQWLF *XOI 6WUHDP Trichodesmium DJJUHJDWHV 3DHUO DQG %ODQG $GGLWLRQDO HYLGHQFH IRU WKH SUHVHQFH RI VPDOO PP VFDOH 2
+ : 3DHUO GHSOHWH LQWHUQDO UHJLRQV FDPH IURP VWXGLHV ZLWK PLFURHOHFWURGHV 3HDUO DQG %HERXW ,Q IXUWKHU VXSSRUW RI )RJJ V K\SRWKHVLV WKHVH VWXGLHV GRFXPHQWHG KRZ WKH R[\JHQ WHQVLRQ RI WKH H[WHUQDO HQYLURQPHQW RI Trichodesmium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richodesmium VSS 2KNL DQG )XMLWD 3UXIHUW HW DO 7KLV IDFLOLWDWHG ZHOO GHILQHG FRQWUROOHG H[DPLQDWLRQV RI &2 DQG 1 IL[DWLRQ LQ WKLV F\DQREDFWHULXP :KLOH VPDOO SXII DQG WXIW DJJUHJDWHV FDQ EH FXOWXUHG VLQJOH WULFKRPHV IUHTXHQWO\ SUHGRPLQDWH 2KNL DQG )XMLWD 3UXIHUW HW DO 6LQJOH WULFKRPHV DUH FDSDEOH RIOLJKWPHGLDWHG 1 IL[DWLRQ DOEHLW DW ORZ UDWHV SHU DPRXQW RI FKORURSK\OO a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richodesmium LV PRVW RIWHQ HQFRXQWHUHG DV GHQVHO\ SDFNHG DJJUHJDWHV )LJ 7KH DELOLW\ RI VLQJOH WULFKRPHV WR IL[ ERWK &2 DQG 1 LQ FXOWXUH PD\ EH UHODWHG WR WKH ORZ OHYHOV RI SKRWRV\QWKHVLV 2 SURGXFWLRQ H[SHULHQFHG DW UHGXFHG LUUDGLDQFH PRO SKRWRQ P V
&KDSWHU 0DULQH 3ODQNWRQ
)LJ 3KRWRV\QWKHWLF &2 EDVHG RQ &2 LQFRUSRUDWLRQ DQG 1 IL[DWLRQ QLWURJHQDVH DFWLYLW\ LQ VLQJOH WULFKRPH FXOWXUHV RID UHFHQW 7ULFKRGHVPLXPLVRODWH,06 GHULYHGIURP$WODQWLF2FHDQZDWHUV3UXIHUW%HERXWHWDO $WPROSKRWRQP V 3$5WKLV LVRODWH LV FDSDEOH RIVLPXOWDQHRXV &2 DQG 1 IL[DWLRQ $W KLJKHU LUUDGLDQFHV ! PRO SKRWRQ P V &2 IL[DWLRQ FRQWLQXHV EXW 1 IL[DWLRQFHDVHVOHDGLQJWR1VWDUYDWLRQDQGGHDWKLQWKHVH1GHSOHWHZDWHUV
$SSDUHQWO\ 1 IL[DWLRQ FDQ DFFRPSDQ\ SKRWRV\QWKHVLV LQ WKLV F\DQREDFWHULXP DV ORQJ DV FHOOXODUUDWHVRI2 SURGXFWLRQGRQRWH[FHHGDFULWLFDO WKUHVKROG ZKLFK LV LQKLELWRU\ WR QLWURJHQDVH 7KHVH FRQGLWLRQV SUHYDLO LQ WKH ORZHU WKLUG RI WKH ZDWHU FROXPQ ZKHUH WKH SRWHQWLDO IRU VLQJOH WULFKRPH SRSXODWLRQV RI Trichodesmium WR H[LVW PD\ EH JUHDWHVW ,PPXQRJROG ODEHOLQJ RI FXOWXUHG DQG QDWXUDO Trichodesmium SRSXODWLRQV GHPRQVWUDWHG WKH DELOLW\ RIFHOOVWR V\QWKHVL]H QLWURJHQDVH 3DHUOHWDO D %HUJPDQ DQG &DUSHQWHU 7KLV LQGLFDWHV WKDW WKH JHQHWLF SRWHQWLDO IRU 1 IL[DWLRQ PD\ EH EURDGO\ GLVWULEXWHG LQ WULFKRPHV DQG DJJUHJDWHV 7KHVH ILQGLQJV FRPELQHG ZLWK WKH REVHUYDWLRQV RI VLQJOHWULFKRPH1 IL[DWLRQ LQ FXOWXUH DQG DJJUHJDWH OHYHO 1 IL[DWLRQ LQ QDWXUH VXJJHVW nif JHQH H[SUHVVLRQ PD\ EH FRQWUROOHG E\ HQYLURQPHQWDO FRQGLWLRQVVXFKDVLUUDGLDQFH WXUEXOHQFH DQGPL[LQJ (QYLURQPHQWDO VWUHVV LQFOXGLQJ KLJK VXUIDFH LUUDGLDQFH PD\ OHDG WR PRUSKRORJLFDO DGDSWDWLRQV
VXFK DV DJJUHJDWLRQ ZKLFK IDFLOLWDWHV FRQWHPSRUDQHRXV SKRWRV\QWKHVLV DQG 1 IL[DWLRQ RQ WKH WULFKRPH DQGRU DJJUHJDWH OHYHO :K\ WKLV SODQNWRQLF F\DQREDFWHULXP GRHV QRW FDUU\ RXW D EXON RI LWV 1 IL[DWLRQ DW QLJKW LQ RUGHU WR PLQLPL]H WKH 2 SUREOHP UHPDLQV XQFOHDU VLQFH WKLV DSSHDUV WR EH D IHDWXUH RI RWKHU QRQKHWHURF\VWRXV IRUPV Lyngbya, Oscillatoria &ORVH DVVRFLDWLRQV EHWZHHQ &\DQREDFWHULD DQG EDFWHULDO RU DOJDO HSLSK\WHV DUH FRPPRQSODFH LQ PDULQH &\DQREDFWHULDO DVVHPEODJHV 3DHUO HW DO E 3DHUO )LJ ,Q WKH FDVH RI WKH KHWHURF\VWRXV JHQHUDNodularia, Aphanizomenon DQG Anabaena, VXFK EDFWHULDO DVVRFLDWLRQV PD\ HQKDQFH 1 IL[LQJ FDSDFLW\ 3DHUO DQG .HOODU .RQRQHQ HW DO 1RQKHWHURF\VWRXV JHQHUD LQFOXGLQJ Trichodesmium, Oscillatoria DQG Lyngbya H[KLELW FORVH DVVRFLDWLRQV ZLWK GLYHUVH PLFURIORUD 7KHVH DVVRFLDWLRQV PD\ KHOS WR RSWLPL]H WKH 1 IL[DWLRQ SRWHQWLDOV RI &\DQREDFWHULD SHUKDSV E\ HQKDQFLQJ ORFDOL]HG 2 FRQVXPSWLRQ DQG QXWULHQW UHJHQHUDWLRQ
+:3DHUO
G D
H
E
F
Fig. 7. 6FDQQLQJ HOHFWURQ DQG OLJKW PLFURVFRS\ REVHUYDWLRQV RI SODQNWRQLFF\DQREDFWHULDOEDFWHULDOLQWHUDFWLRQV +HWHURF\VWVSHFLILF EDFWHULDO DVVRFLDWLRQV LQ D Aphanizomenon flos-aquae, E Anabaena spiroides DQGF Nodularia VS G %DFWHULDODVVRFLDWLRQV ZLWKQRQKHWHURF\VWRXVF\DQREDFWHULDLQFOXGLQJ/\QJE\Daestuarii, H Trichodesmium thiebautii. 1RWHFORVHEDFWHULDODVVRFLDWLRQVZLWK PXFLODJLQRXVVKHDWKVDQGVOLPHOD\HUVVXUURXQGLQJWULFKRPHV
3DHUO DQG .HOODU 3DHUO DQG 3LQFNQH\ 0DQ\ PDULQH DQG IUHVKZDWHU &\DQREDFWHULD DUH QRW FXOWLYDEOH XQGHU EDFWHULDIUHH D[HQLF FRQGLWLRQV )RJJ ,W LV ZHOO NQRZQ WKDW EDFWHULDO HSLSK\WHV FRPPRQO\ REVHUYHG RQ DTXDWLF &\DQREDFWHULD XWLOL]H F\DQREDFWHULDO H[FUHWLRQ SURGXFWV &KUyVW DQG %U]HVND 3DHUO )XUWKHUPRUH EDFWHULD DVVRFLDWHG ZLWK &\DQREDFWHULD ZHUH VKRZQWR EHFKHPRWDFWLFDOO\ DWWUDFWHGWRD UDQJH RI RUJDQLF FRPSRXQGV NQRZQ WR EH FRPPRQ F\DQREDFWHULDO H[FUHWLRQ SURGXFWV 3DHUO DQG *DOOXFFL 7KHVH DVVRFLDWLRQV PD\ EHQHILW WKH &\DQREDFWHULD EHFDXVH JURZWK UDWHV RI WKH KRVW DUH
&KDSWHU0DULQH3ODQNWRQ RIWHQ HQKDQFHG DQG PD[LPL]HG LQ WKH SUHVHQFH RI EDFWHULDO HSLSK\WHV 7KHVHDVVRFLDWLRQV DSSHDUWRKDYH HYROYHG WR PD[LPL]H WKH SK\VLRORJLFDO DFWLYLWLHV RI WKHWZR RUJDQLVPV DQG WR FRXQWHUWKH SUREOHP FDXVHG E\ HOHYDWHG R[\JHQOHYHOV
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nabaena oscillarioides ZDV UDSLGO\ WUDQVIHUUHG WR KHWHURF\VW DVVRFLDWHG EDFWHULD 3DHUO $[HQLF LVRODWHV RI A. oscillarioides H[KLELWHG PD[LPXP JURZWK DQG 1 IL[DWLRQ UDWHV ZKHQ UHLQRFXODWHG ZLWK EDFWHULDO VWUDLQV GHULYHGIURPWKHKRVW3DHUODQG .HOODU 0DULQH F\DQREDFWHULDO EORRP VSHFLHV DUH QRWRULRXVO\ GLIILFXOW WR FXOWXUH D[HQLFDOO\ )RJJ 2KNL DQG )XMLWD 3UXIHUW HW DO 7KH OLWHUDWXUH RQ IUHVKZDWHU IRUPV LQFOXGH PDQ\ UHSRUWV
RQ GLIIHUHQWLDO JURZWK REWDLQHG XQGHU D[HQLF YV EDFWHUL]HG FRQGLWLRQV ZLWK IHZ H[FHSWLRQV EDFWHUL]HG IUHVKZDWHU F\DQREDFWHULDO JHQHUD H[KLELWHG KLJKHU JURZWK UDWHV DQG ZHUH HDVLHU WR PDLQWDLQ LQ FXOWXUH WKDQ D[HQLF VWUDLQV 7KLV JHQHUDO ILQGLQJ ZDV VXPPDUL]HG E\ *LEVRQ DQG 6PLWK “Experience at our laboratory exemplifies this problem (obtaining and maintaining cyanobacterial growth under axenic conditions); when both axenic Oscillatoria redekei and 2 agardhii have been isolated, we have obtained consistent growth only with 2 redekei. The 2 agardhii always appears to grow better in the presence of contaminant heterotrophic bacteria”. ,W ZDV VXVSHFWHGIRU D ORQJWLPHWKDWWKHUHOLDQFHRI &\DQREDFWHULD RQ EDFWHUL]HG FRQGLWLRQV IRU RSWLPDO JURZWK ZDV UHODWHG WR ERWK WKH H[FKDQJH RIPXWXDOO\ EHQHILFLDO PHWDEROLWHV DQG JURZWK IDFWRUV DV ZHOO DV WKH GHWR[LI\LQJ UROHV WKDW SK\FRVSKHUH DVVRFLDWHG EDFWHULD PD\ SOD\ .XHQW]HO¶V DQG /DQJH¶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icrocystis aeruginosa FRORQLHV DV ZHOO DV WKH SUHVHQFH RI DPRHERLG SURWR]RDQ JUD]HUV ZLWKLQ VXFK FRORQLHV DUH PRVW SURIRXQG GXULQJ SHULRGV RI RSWLPDO &2 IL[DWLRQ ,W ZDV VKRZQ WKDW M. aeruginosa FHOOV ZKLFK ERUGHUHG UHJLRQV EHLQJ JUD]HG H[KLELWHG KLJKHU FHOOVSHFLILF UDWHV RI &2 IL[DWLRQ WKDQ FHOOV PRUH GLVWDQW IURP JUD]HUV 7DEOH 7KXV HYHQ WKRXJK SURWR]RDQ JUD]LQJ UHSUHVHQWV FRQVXPSWLRQ RI F\DQREDFWHULDO
+ : 3DHUO
Table 4. 0LFURDXWRUDGLRJUDSKLF GHWHUPLQDWLRQ RIFHOOVSHFLILFUDWHV RISKRWRV\QWKHWLF &2 IL[DWLRQ LQ FRORQLHV RI0LFURF\VWLVDHUXJLQRVD 2ULJLQDOVWRFNVZHUHGHULYHGIURPDEORRPRQWKH1HXVH5LYHU 1& 7KH\ ZHUHFXOWXUHGRQ $60 1PHGLXP DQGH[SRVHGWRPRO SKRWRQ P V 3$5 DW & 'HWHUPLQDWLRQV ZHUH PDGH RQ D[HQLF EDFWHULD DQG SURWR]RDIUHH Amoeba VSSDVVRFLDWHG DQG EDFWHULDDVVRFLDWHGEDFWHULDODUJHO\FRQILQHGWRPXFLODJLQRXVVKHDWKV SRSXODWLRQVRIM. aeruginosa. 0HDQRI LQGHSHQGHQWH[SHULPHQWV 6(GDWDIURP3DHUO 'DWH
SJ& FHOO K $[HQLF
$PRHED$VVRF
%DFWHULD $VVRF
2FW
1RY
1RY
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a P0 S+ UHPDLQV TXLWH FRQVWDQW a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richodesmium VSS LV EXR\DQW
&KDSWHU 0DULQH3ODQNWRQ HYHQ ZKHQ ODUJH FKDQJHV LQ LUUDGLDQFH DUH HQFRXQWHUHG 3HUVLVWHQW EXR\DQF\ ZDV DWWULEXWHG WR H[WUHPHO\ VWURQJ PHPEUDQHERXQG JDV YHVLFOHV 'HVSLWHLWVDELOLW\WRUHPDLQEXR\DQWDQGLWVFDSDFLW\ WR UHPDLQ LQ QHDU VXUIDFH ZDWHUV XQGHU D YDULHW\ RI HQYLURQPHQWDO FRQGLWLRQV Trichodesmium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ynechococcus DQG Synechocystis, RIWHQ FRPSHWH HIIHFWLYHO\ ZLWK HXNDU\RWHV IRU QXWULHQWV DQG OLJKW XQGHU KLJKO\ PL[HG FRQGLWLRQV 7KH\ JHQHUDOO\ H[KLELW KLJK DIILQLWLHV IRU QXWULHQWV DW ORZ FRQFHQWUDWLRQV DQG DV VXFK FDQ IRUP D VXEVWDQWLDO IUDFWLRQ RIWKH SK\WRSODQNWRQ ELRPDVV LQ ZHOOPL[HG HVWXDULHV FRDVWDO DQG SHODJLF ZDWHUV ZKLFK H[SHULHQFHFKURQLF1DQGRWKHUQXWULHQWOLPLWDWLRQV 6PDOO VFDOH WXUEXOHQFH RU VKHDU ZDV VKRZQ WR EH DQ LPSHGLPHQW WR RSWLPDO 1 IL[DWLRQ DQG JURZWK UDWHV LQ VWLUUHG FXOWXUHV RI KHWHURF\VWRXV &\DQREDFWHULD )RJJ .XFHUD HW DO XQSXEOLVKHGGDWD 7KHQHJDWLYHHIIHFWVPD\EHGXHWR EUHDNDJH RI F\DQREDFWHULDO ILODPHQWV VSHFLILFDOO\ DW WKH GHOLFDWH KHWHURF\VWYHJHWDWLYH FHOOMXQFWLRQ /DQJ DQG )D\ SURPRWLRQ RI 2 LQDFWLYDWLRQ RI QLWURJHQDVHLQKHWHURF\VWV )RJJ RUGLVUXSWLRQ RI DVVRFLDWLRQV EHWZHHQ EDFWHULD DQG &\DQREDFWHULD ZKLFK DUH NQRZQ WR SURPRWH WKH SRWHQWLDO RI
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¶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
+:3DHUO
*
e/h &DPSDQG6WHLQ
$VVXPLQJ WKH G\QDPLF YLVFRVLW\ K KDV D YDOXH RI FP V D ORZ YDOXH RI H FP V IRU RSHQ ZDWHUHQYLURQPHQWV DQG DKLJK YDOXH RIH FP V IRU WLGDO FKDQQHOV WKHQ * LV H[SHFWHG WR UDQJH EHWZHHQ V DQG V UHVSHFWLYHO\ 7KRPDV DQG *LEVRQ 'DP HW DO 2XU &RXHWWH &KDPEHUVFDQJHQHUDWHVKHDUUDWHVIURPWRV )RU LOOXVWUDWLYH SXUSRVHV FRQVLGHU D FRPSDULVRQ RI 1 IL[DWLRQ QLWURJHQDVH DFWLYLW\ DQG &2 IL[DWLRQ SULPDU\SURGXFWLYLW\ LQUHSUHVHQWDWLYHKHWHURF\VWRXV VSHFLHV Anabaena oscillarioides DQG A. circinalis, H[SRVHG WR VPDOOVFDOH VKHDU UHSUHVHQWDWLYH RI HVWXDULQH 1HXVH 5LYHU HVWXDU\ PL[HG OD\HU
FRQGLWLRQV +LJKVKHDUUDWHVQHJDWLYHO\DIIHFW1 DQG &2 IL[DWLRQ 7DEOH $IWHU KRXUV WKH LQIOXHQFH RIKLJK VKHDU WUHDWPHQWRQA. oscillarioides ZDVVLJQLILFDQWO\JUHDWHUWKDQWKHLQIOXHQFHRIWKHORZ VKHDU WUHDWPHQW DQG WKH FRQWURO &DUERQ GLR[LGH IL[DWLRQ DQG 1 IL[DWLRQ RI A. circinalis ZHUH VLJQLILFDQWO\ORZHULQWKHKLJKVKHDUWUHDWPHQWUHODWLYH WRWKHFRQWURO 7DEOH 6LQFH1 IL[DWLRQ ZKLFK LV UHOLDQW RQ RSWLPDO SKRWRV\QWKHWLF SHUIRUPDQFH LV NQRZQ WR FRQIHU D VHOHFWLYH DGYDQWDJH WR F\DQREDFWHULDO EORRPJHQHUDLQ 1GHSOHWH ZDWHUV WKH GHWULPHQWDO HIIHFWV RI KLJK VKHDU RQ WKHVH SURFHVVHV FRXOG SRWHQWLDOO\ QHJDWH WKLV DGYDQWDJH 3HUVLVWHQW KLJK VKHDU UHSUHVHQWDWLYH RI ZLQG DQG WLGH H[SRVHG ZDWHUV PD\ WKHUHIRUH DFW DV D LPSHGLPHQW WR WKH HVWDEOLVKPHQWDQG SUROLIHUDWLRQ RIGLD]RWURSKLFEORRP JHQHUD
Table 5. ,PSDFWRIVPDOOPVFDOH VKHDUVWUHVVUHSUHVHQWDWLYHRIZLQGVKHDURQHVWXDULQHDQGFRDVWDOVXUIDFHZDWHUVPL[HGOD\HU RQ&2 DQG1 IL[DWLRQQLWURJHQDVHDFWLYLW\ LQ$QDEDHQDRVFLOODULRLGHVDQG$FLUFLQDOLV.XFHUDDQG3DHUOXQSXEOLVKHGGDWD :HOOGHILQHGVKHDU IRUFHVZHUHLPSDUWHGE\&RXHWWHFKDPEHUV &RQWUROFRQGLWLRQVUHSUHVHQWQRVKHDUVWUHVVORZVKHDUVWUHVVUHSUHVHQWVPRGHUDWHO\ZLQG\ FRQGLWLRQVDQGKLJKVKHDUVWUHVVUHSUHVHQWVKLJKO\ZLQG\FRQGLWLRQV )LJXUHVDUHPHDQ6(
6KHDUVWUHVV
FRQWURO ORZVWUHVV KLJKVWUHVV
A.oscillarioides
A.circinalis
1 )L[DWLRQ QPRO&+ PJFKO K
&2 )L[DWLRQ PJ&PJFKO K
1 )L[DWLRQ QPRO&+ PJFKO K
&2 )L[DWLRQ PJ&PJFKO K
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
&KDSWHU 0DULQH3ODQNWRQ
Fig. 8. &RQFHSWXDOGLDJUDPRIFRQWHPSRUDQHRXVDQGFRQWLJXRXVLQWHUDFWLQJSK\VLFDOFKHPLFDODQGELRWLFFRQWUROVRQSODQNWRQLFF\DQREDFWHULDO JURZWKDQGEORRPG\QDPLFVLQWKHPDULQHHQYLURQPHQWDIWHU3DHUODQG0LOOLH
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
SUHYDLOLQJ SK\VLFDOFKHPLFDO FRQGLWLRQV PD\ \LHOG D ZLGH UDQJH RI FRQGLWLRQV ZKLFK PD\ SRVLWLYHO\ RU DGYHUVHO\LQIOXHQFHERWKWKHVWUXFWXUHDQGIXQFWLRQ RI F\DQREDFWHULDOFRPPXQLWLHVDQGWKHFDSDFLW\RIWKRVH FRPPXQLWLHV WR FRPSHWH ZLWK WKH HXNDU\RWLF FRPSRQHQW RI WKH SK\WRSODQNWRQ FRPPXQLW\ 6XFK FRPSOH[ LQWHUDFWLRQV PXVW EH FRQVLGHUHG FDUHIXOO\ GXULQJIXWXUHHIIRUWVWKDWXVHPRGHOLQJWRHQKDQFHRXU XQGHUVWDQGLQJRIKRZ&\DQREDFWHULDFRPH RUGR QRW FRPHWR GRPLQDWHWKHVH PDULQH HFRV\VWHPV 'HVSLWH WKHLU ³SULPLWLYH´ VWUXFWXUDO DQG HYROXWLRQDU\ SURSHUWLHV WKH F\DQREDFWHULDO SK\WRSODQNWRQ UHSUHVHQWV DQ HFRSK\VLRORJLFDOO\ FRPSOH[ JURXS ZKLFK LQ PDQ\ UHVSHFWV UHPDLQV HQLJPDWLF DQG FKDOOHQJLQJDVZHVWULYHWREHWWHUXQGHUVWDQGVWUXFWXUH DQGIXQFWLRQRIPDULQHHFRV\VWHPV
$FNQRZOHGJPHQWV ,WKDQN-3LQFNQH\-=HKU(&DUSHQWHU'&DSRQH DQG'0LOOLHIRUWHFKQLFDODVVLVWDQFHGLVFXVVLRQVDQG FULWLFDO UHYLHZV RI WKLV FKDSWHU 5HVHDUFK DQG GRFXPHQWDWLRQ VXSSRUWZDV SURYLGHG E\ WKH 1DWLRQDO 6FLHQFH )RXQGDWLRQ 3URMHFWV '(% '(% ,%1 2&( 7KH1RUWK &DUROLQD 6HD *UDQW 3URJUDP 50(5 DQG WKH 86 'HSDUWPHQWRI$JULFXOWXUH 3URMHFW
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richodesmium thiebautii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
+ : 3DHUO %U\FHVRQ , DQG )D\ 3 1LWURJHQ IYWDWLRQ LQ Oscillatoria (Trichodesmium) erythraea LQ UHODWLRQ WR EXQGOH IRUPDWLRQ DQG WULFKRPHGLIIHUHQWLDWLRQ 0DU%LRO &DOGZHOO '( $VVRFLDWLRQV EHWZHHQ SKRWRV\QWKHWLF DQG KHWHURWURSKLF SURNDU\RWHV LQ SODQNWRQ ,Q 1LFKROV -0 HG $EVWUDFWV RI 7KLUG ,QWHUQDWLRQDO 6\PSRVLXP RQ 3KRWRV\QWKHWLF 3URNDU\RWHV &DOZHOO'( DQG&DOGZHOO 6- $Zoogloea VS DVVRFLDWHG ZLWKEORRPV RIAnabaena flos-aquae. &DQ- 0LFURELRO &DPS75DQG6WHLQ3& 9HORFLW\JUDGLHQWVDQGLQWHUQDOZRUN LQIOXLGPRWLRQ - %RVWRQ6RF &LYLO(QJ &DSRQH'* %HQWKLFQLWURJHQIL[DWLRQ,Q &DUSHQWHU(7DQG &DSRQH'*HGV 1LWURJHQLQWKH0DULQH(QYLURQPHQW SS $FDGHPLF3UHVV1HZ
&KDSWHU 0DULQH3ODQNWRQ '¶(OLD &) 6DQGHUV -* DQG %R\QWRQ :5 1XWULHQW HQULFKPHQW VWXGLHV LQ D FRDVWDO SODLQ HVWXDU\ 3K\WRSODQNWRQ JURZWKLQODUJHVFDOHFRQWLQXRXVFXOWXUHV &DQ-)LVK$TXDW6FL 'DP +* 'UDSHDX '7 DQG *UHQLHU * $Q LPSURYHG IORFFXODWLRQGHVLJQ IRUXVHLQ SDUWLFOHDJJUHJDWLRQ H[SHULPHQWV /LPQRO2FHDQRJU 'DUZLQ& -RXUQDORIWKH 5HVHDUFKHV LQWRWKH*HRORJ\ DQG 1DWXUDO +LVWRU\ RI 9DULRXV &RXQWULHV 9LVLWHG E\ WKH +06 %HDJOH +DIQHU3XEOLVKLQJ&R1HZ
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rochlorococcus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odularia spumigena 0HUWHQV&\DQREDFWHULD , )LHOGVWXGLHVRQWKHFRQWULEXWLRQRI EORRPV WR WKH QLWURJHQ EXGJHW RI WKH 3HHO+DUYH\ (VWXDU\ :HVWHUQ$XVWUDOLD +\GURELRORJLD +XDQJ 7& 7X - &KRZ 7- DQG &KHQ 7+ &LUFDGLDQ UK\WKPRIWKHSURNDU\RWHSynechococcus VS5) 3ODQW3K\VLRO ,WWXULDJD 5 DQG0LWFKHOO%* &KURFRFFRLGF\DQREDFWHULD $ VLJQLILFDQW FRPSRQHQW LQ WKH IRRG ZHE G\QDPLFV RI WKH RSHQ RFHDQ 0DU(FRO3URJU6HU -RKQVRQ3:DQG6LHEXUWK-0F1 &KURFRFFRLGF\DQREDFWHULD LQ WKH VHD D XELTXLWRXV DQG GLYHUVH SKRWRWURSKLF ELRPDVV /LPQRO2FHDQRJU .DKUX 0 /HSSlQHQ -0 DQG 5XG 2 &\DQREDFWHULDO EORRPVFDXVHKHDWLQJRIWKHVHDVXUIDFH 0DU(FRO3URJU6HU .DUO '0 /HWHOLHU 5 +HEHO '9 %LUG ') DQG :LQQ &' Trichodesmium EORRPV DQG QHZ QLWURJHQ LQ WKH 1RUWK 3DFLILF J\UH,Q &DUSHQWHU(-&DSRQH'*DQG5XHWHU-*HGV 0DULQH 3HODJLF&\DQREDFWHULD Trichodesmium DQG 2WKHU 'LD]RWURSKV SS .OXZHU'RUGUHFKW7KH1HWKHUODQGV .DUVWHQ* 'DVSK\WRSODQNWRQGHV DWODQWLVFKHQRFHDQVQDFK GHP PDWHULDO GHU GHXWVFKHQ 7LHIVHH([SHGLWLRQ :LVVHQVFKDIWHQ(UJHQ 'HXWVFKHQ 7LHIVHH([SHGLWLRQ 9DOGLYD .DUVWHQ * 'DV LQGLVFKH 3K\WRSODQNWRQ QDFK GHP 0DWHULDO GHU GHXWVFKHQ 7LHIVHH([SHGLWLRQ :LVVHQVFKDIWHQ
(UJHQ 'HXWVFKHQ 7LHIVHH([SHGLWLRQ 9DOGLYD .OHPHU$5DQG.RQRSND$( &DXVHVDQGFRQVHTXHQFHVRI EOXHJUHHQDOJDOF\DQREDFWHULDO EORRPV /DNH5HV0DQDJ .RQRQHQ..XSDULQHQ- 0lNH,l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y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uQHZv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
&KDSWHU 0DULQH 3ODQNWRQ 3DHUO+:%ODQG377XFNHU-DQG%ODFNZHOO- 7KHHIIHFWV RI VDOLQLW\ RQ WKH SRWHQWLDO RI D EOXHJUHHQ DOJDO Microcystis aeruginosa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richodesmium DJJUHJDWHV 5HODWLRQVKLSWR1 IL[DWLRQ SRWHQWLDO $SSO(QYLURQ 0LFURELRO 3DHUO +: DQG 3UXIHUW /( 2[\JHQSRRU PLFUR]RQHV DV SRWHQWLDO VLWHV RI PLFURELDO 1 IL[DWLRQ LQ QLWURJHQGHSOHWHG DHURELFPDULQHZDWHUV$SSO(QYLURQ0LFURELRO 3DHUO +: 3UXIHUW /( DQG $PEURVH :: &RQWHPSRUDQHRXV1 IL[DWLRQDQGR[\JHQLFSKRWRV\QWKHVLVLQWKH QRQKHWHURF\VWRXV PDWIRUPLQJ F\DQREDFWHULXP Lyngbya aestuarii. $SSO(QYLURQ0LFURELRO 3DHUO +: 3UXIHUW%HERXW/(DQG*XR& ,URQVWLPXODWHG 1 IL[DWLRQDQGJURZWKLQQDWXUDODQGFXOWXUHGSRSXODWLRQVRIWKH SODQNWRQLF PDULQH F\DQREDFWHULXP Trichodesmium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
3ODWW 7 6XEED 5DR '9 DQG ,UZLQ % 3KRWRV\QWKHVLV RI SLFRSODQNWRQLQWKHROLJRWURSKLFRFHDQ 1DWXUH 3RUWHU .* DQG 2UFXWW -' 1XWULWLRQDO DGHTXDF\ PDQDJHDELOLW\ DQG WR[LFLW\ DV IDFWRUV WKDW GHWHUPLQH WKH IRRG TXDOLW\ RI JUHHQ DQG EOXHJUHHQ DOJDH IRU Daphnia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ynechococcus SRSXODWLRQV IURP WKH 6DUJDVVR 6HD 0DU (FRO 3URJU6HU 3UXIHUW /( 3DHUO +: DQG /DVVHQ & *URZWK QLWURJHQ IL[DWLRQ DQG VSHFWUDO DWWHQXDWLRQ LQ FXOWLYDWHG Trichodesmium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richodesmium tenue: XOWUDVWUXFWXUHDQGVWRUDJHRIFDUERK\GUDWH SRO\SKRVSKDWHDQGQLWURJHQ -3K\FRO 5RVHQEHUJ * DQG 3DHUO +: 1LWURJHQ IL[DWLRQ E\ EOXH JUHHQDOJDHDVVRFLDWHGZLWKWKHVLSKRQRXVJUHHQVHDZHHGCodium decorticatum: (IIHFWVRQDPPRQLXPXSWDNH 0DU%LRO 5RZDQ . 3KRWRV\QWKHWLF 3LJPHQWV RI$OJDH &DPEULGJH 8QLYHUVLW\3UHVV1HZ
6RXPLD $ /HVF\DQRSK\FpHVGDQVOHSODQFWRQPDULQ $QQ %LRO 6WDO/-DQG%HUJPDQ% ,PPXQRORJLFDOFKDUDFWHUL]DWLRQRI QLWURJHQDVHLQWKHILODPHQWRXVQRQKHWHURF\VWRXVF\DQREDFWHULXP 2VFLOODWRULDOLPRVD 3DQWD 6WDO/-DQG.UXPEHLQ:( 1LWURJHQDVHDFWLYLW\LQWKHQRQ KHWHURF\VWRXV F\DQREDFWHULXP Oscillatoria VS JURZQ XQGHU DOWHUQDWLQJOLJKWGDUNF\FOHV $UFK0LFURELRO 6WHZDUW:'3 1LWURJHQIL[DWLRQ,Q&DUU1*DQG:KLWWRQ %$ HGV 7KH %LRORJ\ RI %OXH*UHHQ $OJDH SS %ODFNZHOO6FLHQWLILF3XEOLFDWLRQV2[IRUG 7DNHGD S, .DPDWDQL$DQG.DZDQREH. (IIHFWVRIQLWURJHQ DQG LURQ HQULFKPHQWV RQ SK\WRSODQNWRQ FRPPXQLWLHV LQ WKH QRUWKZHVW,QGLDQ2FHDQ 0DU&KHP 7D\ORU%)/HH&&DQG%XQW-6 1LWURJHQIL[DWLRQDVVRFLDWHG ZLWKWKHPDULQHEOXHJUHHQDOJD Trichodesmium DV PHDVXUHG E\ WKHDFHW\OHQHUHGXFWLRQWHFKQLTXH$UFK0LFURELRO 7D\ORU'/ 6\PELRVHV ,Q &DUSHQWHU(O DQG &DSRQH'* HGV 1LWURJHQ LQ WKH 0DULQH (QYLURQPHQW SS $FDGHPLF3UHVV1HZ
+ : 3DHUO &\DQREDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 'RUGUHFKW :XOII) 8QGHUVWDQGLQJWKH%DOWLF6HD 6\VWVHPVHFRORJ\LQ WKHRU\DQGSUDFWLFH,Q 'UHSSHU:6HG (FRG\QDPLFVSS 6SULQJHU1HZ
&KDSWHU )UHVKZDWHUEORRPV 5RGHULFN / 2OLYHU *HRUJH * *DQI &RRSHUDWLYH 5HVHDUFK &HQWUH IRU )UHVKZDWHU (FRORJ\ 0XUUD\ 'DUOLQJ )UHVKZDWHU 5HVHDUFK &HQWUH 32 %R[ $OEXU\ 16: $XVWUDOLD 'HSDUWPHQW RI (QYLURQPHQWDO %LRORJ\ 7KH 8QLYHUVLW\ RI$GHODLGH6$ $XVWUDOLD , ,QWURGXFWLRQ ,, %ORRP)RUPLQJ &\DQREDFWHULD
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hitton and M Potts (eds). The Ecology of Cyanobacteria. © Kluwer Academic Publishers Printed in the Netherlands
5/ 2OLYHU DQG** *DQI
;,,, ,QRUJDQLF &DUERQ ;,9 *UD]LQJ ;9 &RQFOXGLQJ 5HPDUNV
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³EORRP´ LV SRRUO\ GHILQHG EXW JHQHUDOO\ LW GHVFULEHV D SK\WRSODQNWRQ ELRPDVV VLJQLILFDQWO\ KLJKHUWKDQWKHODNH¶VDYHUDJH%\WKLVGHILQLWLRQHYHQ ROLJRWURSKLF ZDWHUV PD\ KDYH EORRPV DOWKRXJK WKLV VWUHWFKHV WKH FRQFHSW EH\RQG LWV JHQHUDO PHDQLQJ %ORRPV DUH XVXDOO\ FRPSULVHG RI RQO\ RQH RU WZR VSHFLHVDQGLGHQWLILHGE\WKHGRPLQDQWSK\WRSODQNWRQ W\SH HJ F\DQREDFWHULDO EORRP GLDWRP EORRP Anabaena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³ZDWHUEORRPV´ 5H\QROGV DQG:DOVE\ $ IHZQRQF\DQREDFWHULDOVSHFLHV FDQDOVRIRUPVXUIDFH EORRPV QRWDEO\ WKH JUHHQ DOJDBotryococcus braunii
ZKLFK EHFRPHV EXR\DQW E\ SURGXFLQJ DQG VWRULQJ RLOV 2FFDVLRQDOO\ IODJHOODWHV VXFK DV Euglena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
,, %ORRP)RUPLQJ &\DQREDFWHULD 7KHF\DQREDFWHULDSULQFLSDOO\UHVSRQVLEOHIRUIRUPLQJ EORRPV DUH JDVYDFXRODWH VSHFLHV 7KH\ DUH GLVWULEXWHG DFURVV D QXPEHU RI JHQHUD DQG YDU\ LQ IRUP DQG VL]H IURP VPDOO ILODPHQWV WR ODUJH JOREXODU FRORQLHV7DEOH 7KH ILODPHQWRXV IRUPV RFFXU DV VWUDLJKW VSLUDO RU WZLVWHG FKDLQV RI FHOOV DQG VRPHWLPHV WDNH RQ D VHFRQGDU\ PRUSKRORJ\ DV D UHVXOW RIWKH DJJUHJDWLRQ RU HQWDQJOHPHQW RI PDQ\ ILODPHQWV 5H\QROGV DQG :DOVE\ /HZLV $ FRQVHTXHQFH RIWKLV FRPSOH[ VHFRQGDU\ VWUXFWXUH LV WKH VL]H RI WKH ELRPDVV XQLW )LODPHQWRXV IRUPV GLIIHU VLJQLILFDQWO\
&KDSWHU )UHVKZDWHU%ORRPV
7DEOH 7KHPDMRUJHQHUDRIJDVYDFXRODWHSODQNWRQLFF\DQREDFWHULD
1IL[HU
)DPLO\
2UGHU
)LODPHQWRXV $QDEDHQD Anabaenopsis Aphanizomenon Nodularia Cylindrospermopsis Gloeotrichia Oscillatoria Spirulina
""
1RVWRFDFHDH 1RVWRFDOHV 1RVWRFDFHDH 1RVWRFDFHDH 1RVWRFDFHDH 1RVWRFDFHDH 5LYXODULDFHDH 2VFLOODWRULDFHDH2VFLOODWRULDOHV 2VFLOODWRULDFHDH
1RQILODPHQWRXV Microcystis Gomphosphaeria Coelosphaerium
&KURRFRFFDFHDH&KURRFRFRFFDOHV &KURRFRFFDFHDH &KURRFRFFDFHDH
LQ VL]H IURP VPDOO LQGLYLGXDO ILODPHQWV VXFK DV Anabaena minutissima P ZLGH DQG XS WR P ORQJ :DOVE\ HW DO RU Oscillatoria agardhii var. isothrix P ZLGH DQG XS WR P ORQJ 5H\QROGV D WR VSLUDOO\ FRLOHG ILODPHQWV DV LQ Anabaena circinalis WKDW PD\ EH P ORQJ DQG FDQ DJJUHJDWH WR IRUP PDFURVFRSLF FRORQLHV YLVLEOH WR WKH QDNHG H\H Aphanizomenon ILODPHQWV DJJUHJDWH LQWR UDIWV WKDW DUH UHPLQLVFHQW RI JUDVV EODGHV ZKLOH LQ Gloeotrichia WKH ILODPHQWV DUH FOXVWHUHG DURXQG D FHQWUDO QRGH WR IRUP ODUJH PP XUFKLQOLNH EDOOV 7KH FRORQLDO F\DQREDFWHULD DOVR YDU\ LQ VL]H DQG IRUP 7KH JOREXODU FRORQLHV XVXDOO\ DOWHU VL]H DV D UHVXOW RI JURZWK DQG UHGXFHG FRORQ\ GLVDJJUHJDWLRQ DOWKRXJK ORRVH DVVHPEODJHV RU DJJUHJDWLRQV RI VHSDUDWH FRORQLHV FDQ RFFXU 5H\QROGV HW DO Microcystis aeruginosa UDQJHV LQ VL]H IURP VLQJOH FHOOV RI P GLDPHWHU WKH IRUP PRVW RIWHQ IRXQG LQ FXOWXUH WR ODUJH JOREXODU RU VHPLVSKHULFDO FRORQLHV VHYHUDO PLOOLPHWUHV LQ GLDPHWHU FRQWDLQLQJ WHQV RI WKRXVDQGV RI FHOOV SHU FRORQ\ 5H\QROGV HW DO 7KLV UHSUHVHQWV D FKDQJH LQ XQLW YROXPH RI PRUH WKDQ WKUHH RUGHUV RI PDJQLWXGH DQG KDV LPSRUWDQW LPSOLFDWLRQV IRU D YDULHW\ RI SURFHVVHV LQFOXGLQJ EXR\DQF\ UHJXODWLRQ QXWULHQW XSWDNH JDV H[FKDQJH OLJKW LQWHUFHSWLRQ DQG VXVFHSWLELOLW\ WR JUD]LQJ 6XUIDFH EORRPV FDQ DSSHDU UDSLGO\ RIWHQ ZLWKLQ KRXUV DQG WR WKH FDVXDO REVHUYHU ZLWKRXW SULRU ZDUQLQJ RI WKH SUHVHQFH RI WKH RUJDQLVPV +LVWRULFDOO\ WKLV LPEXHG WKH EORRPV ZLWK D VRPHZKDW P\VWLFDO FKDUDFWHU DQG D QRWLRQ GHYHORSHG WKDW WKH F\DQREDFWHULD FRXOG JURZ H[WUHPHO\ UDSLGO\ ,Q IDFW WKH VXGGHQ DSSHDUDQFH UHVXOWV IURP WKH XSZDUG PLJUDWLRQ RI DQ H[LVWLQJ GLVSHUVHG SRSXODWLRQ
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icrocystis DQG LQ PHVRWURSKLF ODNHV IURP GLDWRPV WKURXJK FKU\VRSK\WHV DQG Sphaerocystis WR GLQRIODJHOODWHV *DVYDFXRODWH F\DQREDFWHULD DUH PRUHW\SLFDO RI HXWURSKLF ODNHV ,Q VKDOORZ ZHOOPL[HG HXWURSKLF ODNHV RI WKH WHPSHUDWH QRUWKHUQ KHPLVSKHUH WKH F\DQREDFWHULD WKDW GRPLQDWH GXULQJ VXPPHU DUH FRPPRQO\ VSHFLHV RI Oscillatoria. 7KHVH ODNHV DUH W\SLFDOO\ YHU\ WXUELG DQG ZKHQ ZLQWHUV DUH QRW WRR FROG GRPLQDQFH FDQ SHUVLVW WKURXJKRXWWKH\HDU 6FKHIIHU HW DO 6DV ,Q VRPH FOHDU ZDWHU WHPSHUDWH ODNHV ZKHUH OLJKW SHQHWUDWHV EH\RQG WKH GHSWK RI WKH HSLOLPQLRQ VSHFLHV VXFK DV Aphanizomenon flos-aquae DQG Oscillatoria agardhii IRUP PHWDOLPQHWLF SRSXODWLRQV RI VLQJOH ILODPHQWV ,Q &URRNHG /DNH ,QGLDQD SHDN FRQFHQWUDWLRQV RI JFKORURSK\OOD / RFFXUUHG DW D GHSWK RI FD P ZKHUH WKHUH ZHUH RSSRVLQJ
5/ 2OLYHUDQG** *DQI JUDGLHQWV RI LUUDGLDQFH DQG QXWULHQW DYDLODELOLW\ .RQRSND ,I FRQGLWLRQV EHFRPH XQVXLWDEOH WKHVH ILODPHQWV PD\ DJJUHJDWH WRJHWKHU DQG PRYH WRZDUGV WKH VXUIDFH DV ZDV REVHUYHG LQ /DNH *MHUVM¡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actylococcopsis VKRZHG JURZWK SXOVHV LQ KDELWDWV FKDUDFWHULVHG E\ KLJK QXWULHQWV DQG PL[LQJ DQG FRQVHTXHQWO\ UHODWLYHO\ ORZ OLJKW DYDLODELOLW\7XUEXOHQFHZDV UHGXFHG ZLWK WKH RQVHW RI WKHUPDO VWUDWLILFDWLRQ DQG WKH OLJKW FOLPDWH LPSURYHG ZKLOH QXWULHQW DYDLODELOLW\ GHFUHDVHG WKH FRPPXQLW\ EHFDPH GRPLQDWHG E\ JUHHQ DOJDH DV ZHOO DV WKH F\DQREDFWHULXP Aphanothece, ZKLFK LV FKDUDFWHULVHG E\ PDQ\ VPDOO FHOOV HPEHGGHG LQ D WKLFN JHODWLQRXV PDWUL[ $V QXWULHQWV EHFDPH OHVV DYDLODEOH DQG WKH OLJKW FOLPDWH UHPDLQHG IDYRXUDEOH F\DQREDFWHULD VKRZHG SXOVHV RI JURZWK DORQJVLGH WKH FKORURSK\WHV Sphaerocystis DQG Selenastrum. Gymnodinium, D GLQRIODJHOODWH ZDV IRXQG LQ DVVRFLDWLRQ ZLWK WKH ODUJHU F\DQREDFWHULD DQG RFFXSLHV WKH KDELWDW ZKHUH OLJKW DYDLODELOLW\ LV KLJK EXWQXWULHQWV DQG WXUEXOHQFH DUH PLQLPDO 7KLV VXFFHVVLRQDO VHTXHQFH PD\ EH LQWHUUXSWHG DW DQ\ WLPH E\ VWRUP HYHQWV RU RWKHU
&KDSWHU )UHVKZDWHU%ORRPV H[FHSWLRQDOFOLPDWLF FRQGLWLRQV ZKLFK HLWKHUUHVHWWKH VXFFHVVLRQDOFORFNRUFDXVHHOHPHQWVWREHVNLSSHG $WRIIVKRUHVWDWLRQVLQ/DNH9LFWRULDGXULQJSHULRGV RI WKHUPDO VWUDWLILFDWLRQ WKH F\DQREDFWHULD Anabaenopsis tanganyikae DQG Anabaena flos-aquae GRPLQDWHWKHFRPPXQLW\EXWJLYHZD\WRGLDWRPVHJ Melosira nyassensis YDU victoriae DV WKH ODNH EHFRPHV LVRWKHUPDO 7KH GHSWK GLVWULEXWLRQ GXULQJ SHULRGV RIWKHUPDO VWUDWLILFDWLRQ DOVR GLIIHUV EHWZHHQ VSHFLHV 'XULQJ SHULRGV RI UHODWLYHO\ VWURQJ VWUDWLILFDWLRQ Lyngbya circumcreta DQG Aphanocapsa elachista PD[LPD DUH FRQILQHG WR WKH WRS P WKH GHSWK RI WKH HXSKRWLF ]RQH LQ WKH RIIVKRUH VWDWLRQV ZDVFD P DQGGHFOLQHWRYHU\VPDOOQXPEHUV LQ WKH GHHSHU ZDWHU ,Q FRQWUDVW GLDWRP VSHFLHV DUH XVXDOO\ FRQILQHG WR RU EHORZ WKH ]RQH RI WKHUPDO GLVFRQWLQXLW\ $V WKHUPDO VWUDWLILFDWLRQ EUHDNV GRZQ ERWK F\DQREDFWHULD EHFRPH PRUH HYHQO\ GLVWULEXWHG ZLWKGHSWK DOWKRXJKWKHODUJHUFRORQLDOAphanocapsa elachista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icrocystis aeruginosa DQG M. flosaquae, Anabaenopsis VSS Aphanizomenon VS DQG Lyngbya VSS 6XUSULVLQJO\ WKH RWKHU PDMRU
F\DQREDFWHULDO FRPSRQHQW RI WKH SK\WRSODQNWRQ FRPPXQLW\ Anabaena flos-aquae, VKRZHG D SURQRXQFHG EXW XQH[SODLQHG VHDVRQDO SHULRGLFLW\ Microcystis VSS DSSHDUZHOOVXLWHGWRWKHGLHOSDWWHUQ RI VWUDWLILFDWLRQ DQG GHVWUDWLILFDWLRQ D WKHUPDO SHULRGLFLW\RIWHQIRXQGLQWKHVXUIDFHZDWHUVRIGHHSHU ODNHV RYHUO\LQJ D VXPPHU WKHUPRFOLQH 7KH UDSLG YHUWLFDOPLJUDWLRQ RIWKHODUJHFRORQLHV RIMicrocystis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icrocystis aeruginosa PD\ FRH[LVW ZLWK Anabaena flos-aquae, LW GRHV QRW FRH[LVW ZLWK Oscillatoria rubescens. 1HYHUWKHOHVV D XQLTXH FKDUDFWHULVWLF RI DOO EORRP IRUPLQJ VSHFLHV LV WKH SUHVHQFH RI JDV YHVLFOHV DQG WKHUH FDQ EH OLWWOH GRXEW WKDW WKH EXR\DQF\SURYLGHGE\WKHVHVWUXFWXUHVLV DVLJQLILFDQW DWWULEXWH
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nabaena ZKHUH WKH JDV YHVLFOH GHQVLW\ LV NJ P LI WKH LQWHUYHQLQJ VSDFH LV ILOOHG ZLWK ZDWHU DW D GHQVLW\ RI NJ P WKHQ WKH RYHUDOO JDV YDFXROH GHQVLW\ ZLOO EH NJP RQHIRXUWK WKH GHQVLW\ RI ZDWHU DQG DQ HIILFLHQW PHFKDQLVP WR SURYLGH OLIW :DOVE\
5/ 2OLYHU DQG ** *DQI 7KH FRPPRQ SHUFHSWLRQ WKDW JDV YHVLFOHV DUH HTXLYDOHQWWREDOORRQV RU EXEEOHV LQ WKH F\WRSODVPLV LQFRUUHFW :DOVE\ KDV VXJJHVWHG D PRUH DSSURSULDWHDQDORJ\ ZRXOGEH DSDLURIROG IDVKLRQHG HDUWKHQZDUH IORZHUSRWV FRDWHG ZLWKRLO RQ WKH LQQHU VXUIDFH WR UHSUHVHQW WKH K\GURSKRELF LQQHU OD\HU RI WKH JDV YHVLFOH DQG JOXHG WRJHWKHU DW WKH ULPV *DV YHVLFOHVDVZLWKWKHIORZHUSRWV DUHSHUPHDEOHWRJDV DQGWKHUHIRUHFRQWDLQ DLU DWDWPRVSKHULFSUHVVXUH EXW WKHDLULVQRWUHTXLUHGWRPDLQWDLQWKHKROORZVSDFHDV WKH JDV YHVLFOH ZDOOV DUH ULJLG ,I WKH JDV YHVLFOH DQDORJ\ LVVXEPHUJHGZDWHUZLOOVHHSLQWRWKHZDOO EXWZLOOEHSUHYHQWHGIURPHQWHULQJWKHYHVLFOHFDYLW\ E\ WKH K\GURSKRELF OD\HU $LU ZLOO GLIIXVH EDFN DQG IRUWKEHWZHHQWKHFDYLW\DQGWKHVXUURXQGLQJPHGLXP DQG WKH YHVLFOH ZLOO EH ILOOHG ZLWK DLU RIFRPSRVLWLRQ VLPLODUWRWKDWLQWKHVXUURXQGLQJOLTXLG
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
&KDSWHU )UHVKZDWHU%ORRPV
Fig. 1 3UHVVXUHV DFWLQJ RQ JDV YHVLFOHV *DV YHVLFOHV PD\FROODSVH LQ UHVSRQVH WR LQFUHDVHG K\GURVWDWLF SUHVVXUHJHQHUDWHG E\ GHSWK DQG LQFUHDVHGWXUJRU SUHVVXUHLQUHVSRQVHWROLJKW:LGHU ZHDNHUYHVLFOHVFROODSVHPRUHUHDGLO\WKDQWKLQQHUZHDNHURQHV .H\ SK K\GURVWDWLFSUHVVXUHSW WXUJRUSUHVVXUHSI SUHVVXUHGXHWRRYHUO\LQJDWPRVSKHUHSJ SUHVVXUHRIJDVLQYHVLFOH
& *DV9HVLFOH&ROODSVH 7KH FULWLFDO SUHVVXUH SF DW ZKLFK RI JDV YHVLFOHV DUH FROODSVHG YDULHV ZLGHO\ EHWZHHQ JDV YDFXRODWHRUJDQLVPVZLWKLQWKHUDQJH WR03D ,Q WKH EORRPIRUPLQJ IUHVKZDWHU F\DQREDFWHULD WKH UDQJH LV OHVV FD WR 03D EXW GLIIHUHQW VSHFLHV VWLOO KDYH SDUWLFXODU OLPLWV ,Q JDV YHVLFOHV LVRODWHG IURP Anabaena flos-aquae WKH FULWLFDO FROODSVH SUHVVXUHV UDQJHG IURP WR 03D :DOVE\ ZKLOH IRU QDWXUDO SRSXODWLRQV RI Microcystis aeruginosa IR aeruginosa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richodesmium thiebautii LQKDELWV WKH GHHS RFHDQV ZKHUH K\GURVWDWLF SUHVVXUH FDQ EH FRQVLGHUDEOH DQG KDV D JDV YHVLFOH ZLGWK RIRQO\ QP *DQWW HW DO ZKLOH Oscillatoria agardhii, ZKLFK LQKDELWV GHHS ODNHV KDV D JDV YHVLFOH ZLGWK RI QP DQG Microcystis aeruginosa, Anabaena flos-aquae DQG Aphanizomenon flos-aquae, ZKLFK DUH IRXQG SULQFLSDOO\ LQ VKDOORZ ODNHV RU LQ ODNHV ZKHUH WKH PL[LQJGHSWKLVUHGXFHGE\WKHUPDOVWUDWLILFDWLRQKDYH JDV YHVLFOH GLDPHWHUV RI DQG QP UHVSHFWLYHO\ :DOVE\ DQG %OH\WKLQJ Dactylococcopsis salina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
5/ 2OLYHUDQG** *DQI UHTXLUHG IRU D ORQJHU JDV YHVLFOH VLQFH LW ZRXOG QRW PXFKLPSURYHWKHSURYLVLRQRIEXR\DQF\
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¶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
&KDSWHU )UHVKZDWHU%ORRPV
Fig. 2 'LDJUDPPDWLF EDODQFH VKHHW GHPRQVWUDWLQJ FKDQJHV LQ FHOO GHQVLW\ RIAnabaena flos-aquae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phanizomenon flos-aquae WKHJDV YHVLFOH FRQWHQWSHUXQLWSURWHLQ ZDV UHJXODWHG E\ WKH VXSSO\ RI HQHUJ\ VR WKDW FHOOV ZHUH QRQ EXR\DQW DW DOO JURZWK UDWHV H[FHHGLQJ WZHQW\ SHUFHQW RIWKH PD[LPXP JURZWK UDWH .RQRSND HW DO .URPNDPS HW DO 6LPLODUO\ ZKHQ Oscillatoria agardhii ZDV VKLIWHG IURPORZWR KLJKHU
OLJKW LQWHQVLWLHV WKH UHVXOWLQJ EXR\DQF\ ORVV ZDV H[SODLQHGE\DGLOXWLRQ RIJDVYHVLFOHV DVFHOOYROXPH LQFUHDVHG EXW ZLWKRXW D PDWFKLQJ LQFUHDVH LQ JDV YHVLFOH V\QWKHVLV 8WNLOHQ HW DO ,Q Pseudanabaena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icrocystis aeruginosa UHVSRQGHGWRDQLQFUHDVHLQWKHOLPLWLQJHQHUJ\VXSSO\ E\ LQFUHDVLQJ JDV YHVLFOH SURWHLQ V\QWKHVLV LQ
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
5/ 2OLYHU DQG ** *DQI EDODQFHG WKHQ WKH UHODWLYH JURZWK UDWH JLYHQ E\ PD[ LVKLJKHYHQLIWKHVSHFLILFJURZWKUDWHLVORZ ,I JURZWK LV UHVWULFWHG E\ D OLPLWLQJ QXWULHQW WKHQ HQHUJ\ FDSWXUH H[FHHGV WKDW XWLOLVHG LQ WKH QXWULHQW UHVWULFWHGJURZWK $VOLJKWVXSSO\LV JUHDWHUWKDQWKDW UHTXLUHG WR PDLQWDLQ WKH 5*5PD[ FDUERK\GUDWH LV VWRUHG)LJ .RQRSNDDQG6FKQXU REWDLQHG FDUERK\GUDWH WR SURWHLQ UDWLRV IRXU WR VHYHQ WLPHV KLJKHU LQ FXOWXUHV OLPLWHG E\ QLWURJHQ SKRVSKRUXV RU VXOSKXU WKDQ LQ QRQOLPLWHG FXOWXUHV RU WKRVH OLPLWHG E\ FDUERQ ,Q JHQHUDO EXR\DQF\ GHFUHDVHV ZKHQ PDMRU QXWULHQWV VXFK DV SKRVSKRUXV RU QLWURJHQ OLPLW FHOO JURZWK EHFDXVH FDUERK\GUDWH DFFXPXODWHV EXW DVVRFLDWHGWXUJRUSUHVVXUH LQFUHDVHV FDQ DOVR FROODSVH JDV YHVLFOHV HVSHFLDOO\ LI WKHUH LV DQ DFFRPSDQ\LQJ ULVH LQ K\GURVWDWLF SUHVVXUH GXH WR FHOO VHGLPHQWDWLRQ 5H\QROGV DQG :DOVE\ .OHPHU .RQRSND :DOVE\ 7KH GHJUHH RI JDV YDFXRODWLRQ PD\ DOVR EH UHGXFHG WKURXJK PROHFXODU FRQWUROV RQ JDV YHVLFOHSURGXFWLRQ EXWWKLV LV VSHFLHV VSHFLILF66 DVVKRZQLQ)LJ ,IQXWULHQWOLPLWDWLRQ JUHDWO\ GHSUHVVHV JURZWK UDWH WKHQ FDUERK\GUDWH DFFXPXODWLRQ DQG EXR\DQF\ ORVV FDQ RFFXU HYHQ DW ORZ OLJKW LQWHQVLWLHV 8QGHU VHYHUH DQG VXVWDLQHG QLWURJHQ OLPLWDWLRQ UHGXFHG JDV YHVLFOH V\QWKHVLV UHVXOWVLQUHGXFHGEXR\DQF\)LJ :KHQDOOQXWULHQWVLQFOXGLQJ FDUERQ DUHSUHVHQWLQ DEXQGDQFH WKHQ WKH EXR\DQF\ RI WKH RUJDQLVPV LV ODUJHO\ D IXQFWLRQ RIWKH LUUDGLDQFH LQWHQVLW\ UHODWLYH WRWKHJURZWKUHTXLUHPHQWVRIWKHFHOOV)LJ ,IWKH LUUDGLDQFH FDSWXUHG LV OHVV WKDQ WKDW UHTXLUHG WR DFKLHYH WKH PD[LPXP JURZWK UDWH XQGHU WKH SUHYDLOLQJ HQYLURQPHQWDO FRQGLWLRQV WKHQ HQHUJ\ VXSSO\ ZLOO EH ORZUHODWLYH WR ZKDW FRXOG EH XWLOLVHG DQG FDUERK\GUDWH UHVHUYHV ZLOO EH UHGXFHG DQG FHOO EXR\DQF\ LQFUHDVHG ,Q RUJDQLVPV OLNH Aphanizomenon flos-aquae, ZKHUH WKH GHJUHH RI JDV YDFXRODWLRQ LV D IXQFWLRQ RI WKH OLPLWLQJ HQHUJ\ VXSSO\ PROHFXODUSURFHVVHV ZLOO LQFUHDVH JDV YHVLFOH V\QWKHVLVDQGHQKDQFHWKHSRVLWLYHEXR\DQF\UHVSRQVH 8WNLOHQ HW DO .RQRSND HW DO .URPNDPS HWDO 'DPHUYDO HW DO $V GLVFXVVHG HDUOLHU IRU Microcystis aeruginosa, QRW DOO RUJDQLVPV QHFHVVDULO\ VKRZ WKLV PROHFXODU UHVSRQVH WR OLJKWFKDQJHV DQG LQ)LJ LW LV GHSLFWHG DV EHLQJ VSHFLHVVSHFLILF66 ,ILUUDGLDQFHV DUH VXIILFLHQW WR VXSSRUW JURZWK UDWHV FORVH WR PD[LPXP XQGHU QXWULHQW VXIILFLHQW FRQGLWLRQV WKHQ FHOOXODUFRPSRVLWLRQ ZLOO DSSURDFK D PHDQ HOHPHQWDO UDWLR VLPLODU WR WKH 5HGILHOG UDWLR & 1 3 E\ DWRPV LQGLFDWLYH RI FHOOV JURZLQJ ZLWKRXW QXWULHQW OLPLWDWLRQ +HFN\ DQG
&KDSWHU )UHVKZDWHU%ORRPV
Fig. 3 )ORZFKDUWRIWKHPDMRUIDFWRUVLQIOXHQFLQJJDVYHVLFOHSURGXFWLRQDQGEXR\DQF\UHJXODWLRQ .H\ &L LQRUJDQLF FDUERQ &+2 FDUERK\GUDWH JY JDV YHVLFOH 1 QLWURJHQ 3 SKRVSKRUXV 5*5PD[ PD[LPXP UHODWLYH JURZWK UDWH 66 VSHFLHV VSHFLILF LQGLFDWLQJ WKH UHVSRQVH LV QRW XQLYHUVDO % FKDQJH LQ EXR\DQF\ ZKLFK FDQ EH SRVLWLYH QHJDWLYH RU HLWKHU GHSHQGLQJRQVSHFLHV
.LOKDP +HFN\ HW DO 7KH EXR\DQF\ VWDWXVRIFHOOVXQGHUWKHVHEDODQFHGJURZWKFRQGLWLRQV LVVSHFLHVVSHFLILF)LJ ZLWKAphanizomenon flosaquae EHLQJ QRQEXR\DQW .RQRSND HW DO .URPNDPS HW DO DQG Microcystis aeruginosa EXR\DQW .URPNDPS HW DO 3UHVXPDEO\ WKH EXR\DQF\ VWDWXV LV VHW E\ PROHFXODU FRQWUROV RQ FHOOXODUVWUXFWXUH $W LUUDGLDQFHV DERYH WKRVH VDWXUDWLQJ QXWULHQW VXIILFLHQW FHOO JURZWK WKH UHVSRQVHV DUH VLPLODU WR WKRVH IRXQG XQGHU QXWULHQW OLPLWDWLRQ 7KH FDUERK\GUDWH VWRUH LQFUHDVHV ZLWK LUUDGLDQFH XS WR D PD[LPXP ZKHQ SKRWRV\QWKHVLV LV VDWXUDWHG 7KLV UHVXOWV LQ DQ HQODUJHG FDUERK\GUDWH VWRUH ZKLFK LQFUHDVHV FHOO GHQVLW\ ZKLOH PROHFXODU FRQWUROV LQ VRPH RUJDQLVPV UHGXFH WKH UDWH RI JDV YHVLFOH V\QWKHVLV )LJ ,I WXUJRU SUHVVXUH LQFUHDVHV VXIILFLHQWO\ WKHQ JDV YHVLFOH FROODSVH &DQ RFFXU DOWKRXJKWKLVPD\QRWEHVXIILFLHQWRQLWVRZQWR
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
9 0L[LQJ 5HJLPHVDQG &\DQREDFWHULD %XR\DQF\ PDLQWDLQV F\DQREDFWHULDO FHOOV LQ VXVSHQVLRQZKLOHLWVUHJXODWLRQHQDEOHVWKHPWRPRYH YHUWLFDOO\WKURXJKWKHZDWHULQUHVSRQVHWRFKDQJLQJ
5/ 2OLYHU DQG** *DQI
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otryococcus WKDW FDQ EHFRPH EXR\DQWDIWHUSURGXFLQJDQGDFFXPXODWLQJRLOV
$ )ORDWLQJ DQG 6LQNLQJ XQGHU 4XLHVFHQW DQG 7XUEXOHQW0L[LQJ &RQGLWLRQV ,Q TXLHVFHQW ZDWHUV WKH VLQNLQJ RU IORDWLQJ UDWH RI SK\WRSODQNWRQ FDQ EH FDOFXODWHG IURP WKH 6WRNHV HTXDWLRQ PRGLILHG LIQHFHVVDU\ E\ WKH LQFOXVLRQ RID ³IRUPIDFWRU´ WKDWDGMXVWVIRUWKH QRQVSKHULFDO VKDSH RIVRPHRUJDQLVPV v= gr 2 S S
>@ ZKHUH WKH WHUPLQDO YHORFLW\ RI WKH RUJDQLVP Y LV GHSHQGHQWRQJUDYLWDWLRQDODFFHOHUDWLRQJ WKHVL]H RI WKH RUJDQLVP UHSUHVHQWHG DV WKH UDGLXV U RI D VSKHUHRIHTXDOYROXPHWKHGHQVLW\RIWKHRUJDQLVP S¶ DQGWKHGHQVLW\S DQGWKHYLVFRVLW\ RIWKH PHGLXP 7KH IRUPUHVLVWDQFHIDFWRU LV GHILQHG DV YYV ZKHUHYV LVWKH WHUPLQDO YHORFLW\RIDVSKHUH RI HTXDOYROXPHDQGGHQVLW\WRWKDWRIWKHRUJDQLVP )RU VRPH VKDSHV WKH IRUP IDFWRU LV NQRZQ IURP HPSLULFDOUHODWLRQVKLSV 0F1RZQDQG0DODLND 'DYH\ DQG :DOVE\ EXWJHQHUDOO\ LWKDV WR EH GHWHUPLQHGDIWHUPHDVXULQJDOORWKHUYDULDEOHV 2OLYHU HW DO 7KH 6WRNHV HTXDWLRQ LV VXLWDEOH IRU FDOFXODWLQJ VLQNLQJ DQG IORDWLQJ YHORFLWLHV LI WKH DVVXPSWLRQ RIODPLQDU IORZ LV QRW YLRODWHG DQG WKLV ZLOO EH WKH FDVH SURYLGHG WKH SDUWLFOH5H\QROGV QXPEHU GRHV QRW H[FHHG :DOVE\ DQG 5H\QROGV 5H\QROGV $V D FRQVHTXHQFH WKH HTXDWLRQ ZLOO QRW EH UHOLDEOH IRU ODUJH SK\WRSODQNWRQ FRORQLHV ZKHUH U!P 5H\QROGV 7KH 6WRNHV IXQFWLRQ H[SOLFLWO\ LGHQWLILHV PDMRU IHDWXUHVWKDWLQIOXHQFHWKHIORDWLQJDQG VLQNLQJUDWHRI
Fig. 4 $ FRPSDULVRQ RI IORDWLQJ DQG VLQNLQJ YHORFLWLHV Y RI VHOHFWHG F\DQREDFWHULD ZLWK WKH FKDUDFWHULVWLF YHORFLW\ RI WXUEXOHQFH X 7KH VROLG OLQH VHSDUDWHV UHJLRQV RI SDUWLFOH HQWUDLQPHQWDQG GLVHQWUDLQPHQWDV GHILQHGE\(TQ 9DOXHV RIX LQGLIIHUHQWKDELWDWVSURYLGHSHUVSHFWLYH DIWHU5H\QROGV D
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
&KDSWHU )UHVKZDWHU%ORRPV 7KH FKDUDFWHULVWLF YHORFLW\ RI WXUEXOHQFH FDQ EH HQYLVDJHG HLWKHU DV WKHURRWPHDQ VTXDUH RIWKHWLPH DYHUDJHG YHUWLFDO YHORFLW\ IOXFWXDWLRQV X X ! ò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
SHQHWUDWLRQKHDW FDSWXUH DQG WKH GHJUHH RI PL[LQJ 7KH GHSWK RI WKH PL[HG OD\HU ZLOO WHQG WR D SRLQW ZKHUH WKH EXR\DQW HQHUJ\ RI WKHVXUIDFHOD\HU DQGWKH NLQHWLF HQHUJ\ RI WKHVXUIDFHZLQG VWUHVVDUHEDODQFHG 7KH LQVWDQWDQHRXV EDODQFH EHWZHHQ WKHVH WZR SURFHVVHV LV GHVFULEHG E\ WKH :HGGHUEXUQ QXPEHU ,PEHUJHU DQG +DPEOLQ ZKLFK LV WKH UDWLR RI WKH HQHUJ\ SHU VXUIDFH DUHD IURP EXR\DQF\ WR WKDW JHQHUDWHG E\ WKHVKHDUIRUFHV w=gh
L
>@ ,Q WKLV H[SUHVVLRQ Z LV WKH GHQVLW\ GLIIHUHQFH EHWZHHQ WKH ZDWHU EHORZ WKH PL[HG OD\HU DQG WKH ZDWHULQWKHPL[HGOD\HU/LVWKHOHQJWKRIWKHODNH DWWKHEDVHRIWKHPL[HGOD\HULQWKHGLUHFWLRQRIWKH ZLQG DQGKLV WKHGHSWK RIWKHGLXUQDO WKHUPRFOLQH 6SLJHODQG,PEHUJHU :KHQ :! WKH VWUXFWXUH LV UREXVW DQG UHVLVWDQW WR IXUWKHUGHHSHQLQJXQOHVVWKHUHLV DVXEVWDQWLDO FKDQJH LQ Z RU X ZKHUHDV LI : WKHQ WKH PL[HG OD\HU GHHSHQV UDSLGO\ XQWLO K DQG Z DUH ODUJH HQRXJK WR PDNH :! 6SLJHO DQG ,PEHUJHU 5H\QROGV D (VWLPDWHV RI WKH GHSWK RI PL[LQJ =P KDYH EHHQ REWDLQHG IURP WKLV IXQFWLRQ E\ VHWWLQJ : 5H\QROGV D ,EHOLQJV HW DO D 0RUH UHFHQWO\ WKH VXLWDELOLW\ RI HTXDWLRQ WR SUHGLFW WKH GHSWKRIWKHPL[HG OD\HUKDV EHHQ TXHVWLRQHG VLQFH LW PRUH FRUUHFWO\ HVWLPDWHV ZKHWKHU RU QRW WKH S\FQRFOLQH ZLOO XQGHUJR IXOO RUSDUWLDO XSZHOOLQJ DQG ZKHWKHU ERXQGDU\ PL[LQJ LV OLNHO\ 0DF,QW\UH SHUV &RP KDV VXJJHVWHGWKDWWKHUHJLRQV ZKHUHWKH ZDWHU FROXPQ LV PL[LQJ DQG WKH PL[LQJ LQWHQVLW\ FDQ EH REWDLQHG PRUH DSSURSULDWHO\ IURP PRGHOOLQJ RU IURP PLFURVWUXFWXUHSURILOLQJ 7KH UDWLR EHWZHHQ WKH GHSWK WR ZKLFK RIOLJKW SHQHWUDWHV HXSKRWLF GHSWK =HX DQG WKH GHSWK RI PL[LQJ LV DQ LPSRUWDQW LQGLFDWRU RIWKH OLJKW UHJLPH HQFRXQWHUHG E\ FHOOV FDSWXUHG LQ WKH ZDWHU PRWLRQ $VWKH =HX=P UDWLR GHFOLQHV EHORZ D YDOXH RIRQHWKH SURSRUWLRQ RI WLPH WKDW WKH FHOOV VSHQG LQ WKH OLJKW GHFUHDVHV XQWLO HYHQWXDOO\ FHOOV HQFRXQWHU LQVXIILFLHQW OLJKW WR JURZ&RQVHTXHQWO\ LQVKDOORZWXUELG ZDWHUV WKDW DUH XQLIRUPO\ DQG FRQWLQXDOO\ PL[HG OLJKW PD\ OLPLW SRSXODWLRQ JURZWK DV DW DQ\ RQH WLPH RQO\ D VPDOOSURSRUWLRQRIWKHSRSXODWLRQFDQ JDLQ DFFHVVWR WKHHXSKRWLF]RQH&RQYHUVHO\ LQFOHDUGHHSODNHVWKH RQVHW RI WKHUPDO VWUDWLILFDWLRQ LQFUHDVHV WKH PHDQ LUUDGLDQFHDVWKHPL[LQJGHSWKGHFUHDVHV ,IWKH =HX]P UDWLR LV JUHDWHUWKDQ RQH LQ DWKHUPDOO\ VWUDWLILHG ODNH WKHQFHOOVPRYLQJZLWKWKHZDWHUDUHDOZD\VLQWKH
5/ 2OLYHUDQG ** *DQI
Fig. 5 &KDQJHV LQ WKH VXUIDFH FXUUHQW VSHHG FV DV DIXQFWLRQ RI ZLQGVSHHG8D
LOOXPLQDWHG]RQHDQGOLJKWSHQHWUDWHVEHORZWKHPL[HG OD\HULQWRWKHWKHUPRFOLQH
& 6LQNLQJ2UJDQLVPV ,Q FRQVLGHULQJ WKH HIIHFW RI WXUEXOHQW PL[LQJ RQ WKH GLVWULEXWLRQ RI VLQNLQJ RUJDQLVPV 5H\QROGV FDUULHGRXWILHOGH[SHULPHQWVPHDVXULQJWKHVHWWOLQJRI /\FRSRGLXP VSRUHV LQ ODUJH ODNH HQFORVXUHV 7KH UHVXOWV GHPRQVWUDWHG WKDW WKH PHDVXUHG effective VLQNLQJ UDWH RIWKH VSRUHV XQGHU WXUEXOHQW FRQGLWLRQV ZDV VLJQLILFDQWO\ OHVV WKDQ WKH intrinsic VLQNLQJ UDWH PHDVXUHG LQ TXLHVFHQW ZDWHU FROXPQV LQ WKH ODERUDWRU\ 5H\QROGV SURSRVHG D PL[LQJ PRGHO WKDW HQYLVDJHG DZDWHU FROXPQ WKDWZDV SHULRGLFDOO\ IXOO\ PL[HG EXW ZLWK D WUDSSLQJ ]RQH DW WKH ERWWRP IURP ZKLFK FHOOV FRXOG QRW EH UHVXVSHQGHG 6PLWK GHYHORSHGDVLPSOHPRGHOWRIRUPDOLVHWKHVHFRQFHSWV DQG GHULYHG D JHQHUDO HTXDWLRQ WR FDOFXODWH WKH QXPEHURISDUWLFOHVUHPDLQLQJLQVXVSHQVLRQ1W DIWHU WLPH W IRU FRQGLWLRQV ZKHUH RUJDQLVPV DUH IXOO\ HQWUDLQHGLQWKHWXUEXOHQFH Nt = No e
-t/t1
>@ +HUH 1R LV WKH RULJLQDO QXPEHU RI SDUWLFOHV LQ VXVSHQVLRQDQGW LVWKHFROXPQFOHDUDQFHWLPH ZKLFK IRU SDUWLFOHV ZLWK D VLQNLQJ YHORFLW\ 9 LQ D ZDWHU FROXPQRIKHLJKW+LVW +9 8QGHUWXUEXOHQWFRQGLWLRQVDQLQILQLWHWLPHLVUHTXLUHG WRFRPSOHWHO\FOHDUWKHZDWHUFROXPQ RISDUWLFOHV DQG DVPDOO QXPEHURIFHOOV PD\UHPDLQ LQ VXVSHQVLRQWR VHHG VXEVHTXHQW JURZWK SKDVHV 6PLWK 5HSODFLQJW ZLWK+9LQ(TQSURYLGHVDPHDQVIRU PHDVXULQJWKHLQWULQVLF VLQNLQJRUIORDWLQJYHORFLW\RI SDUWLFOHV LQ D IXOO\ WXUEXOHQW V\VWHP 6PLWK
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scillatoria redekei WKDW DUH UHVWULFWHG LQ RFFXUUHQFH WR WKH HQGV RIWKH FHOOV DQG GR QRW DSSHDU WR FRQIHU SRVLWLYHEXR\DQF\ 7KH ORVV RI VHGLPHQWLQJ SDUWLFOHV GXH WR WKH SUHVHQFHRIWUDSSLQJ]RQHV DWWKHORZHUERXQGDU\RID PL[HG OD\HU RFFXUV DW DOO LQWHQVLWLHV RI PL[LQJ DQG UHSUHVHQWV D FRQWLQXDO ORVV WR QRQEXR\DQW RU QRQ PRWLOH SK\WRSODQNWRQ SRSXODWLRQV ,Q QDWXUDO ZDWHUV SDUWLFOHV ZLOO EH UHPRYHG IURP WKH PL[LQJ ]RQH LI WKH\ VHGLPHQW LQWR WKH WKHUPRFOLQH ZKHUH WXUEXOHQFH
Fig. 6 3HUFHQW RIWKH SRSXODWLRQ UHWDLQHG ZLWKLQ D P IXOO\ PL[HGZDWHUFROXPQ 7KHWKUHHFXUYHVUHSUHVHQWWKUHHVLQNLQJ YHORFLWLHV XSSHU PLGGOH DQG P K ERWWRP RYHUK
LV VPDOO RU VHWWOH RQWR WKH VXEVWUDWXP ZKHUH ZDWHU YHORFLWLHV DSSURDFK ]HUR %XR\DQWF\DQREDFWHULDFDQ DYRLG WKLV ORVV HQWLUHO\ DQG VR EXR\DQF\ DORQH ZLWKRXW WKH QHHG IRU LWV SUHFLVH UHJXODWLRQ FDQ SURYLGH WKH F\DQREDFWHULD ZLWK D VLJQLILFDQW DGYDQWDJH RYHU VHGLPHQWLQJ SK\WRSODQNWRQ )XUWKHUPRUH LIDGHHS PL[LQJ HSLVRGH PRYHV SDUW RI WKH SK\WRSODQNWRQ FRPPXQLW\ WR D GHSWK EHORZ WKH OHYHO RI WKH WKHUPRFOLQH WKHQ RQFH VWUDWLILFDWLRQ
&KDSWHU )UHVKZDWHU%ORRPV UHIRUPVEXR\DQWFHOOVFDQPRYH EDFNLQWR WKHVXUIDFH PL[HG OD\HU ZKLOH VHGLPHQWLQJ FHOOV FDQQRW +XPSKULHV DQG /\QH
' (QWUDLQPHQW DQG 9HUWLFDO 'LVWULEXWLRQV 0L[LQJWKDWIXOO\HQWUDLQVDSK\WRSODQNWRQ SRSXODWLRQ ZLOO KRPRJHQLVH LWV YHUWLFDO GLVWULEXWLRQ EXW WKH GHJUHH RI HQWUDLQPHQW YDULHV GHSHQGLQJ RQ WKH LQWULQVLF YHORFLW\ RI WKH SK\WRSODQNWRQ UHODWLYHWRWKH YHORFLW\ RI WKH WXUEXOHQW PRWLRQ $ IXQFWLRQ GHVFULELQJ WKLV LQWHUDFWLRQ DQGXVHGWRGLVWLQJXLVKWKH H[WHQW RI HQWHUWDLQPHQW y ZDV SURSRVHG E\ +XPSKULHV DQG ,PEHUJHU y _Y_X
>@ ZKHUH _Y_ LVWKHPRGXOXVLH WKHGLUHFWLRQDOVLJQKDVEHHQ UHPRYHG RIWKH VLQNLQJ RUIORDWLQJ YHORFLW\ RIWKH SK\WRSODQNWRQ XQLW DQG X LV WKH YHORFLW\ RI WKH WXUEXOHQWHGGLHV WKDWWUDQVSRUWWKH FHOOV WKURXJK WKH PL[HGOD\HU+XPSKULHVDQG/\QH 7KLV IRUPXODWLRQ GLIIHUV IURP VRPH SXEOLVKHG YHUVLRQV ZKHUH WKH HTXDWLRQ KDV EHHQ LQFRUUHFWO\ WUDQVFULEHG +XPSKULHV DQG ,PEHUJHU 5H\QROGVD 7KHVL]HRIXFDQEHHVWLPDWHGE\WKHVKHDUYHORFLW\ RIX VHHSUHYLRXVVHFWLRQ :KHQy WKHQ ZDWHU YHORFLWLHVDUHVXIILFLHQWO\ODUJHUHODWLYHWRWKHLQWULQVLF VLQNLQJ RU IORDWLQJ YHORFLWLHV RIWKH SK\WRSODQNWRQ WR HQWUDLQ WKH FHOOV ZLWKLQ WKH ZDWHU PRWLRQ VR WKDW WXUEXOHQFH KRPRJHQLVHV WKH YHUWLFDO GLVWULEXWLRQ RI WKHSRSXODWLRQDVHQYLVDJHGLQ(TQ+XPSKULHVDQG ,PEHUJHU +XPSKULHV DQG/\QH :KHQ y! WKHQWKHLQWULQVLF YHORFLW\ RIWKHSK\WRSODQNWRQ SOD\VDQLQFUHDVLQJUROHLQWKHSRSXODWLRQGLVWULEXWLRQ 7KLVEHJLQVWRRFFXUZKHQWKHWXUEXOHQWYHORFLW\IDOOV EHORZ WLPHVWKHSK\WRSODQNWRQYHORFLW\ ,Q)LJ WKHIORDWLQJRUVLQNLQJVSHHGRIVHOHFWHGRUJDQLVPV LV FRPSDUHG ZLWK WKH PL[LQJ YHORFLW\ XVLQJ HTXDWLRQ WR GLVWLQJXLVK WKH GHJUHH RI HQWUDLQPHQW (VWLPDWHG YDOXHV RI X LQ GLIIHUHQW KDELWDWV DUH LQGLFDWHG IRU FRPSDULVRQ 5H\QROGV D 6LQJOH ILODPHQWV RI Oscillatoria ZLWK VORZ IORDWLQJ UDWHV DUH HQWUDLQHG DW DOO X YDOXHV JUHDWHU WKDQ WKRVH IRXQG LQ WKH PHWDOLPQLRQ ,Q FRQWUDVW ODUJH FRORQLHV RI Microcystis FDQ FRQWLQXH WR XWLOLVH EXR\DQF\ WR DOWHU WKHLUYHUWLFDOGLVWULEXWLRQZLWKLQWKHPL[LQJ]RQHXSWR XYDOXHVW\SLFDO RIUHDVRQDEO\ZHOOPL[HG OD\HUV $W X YDOXHV LQ H[FHVV RI P V DOO IRUPV RI
F\DQREDFWHULD DUH HQWUDLQHG LQ WKH ZDWHU PRWLRQ DQG ZLOOKDYHXQLIRUPYHUWLFDOGLVWULEXWLRQV )LHOG GDWD WR WHVW FULWLFDOO\ WKLV UHODWLRQVKLS DUH VFDUFH EXW LQ /DNH 9LQNHYHHQ LQ 7KH 1HWKHUODQGV ,EHOLQJV HW DO D IRXQG OHVV IORDWLQJ FRORQLHV RI WKH JDVYDFXRODWH F\DQREDFWHULXP Microcystis QHDUWKH VXUIDFHWKDQ DWWKHERWWRPRIWKH PL[HG OD\HU ZKHQWKH VKHDUYHORFLW\XZDV RQO\ILYH WLPHV WKH PD[LPXP IORDWLQJ YHORFLW\ RI WKH Microcystis. 3UHVXPDEO\ WKH SRRU HQWUDLQPHQW RI FRORQLHV LQ WKH ZDWHU PRWLRQ KDG UHVXOWHG LQ D VHSDUDWLRQEHWZHHQWKHWRSDQGERWWRPSRSXODWLRQV :KHQ y! IORDWLQJ RU VLQNLQJ UDWH FRQWUROV FHOO GLVWULEXWLRQ ZKLOH LI y GLIIXVLYH WUDQVSRUW EHFRPHV LQFUHDVLQJO\ LPSRUWDQW LQ GHWHUPLQLQJ FHOO GLVWULEXWLRQ5H\QROGVHWDO %XR\DQWFRORQLHV RIWKH JDV YDFXRODWH F\DQREDFWHULXP Microcystis VS FDQ KDYH UDSLG IORDWLQJ UDWHV XS WR P G FRPSDUHGWRWKHJHQHUDOO\VORZVLQNLQJUDWHVRIPDQ\ QRQEXR\DQW PLFURDOJDH FD P G 6LPXODWLRQ VWXGLHV SUHGLFWHGWKH RFFXUUHQFH RIWRSKHDY\ YHUWLFDO SURILOHVRIEXR\DQWMicrocystis IRU ZKLFKy! ZKLOH XQGHU WKH VDPH FRQGLWLRQV WKH GLIIXVLRQ GRPLQDWHG GLVWULEXWLRQ RI VLQNLQJ FHOOV y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phanizomenon flos-aquae RYHU D GD\ SHULRG LQ WKH %DOWLF 6HD 7KH DQDO\VLV GHPRQVWUDWHG WKDW DYHUDJHG RYHU WKH DOWHUQDWLQJ SHULRGV RI FDOP DQG PL[LQJEXR\DQF\SURYLGHGE\JDVYDFXROHVLQFUHDVHG WKH GDLO\ QHW DUHDO SKRWRV\QWKHVLV RI WKH F\DQREDFWHULDOSRSXODWLRQE\QHDUO\WZRIROG &XUUHQWV JHQHUDWHG E\ ZLQG PL[LQJ QRW RQO\ LQIOXHQFH WKH YHUWLFDO GLVWULEXWLRQ RI SK\WRSODQNWRQ EXWDOVR DGYHFWSDUWLFOHV IURPRQHORFDWLRQWR DQRWKHU FDXVLQJ KRUL]RQWDO DQG YHUWLFDO KHWHURJHQHLW\ RIDOJDO SRSXODWLRQV RQ ERWK D ORFDO DQG EDVLQ VFDOH *HRUJH
5/ 2OLYHUDQG ** *DQI
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
U a U a 2
)RU Ua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
&KDSWHU )UHVKZDWHU%ORRPV KLJKHVWULVNIRUOLYHVWRFNSRLVRQLQJDQGZLQGVSHHGV RIWRPV DUHFULWLFDOWRWKHLUIRUPDWLRQ
) 6XUIDFH%ORRPV 7KH JDVYDFXRODWH F\DQREDFWHULD WKDW FRPPRQO\ SURGXFH VXUIDFH EORRPV DUH WKRVH VSHFLHV WKDW IRUP ODUJHELRPDVVXQLWV HLWKHUFRORQLHV DV LQMicrocystis, RU DJJUHJDWHV RI ILODPHQWV DV LQ Anabaena DQG Aphanizomenon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¡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
EXR\DQW FRORQLHV ,EHOLQJV DQG 0XU :DOVE\ 7KLV SK\VLFDO UHVWUDLQW RQ YHUWLFDO PRYHPHQW KDVEHHQGHPRQVWUDWHGE\,EHOLQJVDQG0XU LQ YHU\ GHQVH VXUIDFH VFXPV RI Microcystis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icrocystis aeruginosa VKRZHG WKDW &2 OLPLWDWLRQ FDQ SURPRWH EXR\DQF\ LQ WKH VKRUW WHUP E\ SUHYHQWLQJ ERWK JDV YHVLFOHFROODSVHDQGWKHDFFXPXODWLRQRIFDUERK\GUDWH EDOODVW 7KLV VXSSRUWV WKH UHVXOWV RI :DOVE\ DQG %RRNHU ZKR IRXQG WKDW FDUERQ OLPLWDWLRQ FDXVHG Anabaena flos-aquae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nabaena Aphanizornenon flos-aquae
oscillarioides WKH\ FRQFOXGHG WKDW &2 ZDV WKH SUHIHUUHG LQRUJDQLF FDUERQ VRXUFHIRUVXVWDLQLQJ KLJK UDWHV RI SKRWRV\QWKHVLV DQG WKDW WKH IRUPDWLRQ RI VXUIDFHEORRPVZRXOGSURYLGHDFFHVVWRWKHSUHIHUUHG IRUP DW WKH DLUZDWHU LQWHUIDFH ,Q SRRUO\ EXIIHUHG ZDWHUVZKHUHS+YDOXHVDUHIUHTXHQWO\LQH[FHVVRI LQGLFDWLQJ GLPLQLVKLQJ VXSSOLHV RI &2 GXH WR VXVWDLQHG SHULRGV RI LQWHQVH SKRWRV\QWKHVLV WKLV ZRXOG SURYLGH DQ HFRORJLFDO DGYDQWDJH RYHU QRQ EXR\DQW VSHFLHV 3DHUO HW DO VKRZHG WKDW QDWXUDO SRSXODWLRQV RI Microcystis KDG RSWLPDO SKRWRV\QWKHWLF UDWHV DQG UHVLVWDQFH WR SKRWRLQKLELWLRQ DW VXUIDFH LUUDGLDQFHV +RZHYHU ,EHOLQJV DQG 0XU ZRUNLQJ DW D YHU\ ILQH VFDOHV P VXJJHVW WKDW WKH FDVH LV QRW \HW SURYHQ DQG SURYLGHG HYLGHQFH WR VKRZ WKDW SKRWRV\QWKHVLV LQ Microcystis VFXPVFDQEHLQKLELWHGDWWKHVXUIDFH 6XUIDFH EORRPV PDUNHGO\ LQIOXHQFH WKH GHSWK WR ZKLFKOLJKWSHQHWUDWHVWKH ZDWHUFROXPQ ODUJHO\ DV D UHVXOW RI VFDWWHULQJ E\ JDV YHVLFOHV :DOVE\ 6FDWWHULQJ IURPWKH YHUWLFDO LQFUHDVHV WKH SDWK OHQJWK RI SKRWRQV WKURXJK KRUL]RQWDO ZDWHU OD\HUV WKHUHE\ HQKDQFLQJ WKH SUREDELOLW\ RI DEVRUSWLRQ ZLWKLQ WKH QHDU VXUIDFH OD\HUV 7KRVH RUJDQLVPV ZKLFK KDYH WDNHQ XS SUHIHUHQWLDO UHVLGHQFH ZLWKLQ WKH HXSKRWLF ]RQH VXFKDVEXR\DQWF\DQREDFWHULDWKHUHIRUHKDYHD JUHDWHU SUREDELOLW\ RI LQWHUFHSWLQJ WKH OLJKW )RU H[DPSOH *DQIHW DO VKRZHG WKDW RIWKH OLJKW VFDWWHUHG LQ 0W %ROG 5HVHUYRLU ZDV GXH WR Microcystis aeruginosa FRORQLHV DQG FRQFOXGHG WKDW WKLV SURYLGHG Microcystis ZLWK D GLVWLQFW DGYDQWDJH ZKHQLQFRPSHWLWLRQZLWKQRQEXR\DQWVSHFLHV
* 0HWDOLPQHWLF3RSXODWLRQV 6RPH VSHFLHV RI JDVYDFXRODWH F\DQREDFWHULD HJ Oscillatoria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
5/2OLYHUDQG***DQI DQG VHQVLWLYHWRYDULDWLRQVLQOLJKWLQWHQVLW\ .RQRSND HWDO IRXQGWKDWDWLQWHQVLWLHVRI! PROP V EXR\DQWILODPHQWV RIO. agardhii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scillatoria agardhii LQ /DNH *MHUVM¡HQ :DOVE\ HW DO 7KH YDULDEOH UHVSRQVLEOH IRU LQGXFLQJ WKH ILODPHQWDJJUHJDWLRQKDVQRWEHHQLGHQWLILHG
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a). 6KDOORZSURGXFWLYHZDWHUVWKDWDUHIUHTXHQWO\ IXOO\PL[HG VXSSRUWSRSXODWLRQV RIQRQDJJUHJDWHG ILODPHQWRXV F\DQREDFWHULD 3ODWH G VXFK DV
&KDSWHU )UHVKZDWHU%ORRPV Oscillatoria VSS *LEVRQ HW DO 6FKHIIHU HW DO 1(b). $QDQDORJRXVPL[LQJUHJLPHRFFXUVLQGHHSHU ODNHV WKDW DUH WKHUPDOO\ VWUDWLILHG DQG KDYH DQ LQWHQVHVHDVRQDOWKHUPRFOLQHEXWZKHUHWKH VXUIDFH OD\HU LV FRQVLVWHQWO\ ZHOO PL[HG 7KHVH WRR DUH IUHTXHQWO\ LQKDELWHG E\ F\DQREDFWHULDFRQVLVWLQJ RI VPDOO VROLWDU\ ILODPHQWV D XQLTXH H[DPSOH EHLQJ Anabaena minutissima LQ /DNH 5RWRQJDLR 1HZ =HDODQG:DOVE\HWDO D ,Q VKDOORZ SURGXFWLYH ZDWHUV ZKHUH WKHUPDO VWUDWLILFDWLRQ RIWKH ZDWHU FROXPQ RFFXUV GLXUQDOO\ RU RFFDVLRQDOO\ SHUVLVWV IRU ORQJHU SHULRGV WKH ODUJHU PRUSKRORJLFDO IRUPV RIF\DQREDFWHULDRFFXU LQFOXGLQJ FRORQLDO IRUPV VXFK DV Microcystis aeruginosa *DQI ,EHOLQJV HW DO E DQGDJJUHJDWHGRUFRQWRUWHGILODPHQWRXVIRUPVVXFK DV Aphanizomenon DQG Anabaena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scillatoria, Anabaena RU Aphanizomenon (GPRQGVRQ .OHPHU :DOVE\ HW DO .RQRSND HW DO EXW PHWDOLPQHWLF SRSXODWLRQV RI RWKHU IRUPV LQFOXGLQJ Microcystis, KDYHEHHQUHFRUGHG:DUGDQG:HW]HO 7KHVH SDWWHUQV VXJJHVW WKDW WKH VPDOOHU IRUPV RI F\DQREDFWHULD RFFXU LQ FRQGLWLRQV ZKHUH WKH LPSRUWDQFH RIEXR\DQF\ UHJXODWLRQ LV UHGXFHG HLWKHU LQ VLWXDWLRQV ZKHUH EXR\DQF\ UHJXODWLRQ LV RYHUZKHOPHG E\ PL[LQJ RU ZKHQ WKH RUJDQLVPV DUH KROGLQJVWDWLRQEHORZWKHGHSWKRIWKHPL[HGOD\HU ,Q FRQWUDVW ZKHQWKHGXUDWLRQRIWKHUPDOVWUDWLILFDWLRQLV SURORQJHG DQG WKH XWLOLW\ RI EXR\DQF\ UHJXODWLRQ LV HQKDQFHG WKHQ WKH ODUJHU IRUPV RI F\DQREDFWHULD EHFRPH PRUH SUHYDOHQW ,Q HVVHQFH PRUSKRORJ\ UHIOHFWV WKH DGYDQWDJH SURYLGHG E\ EXR\DQF\
UHJXODWLRQ DQG WKLV LV GHSHQGHQW RQ WKH PL[LQJ UHJLPH 7KH JHQHUD RI JDVYDFXRODWH F\DQREDFWHULD GR QRW VHSDUDWHRXWVWULFWO\RQWKHEDVLV RIWKHVHWKUHHKDELWDW FDWHJRULHV DV YDULDWLRQV LQ IRUP DQG IXQFWLRQ RFFXU ZLWKLQ DQG EHWZHHQ WD[D ,Q SDUWLFXODU VRPH RI WKH ILODPHQWRXV VSHFLHV VXFK DV Oscillatoria agardhii :DOVE\ HW DO DQGAphanizomenon flos-aquae /\QFK *DQI FDQRFFXUHLWKHUDVVLQJOH ILODPHQWV LQ PHWDOLPQHWLF SRSXODWLRQV RU DV ODUJH DJJUHJDWHV LQ HSLOLPQHWLF SRSXODWLRQV :DOVE\ HW DO (YHQ ZLWKLQ D PRUSKRORJLFDO W\SH WKHUH LV D PL[ RI VSHFLHV ZLWK TXLWH GLIIHUHQW SK\VLRORJLFDO FKDUDFWHULVWLFV )RUH[DPSOHVRPHILODPHQWRXVIRUPV FDQ IL[ PROHFXODU QLWURJHQ ZKLOH RWKHUV FDQQRW 'LIIHUHQFHV EHWZHHQ WKH WKUHH VSHFLHV Oscillatoria rubescens, O. agardhii DQG O. redekei DSSHDU WR VHSDUDWH WKHLU RFFXUUHQFH EHWZHHQ PL[LQJ UHJLPHV O. rubescens RFFXUVPRUHFRPPRQO\DVPHWDOLPQHWLF SRSXODWLRQVZKLOHO. agardhii FDQIRUPPHWDOLPQHWLF SRSXODWLRQV (GPRQGVRQ .OHPHU .RQRSND HWDO EXW DOVR RFFXUV LQ ZHOO PL[HG OD\HUV %URRN HW DO *LEVRQ HW DO redekei DSSHDUV WR EH UHVWULFWHG WR VKDOORZ XQVWUDWLILHGODNHVSUREDEO\EHFDXVHLWKDVWRRIHZJDV YHVLFOHV IRU SRVLWLYH EXR\DQF\ :KLWWRQ DQG 3HDW 0HIIHUWDQG.UDPEHFN O. agardhii DQG O. redekei PD\ FRRFFXU DV LQ WKH VKDOORZ FRPSOHWHO\PL[HG/RXJK1HDJK*LEVRQHWDO
9, 3K\VLFDO&RQWURORI&\DQREDFWHULD 7KH LQWHUDFWLRQ EHWZHHQ PL[LQJ SURFHVVHV EXR\DQF\ UHJXODWLRQDQGVSHFLHVFRPSRVLWLRQ VXJJHVWVWKDWODNH UHVWRUDWLRQ YLD PDQLSXODWLRQ RI WKH WXUEXOHQW HQYLURQPHQWLVDYLDEOHODNHPDQDJHPHQWVWUDWHJ\
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
VHWWOLQJ 5H\QROGV ,I PL[LQJ LV DUWLILFLDOO\ LQGXFHG WKHQ LQWHUPLWWHQW FLUFXODWLRQ PD\EH PRUH DSSURSULDWH WKDQ FRQWLQXRXV FLUFXODWLRQ 6WHLQEHUJ DQG*UXKO 9LVVHUHWDO 7KLVZRXOGDOORZ QRQEXR\DQWFHOOVWRVHGLPHQWPRUHUDSLGO\EHORZWKH ]RQH RI FLUFXODWLRQ DQG LQWR GDUNQHVV ZKHUH WKH V\QWKHVLVRIYHVLFOHV LV PXFK VORZHUEHLQJ GHSHQGHQW XSRQ VWRUHG HQHUJ\ UHVHUYHV 'HDFRQ DQG :DOVE\ IRXQGWKDWDWDQRSWLPDOLUUDGLDQFHRI PRO P V LWWRRNMicrocystis WKUHHGD\V EHIRUHEXR\DQF\ UHFRYHUHG ,QWHUPLWWHQW FLUFXODWLRQ ZRXOG DOVR SURYLGH DQ RSSRUWXQLW\ IRU EXR\DQW FHOOV WR PRYH WRZDUGV WKH LOOXPLQDWHG VXUIDFH OD\HUV ZKHUH WXUJRU SUHVVXUHLV OLNHO\WR LQFUHDVHSULRUWRWKHQH[WPL[LQJ HYHQW .LQVPDQ HW DO IRXQG WKDW WXUJRU SUHVVXUHLQFUHDVHGIURP WR LQ Anabaena flosaquae DIWHU K DW DSKRWRQ LUUDGLDQFH RI PRO P V $Q DOWHUQDWLYH VWUDWHJ\WR ODNHPL[LQJ DQG DGRSWHG E\ WKH IRUPHU 6XIIRON :DWHU %RDUG LQ (QJODQG LV WR DEVWUDFWZDWHUIURPWKHODNHDQGFLUFXODWHLWWKURXJKD PGHHSSLSH ZKHUH WKHK\GURVWDWLF SUHVVXUH DWWKH ERWWRP LV VXIILFLHQW WR FROODSVH DOO JDV YDFXROHV LQ Microcystis aeruginosa &ODUNH DQG :DOVE\ $OWHUQDWLYHO\ :DOVE\ QRWHG WKDW 0HQGD\ DQG %XFN KDG VXFFHVVIXOO\ XVHG H[SORVLRQV WR FUHDWH VKRFNZDYHV WR FROODSVH JDV YDFXROHV EXW WKH VLGHHIIHFWV RI WKLV WUHDWPHQW ZHUH QRW DOZD\V DFFHSWDEOH SDUWLFXODUO\ GXH WR WKHLU HIIHFW RQ RWKHU RUJDQLVPV
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icrocystis aeruginosa LQ UHVSRQVH WR PL[LQJ 7RHW] 5H\QROGVHWDO 9LVVHUHW DO /DNH 1LHXZH 0HHU LV K\SHUWURSKLF DQG
5/2OLYHUDQG***DQI GRPLQDWHG E\ Microcystis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cenedesmus, FHQWULF GLDWRPVDQGIODJHOODWHVGRPLQDWHG ,Q DUWLILFLDO PL[LQJKDGEHHQFRQWLQXRXV IURP 0DUFKWR 6HSWHPEHU ZKHUHDVGXULQJ WR UHGXFH HQHUJ\ FRVWV PL[LQJ ZDV DSSOLHG LQWHUPLWWHQWO\ DQG FRQWUROOHG E\ WKH VXUIDFH DQG ERWWRP WHPSHUDWXUHV DQGR[\JHQFRQFHQWUDWLRQV ,IJUHDWHUWKDQ & RUJ P UHVSHFWLYHO\ WKH FRPSUHVVRU ZDV VZLWFKHG RQ DQG LI WKH GLIIHUHQFH ZDV & RU J P LW ZDV VZLWFKHG RII 7KLV JDYH DQ DQQXDO VDYLQJ RI Microcystis IDLOHG WR GHYHORS LQ HLWKHU RI WKH \HDUV ZKHQ DUWLILFLDO PL[LQJ WRRN SODFH DQG Anabaena QXPEHUV SHU P GHFUHDVHG EXW WKH QXPEHUV RI Aphanizomenon DQG WR D OHVV H[WHQW Aphanocapsa ZHUH JUHDWHU 7KLV REVHUYDWLRQ VXJJHVWV WKDW XQFULWLFDO DGRSWLRQ RI DUWLILFLDO GHVWUDWLILFDWLRQ WR PDQDJH F\DQREDFWHULD LV XQZLVH DOWKRXJK LW FDQ FDXVH PDMRU VKLIWV LQ VSHFLHV FRPSRVLWLRQ 6WHLQEHUJ DQG =LPPHUPDQQ ,Q D UHYLHZ RI WKH HIIHFWLYHQHVV RI DUWLILFLDO GHVWUDWLILFDWLRQ LQ $XVWUDOLDQ UHVHUYRLUV 0F$XOLIIH DQG 5RVLFK IRXQG WKDW LW IDLOHG WR VDWLVIDFWRULO\ FRQWURO SK\WRSODQNWRQLQRYHU RIFDVHV 6WHLQEHUJ DQG *UXKO UHSRUWHG RQ D VHULHV RI H[SHULPHQWVLQYHVWLJDWLQJSK\VLFDOPHDVXUHVWRLQKLELW SODQNWRQLF F\DQREDFWHULD LQ )LVFKNDOWHUVHH D VPDOO %DYDULDQ ODNH KD DQG P PHDQ GHSWK 3HUPDQHQW GHVWUDWLILFDWLRQ LQLWLDOO\ UHVXOWHG LQ SRVLWLYH UHVSRQVHV ZLWK DOPRVW FRPSOHWH UHPRYDO RI WKH F\DQREDFWHULD EXW QHDUO\ D GRXEOLQJ LQ WKH ELRPDVV RI FKORURSK\WHV DQG GLDWRPV +RZHYHU Limnothrix UHDSSHDUHG LQWKHWKLUG\HDUDQGUHDFKHG D SHDNELRPDVV LQ WKH IRXUWK \HDU RIWKH WUHDWPHQW WKDW ZDV KLJKHUWKDQKDG RFFXUUHGSULRU WR GHVWUDWLILFDWLRQ EHLQJ XVHG ,W ZDV FRQFOXGHG WKDW DOWKRXJK WKH FRQWLQXRXV PL[LQJ SURGXFHG UHODWLYHO\ FRQVWDQW PL[LQJ SDWWHUQV ZLWK D ORZ PHDQ LUUDGLDQFH WKH ODNH ZDV QRW VXIILFLHQWO\ GHHS WR UHGXFH WKH PHDQ
&KDSWHU )UHVKZDWHU%ORRPV LUUDGLDQFH EHORZ WKDW UHTXLUHG E\ WKH 2VFLOODWRULDFHDH ,QWHUPLWWHQW GHVWUDWLILFDWLRQ ZDV LQVWLWXWHG RQ WKH EDVLV WKDW LW VKRXOG HOHYDWH ELRPDVV ORVVHVYLDVHGLPHQWDWLRQDQGLPSURYHWKHOLJKWFOLPDWH DV D FRQVHTXHQFH RI GLPLQLVKHG SK\WRSODQNWRQ ELRPDVV DQG WKLV ZRXOG FXUWDLO WKH ORZ OLJKW DGDSWHG F\DQREDFWHULD 8QGHUWKLV UHJLPH Limnothrix SHDNHG RQO\ IRU D IHZ GD\V DQG ZDV RXWFRPSHWHG E\ WKH FKORURSK\WHV DQG GLDWRPV DV KDG RFFXUUHG LQLWLDOO\ ZLWK FRQWLQXRXV GHVWUDWLILFDWLRQ ,Q FRQWUDVW WR SHUPDQHQWPL[LQJ WKH F\DQREDFWHULDGLG QRWUHDSSHDU DQGWKHELRPDVVRIDOJDHUHPDLQHGORZ
& 0L[LQJLQ5HJXODWHG5LYHUV 0DQLSXODWLRQ RI PL[LQJ UHJLPHV LV QRW FRQILQHG WR VWLOO ZDWHU ERGLHV EXW FDQ DOVR EH XVHG LQ UHJXODWHG ULYHUV %XUFK HW DO DQG :HEVWHU HW DO KDYH VKRZQ WKDW WKH RFFXUUHQFH RI Anabaena circinalis LV DIXQFWLRQ RIGLVFKDUJHUDWHV LQUHJXODWHG ULYHUV $W0DXGH:HLU RQ WKH 0XUUXPELGJHH 5LYHU $XVWUDOLD IORZVEHORZWR 0/G FDXVHG D GHFOLQH LQ WKH SRSXODWLRQ GHQVLW\ RI WKH GLDWRP Aulacoseira (Melosira) granulata, ZKLOH Anabaena circinalis LQFUHDVHG6KHUPDQHWDO 7KLV VKLIW LQ VSHFLHV FRPSRVLWLRQ ZDV H[SODLQHG E\ FKDQJHV LQ WKHUPDOVWUDWLILFDWLRQDQGPL[LQJ 'LVFKDUJHV 0 / G SHUPLWWHG WKH GHYHORSPHQW RI SHUVLVWHQW VWUDWLILFDWLRQ & RYHU P DQG WKH FRROHU ULYHU ZDWHUDWWKHXSVWUHDPHQGRIWKHSRROSOXQJHGEHQHDWK WKH VWUDWLILHG HSLOLPQLRQ GLVFKDUJLQJ YLD WKH XQGHUIORZ ZHLU 7KHVH FRQGLWLRQV UHVXOWHG LQ HSLOLPQHWLF ZDWHU YHORFLWLHV FORVHWR ]HUR SURYLGLQJ D UHWHQWLRQ WLPH VXIILFLHQW WR SHUPLW A. circinalis JURZLQJ DW D QHW UDWH RI G WR GHYHORS D VLJQLILFDQW ELRPDVV RYHU D SHULRG RI RQH WR WZR ZHHNV :LWK WKH SHUVLVWHQW VWUDWLILFDWLRQ WKH EXR\DQW Anabaena DFFXPXODWHG LQ WKH WRS P ZKLOH QHJDWLYHO\ EXR\DQW Aulacoseira VDQN DQG DFFXPXODWHGLQWKHERWWRPZDWHUV $VWKHULYHUKDV D HXSKRWLF]RQHRIRQO\ PDQGDPD[LPXPGHSWKRI P=HX=P WKLVSURYLGHGDGLVWLQFWDGYDQWDJH WRWKHF\DQREDFWHULXP 7KHHPHUJHQFHRIAnabaena DVWKHGRPLQDQWJHQXV ZDV DWWULEXWHG WR QLWURJHQ OLPLWDWLRQ ZKLFK KDG EHHQ GHPRQVWUDWHG LQ WKH Aulacoseira SRSXODWLRQ SULRU WR LWV GHPLVH XVLQJ D SK\VLRORJLFDO DVVD\ EDVHG RQ FKORURSK\OODIOXRUHVFHQFH :RRG DQG 2OLYHU ,W ZDV VXJJHVWHG WKDW QLWURJHQ OLPLWDWLRQ SURYLGHG FRQGLWLRQV VXLWDEOH IRU Anabaena WR GRPLQDWH UDWKHU WKDQDQRQQLWURJHQIL[LQJF\DQREDFWHULXP 'XULQJD VLPLODU ORZ IORZ SHULRG WKH A. circinalis SRSXODWLRQ
UHDFKHG D SODWHDX DV WKH WRWDO UHDFWLYH DQG ILOWUDEOH UHDFWLYHSKRVSKRUXVIHOOEHORZPJP VXJJHVWLQJ WKDW SKRVSKRUXV DYDLODELOLW\ PD\ KDYH FRQWUROOHG WKH SRSXODWLRQPD[LPXP:HEVWHUHWDO %XUFK HW DO LQ WKH 6RXWK $XVWUDOLDQ VHFWLRQ RIWKH5LYHU0XUUD\ GHPRQVWUDWHG WKDWSRSXODWLRQV RI A. circinalis RFFXUUHGLQWKHULYHUZKHQGLVFKDUJHZDV VPDOO HQRXJK WR SHUPLW WKHUPDO VWUDWLILFDWLRQ 0RUH LPSRUWDQWO\ WKH GURS LQ ULYHUOHYHOV GXH WR LUULJDWLRQ GHPDQG DW ORZ IORZV DOORZHG ZDWHU IURP WKH QHLJKERXULQJ ZHWODQGV ELOODERQJV WR IORZ EDFNLQWR WKHPDLQFKDQQHOFDUU\LQJZLWKLWVLJQLILFDQWQXPEHUV RI A. circinalis 3 %DNHU SHUV FRPP ,Q WKLV V\VWHPLW DSSHDUV DV WKRXJK WKH PDLQ LQRFXOXP RIA. circinalis FRPHVIURPWKHFDOPDGMDFHQWZHWODQGV
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icrocystis ZKLFKKDGD ODUJH T YDOXH RIFD DQG DOVR D KLJKHU PLQLPXP WHPSHUDWXUH IRU JURZWK 7KRPDV DQG :DOVE\ 5REDUWVDQG=RKDU\ 7KH EHOLHI WKDW F\DQREDFWHULD SUHIHU KLJKHU WHPSHUDWXUHV LV EDVHG PDLQO\ RQ ILHOG VWXGLHV DQG VHDVRQDO FRUUHODWLRQV EXW WKHVH PD\ EH VSXULRXV DV
KLJK WHPSHUDWXUHV DUH DOVR DVVRFLDWHG ZLWK WKHUPDO VWUDWLILFDWLRQ DQG FKDQJHV LQ WXUEXOHQW PL[LQJ WKDW SURYLGH DQ HQYLURQPHQW FRQGXFLYH WR WKH JDV YDFXRODWHF\DQREDFWHULD )R\HWDO FRQFOXGHG WKDW WKH WHPSHUDWXUH RSWLPD RIFXOWXUHV RIAnabaena flos-aquae, Aphanizomenon flos-aquae IR gracile, Oscillatoria agardhii DQG Oscillatoria redekei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
Fig. 7 $EVRUSWLRQ VSHFWUD RI FKORURSK\OOD FKOD FDURWHQRLGV FDURWQ DQG WKH SK\FRELOLSURWHLQV FSK\FRHU\WKULQ &3( F SK\FRF\DQLQ &3& DQG DOORSK\FRF\DQLQ $3& FRPSLOHG IURPD QXPEHURIVRXUFHV 7KH DEVRUSWLRQ VSHFWUXP IRUHDFK SLJPHQW LV VKRZQUHODWLYHWRLWVDEVRUSWLRQSHDN
FRQFHQWUDWHG LQ WKHVH ZDYHEDQGV RU ZKHQ WKHUH DUH VXEVWDQWLDOIOXFWXDWLRQVLQOLJKWTXDOLW\RYHUWLPH 7KH XQGHUZDWHU OLJKW FOLPDWH FKDQJHV ERWK LQ TXDQWLW\ DQG LQ TXDOLW\ ZLWK GHSWK 7KH LQWHQVLW\ GHFUHDVHV H[SRQHQWLDOO\ DV D UHVXOW RIDEVRUSWLRQ DQG VFDWWHULQJ E\ SDUWLFOHV DQG FRORXUHG FRPSRXQGV DQG WKH VHOHFWLYHUHPRYDO RIZDYHOHQJWKV FDXVHV VKLIWV LQ WKHVSHFWUDOGLVWULEXWLRQ.LUN 2OLYHUDQG*DQI 2OLYHU .LUN DQG 2OLYHU :DWHU
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
Fig. 8 6SHFWUDO GLVWULEXWLRQ RITXDQWD FDOFXODWHG IRUFOHDU QDWXUDO ZDWHU DW WKH VXUIDFH DQG DW P GHSWK WRS ILJXUH DQG IRU RUJDQLFDOO\ FRORXUHG ZDWHU IURP 0W %ROG 5HVHUYRLU 6RXWK $XVWUDOLDDWWKHVXUIDFHDQGDWPGHSWKERWWRPILJXUH
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
QXPEHU RI SKRWRV\QWKHWLF XQLWV )DONRZVNL DQG /D5RFKH ,I SKRWRV\QWKHVLV EHFRPHV OLPLWHG E\ WKH UDWH RI GHOLYHU\ RI OLJKW HQHUJ\ WR WKH SKRWRV\VWHPV DV XQGHU ORZ LUUDGLDQFH WKHQ DQ LQFUHDVH LQ DQWHQQD VL]H SURYLGHV RQH PHDQV RI LQFUHDVLQJ WKH SKRWRQ VXSSO\ ,I WKH VXSSO\ RI SKRWRQV IURP WKH DQWHQQD DSSURDFKHV WKH PD[LPXP WXUQRYHUUDWH RIWKH SKRWRV\VWHP WKHQ DQ LQFUHDVH LQ WKH QXPEHU RISKRWRV\QWKHWLF XQLWV ZLOO LQFUHDVH WKH WRWDO VXSSO\ RI HQHUJ\ WR WKH FHOO IRU SKRWRV\QWKHVLV DQG JURZWK )DONRZVNL DQG /D5RFKH 7KLV DGDSWDWLRQ FRXOG EH SDUWLFXODUO\ LPSRUWDQW LI FHOOV ZHUH UHJXODUO\ H[SRVHG WR VDWXUDWLQJ OLJKW LQWHQVLWLHV IRU VKRUW SHULRGV RI WLPH IRU H[DPSOH LI PL[HG UDSLGO\ EHWZHHQ WKH VXUIDFH DQG DSKRWLF ]RQH DV WKH LQFUHDVH LQWKH QXPEHU RISKRWRV\QWKHWLF XQLWV ZRXOG KHOS PD[LPLVH WKH FDSWXUH RIHQHUJ\ GXULQJ WKH EULHI OLJKWSHULRG 7LO]HU :KHQ FHOOV DUH H[SRVHG WR KLJK LUUDGLDQFH LQWHQVLWLHV IRU H[WHQGHG SHULRGV WKHLU LV DQ LQFUHDVHG ULVN RI SKRWRGDPDJH WR WKH SKRWRV\VWHPV 7KLV ULVN FDQ EH FXUWDLOHG E\ D UHGXFWLRQ LQ DQWHQQD VL]H DQG IXUWKHU LI WKH UDWH RI HQHUJ\ FDSWXUH H[FHHGV WKH FDSDFLW\ RI WKH FHOO PDFKLQHU\ WKHQ WKH QXPEHU RI SKRWRV\QWKHWLF XQLWV FDQ EH UHGXFHG &KDQJHV LQ DQWHQQD VL]H DQGRU FKDQJHV LQ WKH QXPEHU RI SKRWRV\QWKHWLF XQLWV DUH SKRWRDFFOLPDWLRQ VWUDWHJLHV HPSOR\HG E\ ERWK F\DQREDFWHULD DQG PLFURDOJDH XQGHU YDULRXV FRQGLWLRQV ,W PLJKW EH H[SHFWHG IURP FRPSDULVRQV RI DQWHQQD VWUXFWXUH DQG VSHFWUDO DEVRUSWLRQ FKDUDFWHULVWLFV WKDW WKH F\DQREDFWHULD ZRXOG UHVSRQG TXLWH GLIIHUHQWO\ WR FKDQJHV LQ 3$5 FRPSDUHG WR WKH PLFURDOJDH 7R GDWH WKHUH LV OLWWOH H[SHULPHQWDO GDWD WR DVVHVV WKLV H[SHFWDWLRQDVWKHPDMRULW\RIODERUDWRU\VWXGLHVKDYH XVHG ZKLWH OLJKW DQG LQYHVWLJDWHG UHVSRQVHV WR FKDQJLQJ LUUDGLDQFH LQWHQVLW\ 7KH GLIIHUHQW FKDUDFWHULVWLFV RI WKH SKRWRV\QWKHWLF XQLWV VXJJHVWV WKDWLQFUHDVHGHIIRUWVKRXOGEHJLYHQWRGHVFULELQJWKH HIIHFWV RI VSHFWUDO FKDQJHV RQ SKRWRV\QWKHVLV DQG JURZWK 3HUKDSV DQ LQGLFDWLRQ RI WKH LPSRUWDQFH RI WKLVLVWKHFRPPRQREVHUYDWLRQWKDWF\DQREDFWHULDDUH IUHTXHQWO\ DVVRFLDWHG ZLWK RUJDQLFDOO\ULFK ZDWHUV $OWKRXJK RIWHQ LQWHUSUHWHG DV LQGLFDWLQJ D UROH IRU SKRWR KHWHURWURSK\ LW PLJKW DOVR UHIOHFW D PRUH FRQGXFLYH VSHFWUDO GLVWULEXWLRQ RI 3$5 IRU F\DQREDFWHULD
$ /LJKW,QWHQVLI\ &\DQREDFWHULD VKRZ DQ LQYHUVH FRUUHODWLRQ EHWZHHQ SLJPHQW FRQWHQW DQG LUUDGLDQFH LQWHQVLW\ D UHVSRQVH
5/2OLYHUDQG***DQI FRPPRQO\ IRXQG LQ SKRWRWURSKLF RUJDQLVPV .RQRSNDDQG 6FKQXU )R\DQG *LEVRQ 5LFKDUGVRQ HW DO :\PDQ DQG )D\ D 7DQGHDX GH 0DUVDF DQG +RXPDUG 7KLV UHVSRQVH RFFXUV LQ F\DQREDFWHULD ZKHWKHU WKH\ DUH JURZLQJ XQGHU D VKRUW OLJKWGDUN F\FOH /' )R\ D ORQJ /' F\FOH 3RVW HW DO D RU FRQWLQXRXV LOOXPLQDWLRQ 9DQ /LHUH DQG 0XU 3RVWHWDO ,Q HLJKW VWUDLQV RI F\DQREDFWHULD Anabaena flosaquae, Anabaena solitaria, Anabaena circinalis [ Oscillatoria redekei, Oscillatoria agardhii, Microcystis aeruginosa, Gloeotrichia echinulata VWXGLHG E\ :\PDQ DQG )D\ D WKH UDWLR RI FKORURSK\OOD WR SK\FRELOLSURWHLQV FK3%3 UHPDLQHGFRQVWDQWGXULQJOLJKWOLPLWHGJURZWK GHVSLWH WKH UHGXFWLRQ LQ SLJPHQWV DV LUUDGLDQFH LQWHQVLW\ LQFUHDVHG 7KLV LQGLFDWHG WKDW WKH QXPEHU RI SKRWRV\QWKHWLF XQLWV SHU FHOO GHFOLQHG LQ UHVSRQVH WR LQFUHDVLQJLUUDGLDQFH:\PDQDQG)D\ D3RVWHW DO 7DQGHDX GH 0DUVDF DQG +RXPDUG 2QFHPD[LPXPJURZWKUDWHZDVDWWDLQHGWKHFKO3%3 UDWLR LQFUHDVHG ZLWK IXUWKHU LQFUHDVHV RILUUDGLDQFH LQ DOO VSHFLHV H[FHSW WKH SK\FRHU\WKULQULFK VWUDLQV Oscillatoria agardhii and Gloeotrichia echinulata 7KLVFKDQJHLQUDWLR LQGLFDWHG DGHFOLQH LQWKH VL]HRI WKH 3%6 :\PDQ DQG )D\ D SUHVXPDEO\ UHGXFLQJ H[FLWRQ GHOLYHU\ WR 36,, DQG KHOSLQJ SRVWSRQH WKH RQVHW RISKRWRLQKLELWLRQ DQG GDPDJH WR WKHUHDFWLRQ FHQWUH $OWKRXJKWKH WZRSK\FRHU\WKULQ ULFK VWUDLQV GLG QRW LQFUHDVH WKHLU FKO3%3 UDWLR DW LUUDGLDQFHV LQ H[FHVV RI WKDW UHTXLUHG IRU PD[LPXP JURZWK UDWH WKH SK\FRHU\WKULQ FRQWHQW RI WKH 3%6 GHFOLQHG UHGXFLQJ LWV HIIHFWLYH VSHFWUDO VL]H 7KHVH VDPH DGMXVWPHQWV ZHUH REVHUYHG RQ VKLIWLQJ Synechococcus 3&& IURP KLJK WR ORZ OLJKW ZLWK DQ LQLWLDO LQFUHDVH LQ WKH 3%6 DQWHQQD VL]H IROORZHGE\ DQ LQFUHDVH LQ3%6 SHU DUHDRIWK\ODNRLG 7DQGHDXGH0DUVDFDQG+RXPDUG $OWKRXJK VLPLODUFKDQJHV LQSLJPHQW FRQWHQWRFFXU LQ WKH HXNDU\RWLF DOJDH LQ UHVSRQVH WR DOWHUDWLRQV LQ OLJKW LQWHQVLW\ WKHUH DUH SUHOLPLQDU\ LQGLFDWLRQV WKDW WKH DFFOLPDWLRQ WR SKRWRSHULRG LV GLIIHUHQW .URPNDPS )ODPHOLQJDQG.URPNDPS ,Q Scenedesmus protuberans WKH SLJPHQW FRQWHQW LQFUHDVHG DV H[SHFWHG ZKHQ WKH JURZWK LQWHQVLW\ ZDV GHFUHDVHG XQGHU D /' F\FOH *RQV DQG 0XU 3RVWHWDO E +RZHYHUWKHUHVSRQVHZDV PRGLILHG ZKHQ WKH LQFLGHQW LUUDGLDQFH RQ D FXOWXUH ZDVLQFUHDVHGEXWWKHWRWDOOLJKWGRVHNHSWFRQVWDQWE\ LQWURGXFLQJ D OLJKWGDUN F\FOH 7KH FHOOXODU FKORURSK\OO FRQWHQW DQG WKH VL]H RI SKRWRV\QWKHWLF
&KDSWHU )UHVKZDWHU %ORRPV XQLWV GHFOLQHG DV H[SHFWHG EXW WKH QXPEHU RI SKRWRV\QWKHWLF XQLWV LQFUHDVHG )ODPHOLQJ DQG .URPNDPS 7KLV LQFUHDVHG WKH PD[LPXP SKRWRV\QWKHWLF FDSDFLW\ DQG HQDEOHG WKH RUJDQLVP WR WDNH EHWWHU DGYDQWDJH RI WKH SHULRGLF KLJK OLJKW LQWHQVLWLHV ,EHOLQJV DOVR VXJJHVWHG WKDW WKH DELOLW\ RI Scenedesmus WR DFFOLPDWLVH PRUH UHDGLO\ WR KLJK SKRWRQ GHQVLWLHV DQG WR WDNH DGYDQWDJH RI VKRUW GXUDWLRQ SHDNV LQ LUUDGLDQFH E\ LQFUHDVLQJ LWV PD[LPXP UDWH RI SKRWRV\QWKHVLV ZDV UHVSRQVLEOH IRU LWV FRPSHWLWLYHDGYDQWDJHRYHUMicrocystis LQDKLJKO\ WXUEXOHQW HQYLURQPHQW ,W ZDV IRXQG WKDWWKH WLPH IRU DGDSWDWLRQ WR KLJK OLJKW LQWHQVLWLHV ZDV ORQJHU IRU Microcystis VR WKDW LI WKH WZR VSHFLHV ZHUH UDSLGO\ FLUFXODWLQJ WKURXJK D VWURQJ YHUWLFDO OLJKW JUDGLHQW Scenedesmus ZRXOG RXWFRPSHWH Microcystis. ,Q WKH K\SHUWURSKLF %DXW]HQ 5HVHUYRLU M. aeruginosa FRQWULEXWHV ! RI WKH WRWDO SK\WRSODQNWRQ ELRPDVV IURP -XQH WR 6HSWHPEHU .RKOHU 7KH JURZWK RI Microcystis ZDV UHVWULFWHG WR SHULRGV ZKHQ WKH ZDWHU FROXPQ ZDV VWUDWLILHG 8QGHU PL[LQJ FRQGLWLRQV SULPDU\ SURGXFWLRQ IHOO DOWKRXJK WKH ELRPDVV UHPDLQHG KLJK 6XUIDFH SKRWRV\QWKHVLV ZDV LQKLELWHG DW LQWHQVLWLHV RI ! PRO P V GXULQJ DQG LPPHGLDWHO\ DIWHU SHULRGV RI PL[LQJ EXW LQKLELWLRQ ZDV QRW REVHUYHG DIWHU WZR RU PRUH FDOPGD\V .RKOHU VXJJHVWV WKDW Microcystis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icrocystis aeruginosa IURP WKH 1HXVH 5LYHU ZKHQ WKH\ ZHUH LQFXEDWHG LQ JODVV DQG TXDUW] ERWWOHV +H UHODWHG WKH ODFN RI SKRWRLQKLELWLRQ
WR WKH SUHVHQFH RI FDURWHQRLGV ZKLFK SURWHFWHG WKH SKRWRV\VWHPV IURP SKRWRR[LGDWLRQ DQG VHUYHG DV DFFHVVRU\SLJPHQWV ,QFRQWUDVWODERUDWRU\FXOWXUHV RI Microcystis aeruginosa ZLWK ORZ OHYHOV RI FDURWHQRLGV GLVSOD\HG VLJQLILFDQW UHGXFWLRQV LQ SKRWRV\QWKHVLV ZKHQ H[SRVHG WR VXQOLJKW LQ WKH VDPH ZD\ $OWKRXJK SKRWRLQKLELWLRQ UHGXFHV WKH UDWH RI SKRWRV\QWKHVLV LWV LPSDFW YDULHV ZLWK OLJKW LQWHQVLW\ DQG H[SRVXUH SHULRG DQG PD\ EH TXLFNO\ UHYHUVHG RQ UHWXUQ WR ORZ OLJKW &RQVHTXHQWO\ WKH GHJUHH RI SKRWRLQKLELWLRQ ZLOO EH LQIOXHQFHG E\ ZDWHU PL[LQJ DQG WKH YHUWLFDO PRYHPHQW RI RUJDQLVPV )XUWKHUPRUH DV PD[LPXP UDWHV RI SKRWRV\QWKHVLV JHQHUDOO\ RFFXU DW LUUDGLDQFHV H[FHHGLQJ WKDWUHTXLUHG WR VDWXUDWH JURZWK WKH LPSDFW RI SKRWRLQKLELWLRQ RQ JURZWK UDWHV LV XQOLNHO\ WR EH GLUHFW DOWKRXJK LW LV JHQHUDOO\ SUHVXPHG WKDW SKRWRLQKLELWLRQ UHGXFHV JURZWK UDWHV /DERUDWRU\ H[SHULPHQWV XVLQJ ZKLWH OLJKW KDYH JHQHUDOO\ LQGLFDWHG WKDW DW YHU\ ORZ LUUDGLDQFH LQWHQVLWLHV F\DQREDFWHULD KDYH ORZHU OLJKW UHTXLUHPHQWV IRU JURZWK WKDQ JUHHQ DOJDH GXH WR D ORZHU PDLQWHQDQFHHQHUJ\0XU 5LFKDUGVRQHW DO ,W ZRXOG EH SUHGLFWHG IURP WKLV WKDW F\DQREDFWHULD ZRXOG RXWFRPSHWH WKH PLFURDOJDH ZKHQ OLJKW ZDV LQ YHU\ OLPLWLQJ VXSSO\ +RZHYHU EHFDXVH RI VSHFWUDO GLIIHUHQFHV LQ HQHUJ\ FDSWXUH DQG DSSDUHQW FKDQJHV LQ WKH PDLQWHQDQFH HQHUJ\ UHTXLUHPHQWGHSHQGLQJRQ OLJKWKLVWRU\ WKLV PLJKWQRW DOZD\V EH WKH FDVH *LEVRQ DQG IXUWKHU LQYHVWLJDWLRQLVUHTXLUHG :LWKLQ WKH F\DQREDFWHULD WKHUH DUH GLIIHUHQFHV EHWZHHQ VSHFLHV LQ WKH OLJKWUHTXLUHPHQWV IRU JURZWK )R\ HW DO FRPSDUHG WKH JURZWK RI Anabaena flos-aquae, Aphanizomenon flos-aquae, Oscillatoria agardhii DQG Oscillatoria redekei XQGHU GLIIHUHQW OLJKW LQWHQVLWLHV RQ D KK OLJKWGDUN F\FOH 7KH UHVXOWV VKRZHG WKDW Oscillatoria redekei LV DGDSWHG WR XVH ORZ OLJKW LQWHQVLWLHV DQG ZRXOG EH H[SHFWHG WR GRPLQDWH WKH RWKHU VSHFLHV RQO\ DW OLJKW LQWHQVLWLHV EHORZ FD PRO SKRWRQ P V DVVXPLQJ OX[ PRO SKRWRQ P V YDQ /LHUH DQG :DOVE\ $W KLJKHU LQWHQVLWLHV WKH FHOO JURZWK RI Oscillatoria redekei ZDV SKRWRLQKLELWHG 7KHVH UHVXOWV DUH LQ DFFRUG ZLWK ILHOG REVHUYDWLRQV IURP /RXJK 1HDJK ZKHUH Oscillatoria redekei KDV LWV PD[LPXP SRSXODWLRQ LQ HDUO\ VSULQJ DQG GHFOLQHV GXULQJ WKH VXPPHU PRQWKV ,Q FRQWUDVW Oscillatoria agardhii UHDFKHV LWV SRSXODWLRQ PD[LPXP LQ PLG VXPPHU )R\ HW DO $FURVV WKH VSHFLHV WKHUH ZDV D GHFUHDVLQJ HIILFLHQF\ RI OLJKWOLPLWHG JURZWK LQ WKH RUGHU O. redekei, O. agardhii, Anabaena flos-
5/2OLYHUDQG** *DQI
aquae, M. aeruginosa, ZLWK Oscillatoria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scillatoria DQG WKH VKDGH IDFWRU 7KLV VXJJHVWV WKDW WKH 2VFLOODWRULDFHDH UHVSRQG PRUH WR WKH FKDQJLQJ OLJKW FOLPDWH WKDQ WR FKDQJHV LQ QXWULHQW DYDLODELOLW\ DQG LV IDYRXUHG E\ ORZ OHYHOV RI LUUDGLDQFH VXSSRUWLQJ WKH ILQGLQJV RI )R\ HW DO 7KH GHWDLOHGDQDO\VLV RI 0XU HW DO LQ 6FKHIIHU HW DO IXUWKHU VXSSRUWV WKLV YLHZ DQG GHPRQVWUDWHV WKDW WKH GHFOLQH RI Oscillatoria SRSXODWLRQV LV PRUH FORVHO\ FRUUHODWHG ZLWK LQFUHDVHV LQ WKH PHDQ LUUDGLDQFH DV =HX=PL[ ULVHV DERYH D FULWLFDO YDOXH RI FD WKDQ ZLWK GHFOLQLQJ 73 FRQFHQWUDWLRQV $V 6DV KDV QRWHG D FKDUDFWHULVWLF RI O. agardhii SRSXODWLRQV LV WKDW WKH\ UHWUHDW WR WKH PHWDOLPQLRQ DV WKH OLJKW FOLPDWHLPSURYHV LQ UHVSRQVHWR QXWULHQWDQG WXUELGLW\ UHGXFWLRQ DQG PD\ GLVDSSHDU DOWRJHWKHU DV ZDWHU FODULW\ LPSURYHVIXUWKHU 6FKHIIHUHW DO XVHG D VLPSOH JUDSKLFDOPRGHOEDVHG XSRQ ILHOGREVHUYDWLRQV WR GHPRQVWUDWH WKDW ZLWK LQFUHDVLQJ 73 FRQFHQWUDWLRQV WXUELGLW\ LQFUHDVHV GXH WR ERWK WKH SUHVHQFH RI RWKHU DOJDH DQG VXVSHQGHG SDUWLFXODWH PDWWHU $W D FULWLFDO DWWHQXDWLRQ FRHIILFLHQW KLJK VKDGH Oscillatoria agardhii LV IDYRXUHG DQG ZLOO VXGGHQO\ GRPLQDWH WKH FRPPXQLW\ ,I WKH QXWULHQW ORDG LV WKHQ GHFUHDVHG O. agardhii ZLOO FRQWLQXH WR GRPLQDWH XQWLO WKH ELRPDVV LV UHGXFHG WR D SRLQW ZKHUH WKH FULWLFDO DWWHQXDWLRQ FRHIILFLHQW LV UHDFKHG EXW EHFDXVH RI VHOIVKDGLQJWKLV ZLOO EH DWDORZHU 73 WKDQ WKDW ZKLFK ZDV SUHVHQW ZKHQ WKH EORRP ZDV LQLWLDWHG 7KHVH FULWLFDO EUHDNSRLQWV RIIHU DSRVVLEOH H[SODQDWLRQ RI ZK\ Oscillatoria ODNHV VZLWFK VR UDSLGO\ IURP RQH VWDWH WR DQRWKHU DQG ZK\ LQ VRPH ODNHV ZLWK D KLJK EDFNJURXQG WXUELGLW\ O. agardhii PD\ QRW GLVDSSHDU LQ UHVSRQVHWR QXWULHQW UHGXFWLRQV
% 6SHFWUDO &KDQJHV :\PDQ DQG )D\ E JUHZ HLJKW VWUDLQV RI F\DQREDFWHULD XQGHU HTXLYDOHQW SKRWRQ IOX[HV RI UHG JUHHQ EOXH DQG ZKLWH OLJKW DQG IRXQG ODUJH GLIIHUHQFHV LQ WKH FHOO FRQFHQWUDWLRQV RI SKRWRV\QWKHWLF SLJPHQWV ,Q UHG OLJKW WKHUH ZDV D GHFOLQH LQ FKORURSK\OO DQG SK\FRELOLSURWHLQ FRQWHQW EXW DOO VWUDLQV JUHZ DW D VLJQLILFDQWO\ IDVWHU JURZWK UDWH WKDQ XQGHU DQ HTXLYDOHQW SKRWRQ IOX[ RI ZKLWH OLJKW )RU H[DPSOH Anabaena solitaria JUHZ WLPHV IDVWHU LQ UHG OLJKW WKDQ LQ ZKLWH OLJKW 8QGHU JUHHQ OLJKW WKH SLJPHQW FRPSRVLWLRQV ZHUH VLPLODU WR WKRVH LQ ZKLWH OLJKW EXW RQO\ WKH WZR SK\FRHU\WKULQ ULFK VWUDLQV Oscillatoria agardhii and Gloeotrichia echinulata JUHZ VLJQLILFDQWO\ IDVWHU WKDQ LQ ZKLWH OLJKW DOO RWKHU VWUDLQV JURZLQJ DW WR RI WKHLU ZKLWH OLJKW UDWH ,Q EOXH OLJKW WKH SLJPHQW FRPSRVLWLRQV ZHUH DJDLQ VLPLODU WR WKRVH LQ ZKLWH OLJKW DOWKRXJK D PDMRULW\ RI WKH SK\FRF\DQLQULFK VWUDLQV VKRZHG D UHGXFWLRQ LQ FKORURSK\OO FRQWHQW 7KH SK\FRF\DQLQ ULFKIRUPV KDG JURZWK UDWHV RI WKHLU ZKLWH OLJKW UDWH ZKLOH WKH SK\FRHU\WKULQ ULFK VWUDLQV O. agardhii DQG G. echinulata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
&KDSWHU )UHVKZDWHU%ORRPV VWRUDJHLVD YDOXDEOHDWWULEXWHHQDEOLQJFHOOVWRXWLOLVH SRROV RI QXWULHQWV WKDW DUH VSDWLDOO\ DQG WHPSRUDOO\ VHSDUDWHG VR WKDW JURZWK LV PDLQWDLQHG GXULQJSHULRGV RI QXWULHQW VFDUFLW\
; 3KRVSKRUXV 8QGHU SKRVSKRUXV OLPLWLQJ FRQGLWLRQV FHOOXODU SKRVSKRUXV FRQFHQWUDWLRQV GHFOLQH DV SKRVSKRUXV OLPLWHG JURZWK UDWH GHFOLQHV ZKLOH WKH SKRVSKRUXV XSWDNHSRWHQWLDOLQFUHDVHV $VDFRQVHTXHQFH DSXOVH RI SKRVSKRUXV GHOLYHUHG WR SKRVSKRUXV OLPLWHG FHOOV UHVXOWV LQ VXEVWDQWLDO IRUPDWLRQ RI SRO\SKRVSKDWH UHVHUYHV WKH SRO\SKRVSKDWH ³RYHUSOXV´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scillatoria agardhii DQG WKH WZR GLDWRPV Cyclotella meneghiniana DQG Asterionella formosa. 5DWKHUWKDQ D FRQVLVWHQW SK\ORJHQHWLF GLIIHUHQFH LQ SKRVSKRUXV XSWDNH DQG DFFXPXODWLRQ FKDUDFWHULVWLFV WKH JDVYDFXRODWH F\DQREDFWHULD DSSHDU WR EH DGYDQWDJHG E\ EHKDYLRXUDO IHDWXUHV RI WKHLU OLIH KLVWRU\WKDWHQDEOHWKHPWRXWLOLVHVSHFLILFSKRVSKRUXV FRQGLWLRQV PRUH HIIHFWLYHO\ )RU H[DPSOH Microcystis KDV D KLJK9PD[ IRU SKRVSKRUXV XSWDNH D ORZ PLQLPXP 3 FRQWHQW DQG D ODUJH FDSDFLW\ WR DFFXPXODWH SKRVSKRUXV .URPNDPS HW DO 7KHVH DWWULEXWHV VXJJHVW LW LV D VWRUDJH VSHFLDOLVW DQG WKLV LV LQ DFFRUGZLWK LWV DELOLW\WR UHJXODWH EXR\DQF\ WR JDLQ DFFHVV WR SKRVSKRUXV LQ GHHSHU ZDWHU OD\HUV ZKHQ HSLOLPQHWLF FRQFHQWUDWLRQV DUH ORZ *DQI DQG 2OLYHU ,QFRQWUDVWOscillatoria agardhii KDGD VLPLODUSKRVSKDWHXSWDNHUDWHEXWDODUJHUPLQLPXP3 FRQWHQW D ORZHU 9PD[ DQG ZDV OHVV DGHSW DW DFFXPXODWLQJ SKRVSKRUXV ,Q FRPSHWLWLRQ
H[SHULPHQWV ZLWK D SXOVHG VXSSO\ RI SKRVSKRUXV DW VDWXUDWLQJ FRQFHQWUDWLRQV Microcystis RXWFRPSHWHG Oscillatoria EHFDXVH RI LWV ODUJHU 9PD[ DQG PRUH HIILFLHQWXVHRISKRVSKRUXV.URPNDPSHWDO $PRUH H[WUHPH H[DPSOH RIEHKDYLRXUDO DGDSWDWLRQ LV DIIRUGHG E\ Gloeotrichia echinulata ZKHUH RIWHQ D ODUJH SHUFHQWDJH RI SODQNWRQLF SRSXODWLRQV DUH FRPSULVHG RI FRORQLHV QHZO\ UHFUXLWHG IURP WKH VHGLPHQWV %DUELHUR DQG:HOFK ,VWYiQRYLFV HW DO 3HUDNLV HW DO G. echinulata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echinulata ZHUH IRXQG LQWKHWRS FP RI VHGLPHQWV LQ DUHDV RI WKH ODNH ZLWK D GHSWK P3HWWHUVVRQHWDO 'XULQJ G. echinulata EORRPV WKH GLUHFW FRQWULEXWLRQ WR LQFUHDVHV LQ WKHSODQNWRQLF SRSXODWLRQ E\ QHZO\ UHFUXLWHG FRORQLHV IURP WKH VHGLPHQWV FDQ EHRYHU %DUELHURDQG:HOFK 3HUDNLVHW DO VXJJHVWLQJWKDWWKH\UHTXLUHODUJH VXVWDLQHG LQSXWV RIFRORQLHVIURPWKH VHGLPHQWV WRSHUVLVW 7KH PDVVLYHUHFUXLWPHQWRIEHQWKLFG. echinulata FRORQLHV WR WKH SODQNWRQ LV DVVRFLDWHG ZLWK D ODUJH WUDQVIHU RI SKRVSKRUXV LQWR WKH ZDWHU FROXPQ ,VWYiQRYLFV HW DO LQYHVWLJDWHG WKH SKRVSKRUXV XSWDNH FKDUDFWHULVWLFV RI G. echinulata LQ /DNH (UNHQ DQG VKRZHG WKDW LW ZDV XQDEOH WR XWLOLVH WKH ORZ HSLOLPQHWLF SKRVSKRUXV FRQFHQWUDWLRQV SUHVHQW GXULQJ WKHEORRP ,W GLG QRW VHHP WR UHJXODWH EXR\DQF\ WR DFFHVV SKRVSKRUXV ULFK VXEVXUIDFH ZDWHU OLNH Microcystis EXW LQVWHDG UHPDLQHG FRQWLQXRXVO\ EXR\DQW DQG WKH\ FRQFOXGHG WKDW WKH FRORQLHV DVVLPLODWH SKRVSKRUXV LQ WKH VHGLPHQWV SULRU WR WKHLU PLJUDWLRQLQWRWKHSODQNWRQDQGXVHWKLVLQWHUQDOVWRUH WR VXSSRUWSODQNWRQLF JURZWK 2Q WKH EDVLV RIWKHVH UHVXOWV G. echinulata ZDV GHVFULEHG DV DQ H[WUHPH VWRUDJHDGDSWHG VSHFLHV ZLWK LWV 3DVVLPLODWLRQ DQG JURZWKSKDVHVFRPSOHWHO\VHSDUDWHGLQWLPHDQGVSDFH ,VWYiQRYLFVHWDO 5HFUXLWPHQW RI SRSXODWLRQV IURP WKH VHGLPHQWV RFFXUV LQ VHYHUDO RWKHUEORRPIRUPLQJF\DQREDFWHULD LQFOXGLQJ Microcystis, Anabaena, Aphanizomenon
DQG Coelosphaerium DQG RFFDVLRQDOO\ WKHVH DUHODUJH HQRXJK WR LQIOXHQFH WKHLU SRSXODWLRQ G\QDPLFV 3HUDNLV HW DO +RZHYHU LQ JHQHUDO WKHVH JHQHUD LQFUHDVH IURP JURZWK LQ WKH ZDWHU FROXPQ 3UHVWRQ HW DO 7ULPEHH DQG +DUULV %DUELHUR DQG:HOFK 3HUDNLV HW DO DQG DUH UHOLDQW RQ REWDLQLQJ RQJRLQJ VXSSOLHV RI SKRVSKRUXV
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nacystis nidulans DVVLPLODWLRQ DQG JURZWKRQQLWUDWHRUDPPRQLDLVFRPSDUDEOHLIOLJKWLV VDWXUDWLQJ *XHUUHUR DQG /DUD EXW DW OLJKW LQWHQVLWLHV KDOI VDWXUDWLQJ WR SKRWRV\QWKHVLV QLWUDWH DVVLPLODWLRQLVUHGXFHG ZKLOH DPPRQLXPDVVLPLODWLRQ UHPDLQV XQFKDQJHG *DUFLD*RQ]DOH] HW DO :DUG DQG:HW]HO VKRZHGWKDWWKHORZHVWOLJKW LQWHQVLW\ DW ZKLFK JURZWK RI Aphanizomenon flosaquae, Microcystis aeruginosa DQG Anabaena jlosaquae ZRXOG RFFXU ZDV GHWHUPLQHG E\ WKH QLWURJHQ VRXUFH 2I WKH WKUHH WHVWHG DPPRQLD QLWUDWH DQG QLWURJHQ JDV DPPRQLD VXSSRUWHG WKH KLJKHVW JURZWK UDWH XQGHU DOO OLJKW UHJLPHV 7KHVH FRPSDULVRQV LQGLFDWH D SUHIHUHQFH IRU 1+ E\ F\DQREDFWHULD SDUWLFXODUO\ DW ORZ OLJKW LQWHQVLWLHV EXW WKH\ GR QRW VXSSRUWWKHQRWLRQWKDWWKLV HQDEOHV WKHPWR GRPLQDWH WKHPLFURDOJDH ZKHQ DPPRQLXPLV WKHPDMRUVRXUFH RIQLWURJHQ 6LPLODUO\ WKH UHVXOWV GR QRW VXSSRUW WKH FRQWHQWLRQ WKDW WKH PLFURDOJDH KDYH D SDUWLFXODU SUHIHUHQFHIRUQLWUDWH +RZHYHUWKH\ GR LQGLFDWHWKDW XQGHU VXERSWLPDO OLJKW FRQGLWLRQV F\DQREDFWHULDO JURZWKLVUHGXFHGZKHQWKHQLWURJHQVRXUFHLVQLWUDWH ZKHUHDV JURZWKUDWHV RIPLFURDOJDHDUHQRWDIIHFWHG 6R UDWKHU WKDQ QRQQLWURJHQ IL[LQJ F\DQREDFWHULD EHLQJ IDYRXUHG E\ DPPRQLXP DQG HXNDU\RWLF DOJDH E\QLWUDWH QLWURJHQ %ORPTYLVWHW DO LW ZRXOG VHHPWKDWWKHF\DQREDFWHULDPD\EHGLVDGYDQWDJHGE\ XVLQJQLWUDWHQLWURJHQXQGHUORZOLJKWFRQGLWLRQV
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nabaena, Aphanizomenon DQG Gloeotrichia. :KHQ FRPELQHG QLWURJHQ LV DEVHQW WKHVH RUJDQLVPV GLIIHUHQWLDWH WKLFN ZDOOHG FHOOV FDOOHG KHWHURF\VWV WKDW LVRODWH WKH QLWURJHQIL[LQJ HQ]\PH V\VWHP QLWURJHQDVH IURP LQDFWLYDWLRQ E\ R[\JHQ 7KH KHWHURF\VW SURYLGHV WKLV SURWHFWLRQE\HQKDQFHGUHVSLUDWLRQ DQG E\WKHEDUULHU RIWKHKHWHURF\VWHQYHORSH :RONHWDO 6RPH QRQKHWHURF\VWRXV VSHFLHV DUH DOVR DEOH WR IL[ 1 LQFOXGLQJ VSHFLHV RI Oscillatoria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
5/ 2OLYHUDQG** *DQI ;,, 5HVSRQVHV RI &\DQREDFWHULD WR 1 DQG 3 $ :KROH/DNH3KRVSKRUXV(QULFKPHQW 7KH FODVVLFDO ZRUN RI D QXPEHU RI DXWKRUV LQ WKH VHJ 6DNRPRWR 9ROOHQZHLGHU OHG WR WKH UHFRJQLWLRQ RI WKH LPSRUWDQFH RI LQFUHDVHG SKRVSKRUXV ORDGLQJV LQ WKH SURFHVV RIHXWURSKLFDWLRQ RIODNHV 7KHVH VWXGLHV RQ SKRVSKRUXV DQG ODWHU RQ WKH LQWHUDFWLRQ EHWZHHQ QLWURJHQ DQG SKRVSKRUXV HJ 6PLWK OHG WR HFRORJLFDO UHVHDUFK IRFXVHG RQ WKH PDQLSXODWLRQ RIZKROH ODNHV RU SRUWLRQV RI WKHP WR H[SORUH WKH UHVSRQVHV RISK\WRSODQNWRQ DEXQGDQFH DQG FRPPXQLW\ VWUXFWXUH WR QXWULHQW FRQGLWLRQV 6FKLQGOHU /XQGDQG5H\QROGV $Q LPSOLFDWLRQ RI SKRVSKRUXV ORDGLQJ PRGHOV LV WKDWGLVFKDUJLQJQXWULHQWULFKZDWHUVLQWRDZDWHUERG\ ZLOO LQFUHDVH SURGXFWLYLW\ DQG LISK\VLFDO FRQGLWLRQV SHUPLW OHDG WR WKH SUROLIHUDWLRQ DQG HYHQWXDO GRPLQDQFH RIEORRPIRUPLQJ F\DQREDFWHULD 3HDUVDOO VXJJHVWHG D UHODWLRQVKLS EHWZHHQ LQFUHDVLQJ QXWULHQWV GLVVROYHG RUJDQLF PDWWHU DQG WKH SUHVHQFH RI EORRPIRUPLQJ F\DQREDFWHULD LQ (QJOLVK ODNHV 3LFNDQG/HDQ UHFDOFXODWHG GDWDIURP*RUKDP HW DO IRU ODNHV LQQRUWKHUQ (QJODQG DQG FDPH WR WKH FRQFOXVLRQ WKDW WKHUH ZDV D VLJQLILFDQW UHODWLRQVKLS EHWZHHQ WKH PHDQ DQQXDO SK\WRSODQNWRQ ELRPDVVDQGWKDWRIWKHEORRPIRUPLQJF\DQREDFWHULD ZLWKWKHSURSRUWLRQ RIF\DQREDFWHULDLQFUHDVLQJ DV WKH WRWDO SK\WRSODQNWRQ ELRPDVV URVH DERYH J P IUHVK ZHLJKW 5H\QROGV 6WHLQEHUJ DQG +DUWPDQQ DQG 6WHLQEHUJ DQG *UXKO VXJJHVW HXWURSKLFDWLRQ HVSHFLDOO\ E\ SKRVSKRUXV RIWHQ OHDGV WR VLJQLILFDQW VKLIWV LQ SK\WRSODQNWRQ VSHFLHV FRPSRVLWLRQ WRZDUGV EORRPIRUPLQJ F\DQREDFWHULD $GGLWLRQDO HYLGHQFH WR VXSSRUW WKLV FRQFOXVLRQ FRPHV IURP (XURSH DQG 1RUWK $PHULFD +DUULV &XOOHQ DQG )RUVEHUJ 6DV &RRNH HW DO DV ZHOO DV IURP $XVWUDOLD DQG $IULFD 7KH3HHO+DUYH\(VWXDU\LQ:HVWHUQ$XVWUDOLDDQGWKH +DZNHVEXU\ 5LYHU LQ 1HZ 6RXWK :DOHV ERWK LOOXVWUDWH WKDW LQFUHDVHV LQ QXWULHQW ORDGV IURP HLWKHU DJULFXOWXUDO UXQRIIRU VHZDJH WUHDWPHQWSODQWV UHVXOWV LQ H[WHQVLYH SRSXODWLRQV RI EORRPIRUPLQJ F\DQREDFWHULD LQ WKHVH H[DPSOHV Nodularia spumigenea 3ODWH H DQGMicrocystis aeruginosa Anabaena VSUHVSHFWLYHO\ *HRUJH %UDGE\ &XOOHQ +XPSKULHV 5RELQVRQ
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
% %LRPDVV5HVSRQVHWR35HPRYDO /DNH0F,OZDLQH QHDU WKH FLW\ RI+DUDUH =LPEDEZH H[SHULHQFHG DQWKURSRJHQLF HXWURSKLFDWLRQ DV D UHVXOW RI VHZDJH GLVFKDUJH LQWR WKH ODNH 7KRUQWRQ $V QLWURJHQ DQG SKRVSKRUXV FRQFHQWUDWLRQV URVH WR WLPHV WKHLU RULJLQDO OHYHOV FKORURSK\OOD FRQFHQWUDWLRQV SHDNHG DWFD PJ P DQG GLG QRW IDOOEHORZFDPJP WKURXJKRXW 'XULQJ WKLVSHULRGWKHODNHZDVGRPLQDWHGDOPRVWH[FOXVLYHO\ E\ Microcystis aeruginosa DQG Anabaena flos-aquae. $IWHU GLYHUVLRQ RI WKH VHZDJH WR SDVWXUH LUULJDWLRQ EHWZHHQ DQG WKH VROXEOH UHDFWLYH SKRVSKDWH OHYHOV IHOO E\ DQ RUGHU RI PDJQLWXGH DQG FKORURSK\OODFRQFHQWUDWLRQVIHOO WR DPHDQRI PJ P $OWKRXJK WKH ODNH VWLOO VXSSRUWHG SRSXODWLRQV RI F\DQREDFWHULD DV ZHOO DV Melosira granulata, WKHUH ZDV D UHDSSHDUDQFH RI VLJQLILFDQW SRSXODWLRQV RI RWKHUHXNDU\RWLFJHQHUD 7KH/DNH0F,OZDLQHH[SHULHQFHVXJJHVWVWKDWZKHQ QXWULHQW ORDGV DUH UHGXFHG SK\WRSODQNWRQ ELRPDVV GHFUHDVHV EXW WKH FRQFOXVLRQ WKDW F\DQREDFWHULD ZLOO DOVR GLVDSSHDU LV D FRQYHQLHQW UDWKHU WKDQ D UHDOLVWLF RQH 1HYHUWKHOHVV WKHUH DUH VLWXDWLRQV ZKHUH WKLV GRHV RFFXU )RU H[DPSOH DV DUHVXOWRIWKH GLYHUVLRQ RI VHZDJH IURP /DNH :DVKLQJWRQ (GPRQGVRQ DQG /HKPDQ 6HLS HW DO WKH SHUFHQWDJH RI WRWDO SKRVSKRUXV IURP VHZDJH ZDV UHGXFHG IURP FD WR ]HUR DQG WKH WRWDO SKRVSKRUXV FRQWHQW RIWKH ODNH IHOO IURP [ NJ WR [ NJ 7KLV UHVXOWHG LQ D GHFUHDVH LQ PD[LPXP FKORURSK\OOD FRQFHQWUDWLRQV IURP FD PJ P WR OHVVWKDQ PJP &RQFXUUHQWO\WKH SURSRUWLRQ RI WKH WRWDO SK\WRSODQNWRQ ELRPDVV DWWULEXWDEOH WR ILODPHQWRXV F\DQREDFWHULD
Oscillatoria rubescens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eratium VSS EXW GXH WR LQWHQVH SDUDVLWLVPE\Aphanomycopsis cryptica, DELIODJHOODWH IXQJXVWKHCeratium GLPLQLVKHGIURP ZDVZHUH UHSODFHG E\ Aphanizomenon flos-aquae IR gracile, Anabaena flos-aquae DQG A. solitaria. 6XEVHTXHQW FKDQJHV LQ WKH WLPLQJ DQG PD[LPD RI WKH F\DQREDFWHULDZHUH DWWULEXWDEOH WR WKH LQWHQVH JUD]LQJ E\WKH SURWR]RDQ FLOLDWHNassula aurea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
5/ 2OLYHUDQG** *DQI
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cenedesmus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
   Â
&KDSWHU )UHVKZDWHU %ORRPV
Fig. 9 +\SRWKHWLFDOH[DPSOHRIWKHUHVSRQVHRIODNHFKORURSK\OOD FRQFHQWUDWLRQWRUHGXFWLRQVLQSKRVSKRUXV FRQFHQWUDWLRQ DFFRUGLQJ WR6DV 7KHWKHRUHWLFDOFKORURSK\OOSKRVSKRUXV UHODWLRQVKLS LV VKRZQ E\ WKH VWUDLJKW OLQH GDVKHG ZKLOH WKH DFWXDO UHVSRQVH IROORZV WKH VROLG OLQH $W KLJK SKRVSKRUXV FRQFHQWUDWLRQV WKH GLIIHUHQFH EHWZHHQ WKH WKHRUHWLFDO DQG DFWXDO FKORURSK\OO FRQFHQWUDWLRQV LV GXH WR UHVWULFWLRQ RIWKH DOJDO ELRPDVV E\ VRPH RWKHUIDFWRUDQGSKRVSKRUXVLVSUHVHQWLQH[FHVV
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
ZKHUH &% DQG &$ GHQRWH WKH JURZWKVHDVRQDO DYHUDJH FKORURSK\OOD FRQFHQWUDWLRQV SUH DQG SRVW UHVWRUDWLRQ UHVSHFWLYHO\ 3% DQG 3$ WKH SUH DQG SRVWUHVWRUDWLRQ YDOXHV RI WKH ZKROHODNH DQQXDO PHDQ WRWDO SKRVSKRUXV FRQFHQWUDWLRQ DQG m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scillatoria VSS WKH GDWD VXJJHVW WKDW WKH WUHQG RI LQFUHDVLQJ FHOO QXPEHUV ZLWK LQFUHDVLQJ QXWULHQW VXSSO\ LV PRUH RU OHVV UHYHUVLEOH DV WKH QXWULHQW ORDG GHFUHDVHV EHORZ WKUHVKROG YDOXHV ,Q VKDOORZ ODNHV WKLVWKUHVKROGDSSHDUVWREH PJP )53EXW LV UDWKHU ORZHU IRU GHHS ODNHV DW PJ P 6DV
' 5HVSRQVHVRI&\DQREDFWHULD WR3 5HPRYDO 7KH VORSH RI WKH OLQHDU ORJORJ UHODWLRQVKLS EHWZHHQ FRQFHQWUDWLRQV RIFKORURSK\OOD DQG WRWDO SKRVSKRUXV ZHUH XVHG WR DVVHVV ZKHWKHU RU QRW F\DQREDFWHULD UHVSRQGHG WR FKDQJHV LQ WKH 73 FRQFHQWUDWLRQ LQ D VLPLODU RU GLIIHUHQW PDQQHU WR PLFURDOJDH 7KH IUHTXHQF\RIRFFXUUHQFHRISDUWLFXODUVORSHV ZDV XVHG WR FRPSDUH WKH WUDMHFWRU\ RIWKH UHODWLRQVKLS EHWZHHQ FKORURSK\OOD DQG 73 IRU F\DQREDFWHULD DQG IRU PLFURDOJDH6HLSHWDO 3RVLWLYHVORSHVRI LQGLFDWHG D UHODWLRQVKLS EHWZHHQ FKORURSK\OOD DQG 73 6ORSHV QHDU LQGLFDWH VPDOO UDWLRV WKDW VXJJHVWFKDQJHV LQ73FRUUHVSRQG ZLWKPXFK VPDOOHU
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scillatoria VSS ZKLFK HLWKHU GHFUHDVHG VLJQLILFDQWO\ GLVDSSHDUHG RU UHWUHDWHG WR WKH PHWDOLPQLRQ DV ZDWHU WUDQVSDUHQF\ LQFUHDVHG LQ UHVSRQVH WR UHGXFWLRQV LQELRPDVV /HVV REYLRXV ZDV WKH UHVSRQVH RI F\DQREDFWHULDO VSHFLHV VXFK DV Aphanizomenon VSS Anabaena VSS DQG Microcystis VSS WKDW FDQ IRUP VXUIDFH EORRPV ,Q D VLJQLILFDQW QXPEHURIFDVHVZKHUHWHPSHUDWXUHVZHUHIDYRXUDEOH WKHVHWHQGHGWRLQFUHDVHDVOscillatoria VSSGHFUHDVHG LQ DEXQGDQFH DQG WKLV ZDV FRUUHODWHG ZLWK DGHFOLQH LQWKHUDWLR RI73 Zeu/Zmix LH WKHUDWLR RISKRVSKRUXV WRDYDLODEOHOLJKW 7KLV VSHFLHVUHVSRQVHWR ODNHUHVWRUDWLRQ LV DSULPH DYHQXH IRU IXWXUH UHVHDUFK ,W LV HYLGHQW WKDW VSHFLHV FKDQJHV DUH QRW VLPSO\ D IXQFWLRQ RI WKH UHODWLYH JURZWKUDWHVRIGLIIHUHQWVSHFLHVDOWKRXJKWKLVPD\EH FRQWULEXWRU\ UDWKHU LW LV WKH LQWHUDFWLRQ EHWZHHQ D QXPEHU RIELRORJLFDO IHDWXUHV DQG WKH VWDELOLW\ RIWKH ZDWHUFROXPQDVZHOODVWKHTXDOLW\RIWKHXQGHUZDWHU OLJKWILHOG
( %LRDVVD\V /DNH(QULFKPHQWDQG 1LWURJHQ /LPLWDWLRQ (OVHUHWDO UHYLHZHG1RUWK$PHULFDQODNHV IURP ZKLFK GDWD ZHUH DYDLODEOH RQ WKH UHVSRQVH RI DOJDO ELRPDVV FKORURSK\OO &IL[DWLRQ FHOO FRXQWV LQ YLYR FKORURSK\OO IOXRUHVFHQFH WR QXWULHQW HQULFKPHQW RI 1 3 RU 13 YHUVXV D FRQWURO QR DGGLWLRQ LQ HQULFKPHQW ELRDVVD\V DQG ZKROHODNH H[SHULPHQWV 7KH DYHUDJHFRQFHQWUDWLRQVDGGHGZHUH 0 1 DQG 0 3 6LPXOWDQHRXV 1 DQG 3 HQULFKPHQW LQ ELRDVVD\V QHDUO\ DOZD\V HOLFLWHG D
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cenedusmus DQG RWKHU DOJDH XQDEOH WR IL[ DWPRVSKHULF QLWURJHQ +RZHYHU ZKHQ WKH 13 UDWLR ZDV FXWEDFNWR D EORRP RI WKH QLWURJHQIL[LQJ Aphanizomenon gracile RFFXUUHGZKLFKIL[HGVLJQLILFDQWDPRXQWVRIQLWURJHQ $ VLPLODU UHVSRQVH ZDV QRWHG LQ /DNH ZKHQ IHUWLOLVHG ZLWK D7173UDWLR RI DOWKRXJKKHUHWKH GRPLQDQWJHQXVZDVAnabaena. ,Q WKH VHFRQG H[DPSOH 6WRFNQHU DQG 6KRUWUHHG WKH &OD\RTXRW $UP RI ROLJRWURSKLF .HQQHG\ /DNHH ZDV HQULFKHG ZLWK VROXWLRQV RI 1+12 DQG 1+ +32 R WR PJ 3 P ZHHN WR JLYH 13
PRODUUDWLRVWKDWURVH IURP FD E\ ZHLJKW LQ WKH ILUVW \HDU RIIHUWLOLDVDWLRQ WR LQ DQGWKHQIXUWKHULQFUHDVHGWRDQGRYHUWKH QH[WWKUHH \HDUVAnabaeuna circinalis FRQFHQWUDWLRQV ZHUH FHOOV PO SULRU WR IHUWLOLVDWLRQ EXW UHDFKHG !FHOOVPO LQ DQGVXEVHTXHQWO\IHOOWRDQ DYHUDJH FHOO FRQFHQWUDWLRQ RI PO LQ &RQFXUUHQWO\ WKH JURZLQJ VHDVRQ DYHUDJH VHFFKL GHSWKV GHFUHDVHG IUSP GRZQ WR P LQ EHIRUH GHHSHQLQJ DJDLQ WR P LQ 'XULQJ WKLV SHULRG WKH WRWDO SKRVSKRUXV FRQFHQWUDWLRQ LQFUHDVHG WR D PD[LPXP RI RQO\ PJ P 7KH DXWKRUV FRQFOXGHG WKDW WKH ORZ LQLWLDO 13 UDWLRV LQ WKH SUHVHQFH RI ULVLQJ SKRVSKRUXV FRQFHQWUDWLRQV SURPRWHG A. circinalis EXW DV WKH13 UDWLR URVH DQG QLWURJHQ ZDV QR ORQJHU OLPLWLQJ WKHQ RWKHU VSHFLHV SUHGRPLQDQWO\ Synechococcus VS EHFDPHGRPLQDQW
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
Table 2. 7KHSHUFHQWSUREDELOLWLHVRIRFFXUUHQFHRIIRXUF\DQREDFWHULDXQGHUGLIIHUHQWPL[LQJFRQGLWLRQV 0 DQGWKHSUHYDLOLQJ7173 UDWLRV+DUULV86(3$GDWD
6SHFLHV
0
7173
0 7173
0
7173
0
Anabaena Aphanizomenon Microcystis Oscillatoria
67
!
50
!
100
!
7173
0 ! 7173 38
5/ 2OLYHUDQG** *DQI
Anabaena's GLVWULEXWLRQ VSDQQHG WKH FRPSOHWH UDQJH RI 0 IURP VWDEOH WR KLJKO\ WXUEXOHQW ZDWHU FROXPQV EXW ZDV IDYRXUHG E\ 7173 UDWLRV RI VXJJHVWLQJWKDWLWFRPSHWHVEHVWLQKDELWDWV ZKHUHWKH IOX[ RI 1 LQWR WKH ZDWHU PD\EH OLPLWLQJ Aphanizomenon DOVR RFFXUUHG LQ KDELWDWV ZLWK ORZ 7173 UDWLRV FRQVLVWHQW ZLWK WKH REVHUYDWLRQ WKDW ZKHQWKHORDGLQJ ZDV UHGXFHG IURP WR LQWKH ([SHULPHQWDO/DNHV $UHDDEORRPRIAphanizomenon GHYHORSHG ZKHUHDV LW KDG QRW EHHQ REVHUYHG EHIRUH 6FKLQGOHU Aphanizomenon KDG D FOHDU SUHIHUHQFH IRU VWDEOH VXPPHU FRQGLWLRQV DOWKRXJK LW DOVR RFFXUUHG DW 0! 7KH JURZWK RI WKH WZR ILODPHQWRXV VSHFLHV RYHU D ZLGH UDQJH RI PL[LQJ UHJLPHV PD\ UHIOHFW WKHLU DELOLW\ WR RFFXU ERWK DV VLQJOHILODPHQWVDQGDVODUJHDJJUHJDWHGFRORQLHV DQG LV FUXGHO\ LQ DFFRUG ZLWK WKH FODVVLILFDWLRQ GLVFXVVHG LQWKHHDUOLHUVHFWLRQ +DELWDWVVWUXFWXUHGE\WXUEXOHQW PL[LQJ Oscillatoria DSSHDUHGLQGHSHQGHQWRI7173 UDWLRV EXW SUHIHUUHG OHVV VWDEOH ZDWHU FROXPQV VXJJHVWLQJ WKDW PRVW VDPSOHV ZHUH IURP ZHOO PL[HG ODNHV DQG WKDW WKRVH ZLWK PHWDOLPQHWLF SRSXODWLRQV ZHUH QRW ZHOO UHSUHVHQWHG LQ WKH GDWD ,Q FRQWUDVW Microcystis RFFXUUHG LQ VWDEOH KDELWDWV ZKHUH WKHUH ZDV OLWWOH HYLGHQFHIRUQLWURJHQ OLPLWDWLRQ
;,,,
,QRUJDQLF&DUERQ
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
3K\WRSODQNWRQ SKRWRV\QWKHVLV UHPRYHV &2 IURP VROXWLRQUHVXOWLQJLQGLVUXSWLRQWRWKHHTXLOLEULXPDQG DULVH LQS+ WKH PDJQLWXGH RIZKLFK GHSHQGV RQ WKH EXIIHULQJ LQWHQVLW\ RI WKH ZDWHU 6KDSLUR ,W KDV EHHQ VKRZQ E\ PDQLSXODWLQJ &2 FRQFHQWUDWLRQ DQG S+ LQ HQFORVXUH DQG ODNH H[SHULPHQWV WKDW F\DQREDFWHULD DUH EHWWHU DGDSWHG WR WDNHDGYDQWDJH RI ORZ &2 FRQFHQWUDWLRQV WKDQ VRPH EXW QRW DOO PLFURDOJDH 7DOOLQJ 6KDSLUR 6RPH F\DQREDFWHULD KDYH EHHQ VKRZQ WR XWLOLVH +&2 7DOOLQJ EXW DV WKH\ JHQHUDOO\ FRQWDLQ FDUERQLF DQK\GUDVH WKDW FDWDO\VHV WKH GHK\GUDWLRQ RI ELFDUERQDWH LW LV OLNHO\ WKDW &2 LV XVHG ZLWKLQ WKH FHOO +RZHYHU WKHVH IHDWXUHV DUH QRW XQLTXH WR WKH F\DQREDFWHULDDQGVRPHHXNDU\RWLFDOJDHDUHDOVRDEOH WRXWLOLVHYHU\ORZ&2 FRQFHQWUDWLRQV7DOOLQJ DQG VRPH FDQ DFFHVV +&2 DV WKH\ DOVR FRQWDLQ FDUERQLF DQK\GUDVH 7KH GHEDWH RYHU WKH LPSRUWDQFH RIFRPSHWLWLRQ IRU LQRUJDQLF FDUERQ KDV EHHQ IXUWKHU FRPSOLFDWHG E\ WKH GLVFRYHU\ RI WKH FDUERQ FRQFHQWUDWLQJPHFKDQLVPLQSK\WRSODQNWRQ ,WLV VWLOO XQFOHDUZKHWKHUWKLV PHFKDQLVPLV PRUH EHQHILFLDO LQ F\DQREDFWHULD RU PLFURDOJDH DQG VR WKH UROH RI LQRUJDQLFFDUERQOLPLWDWLRQRQFRPSHWLWLRQLVGLIILFXOW WRDVVHVV )LHOG H[SHULPHQWV ZKHUH PDQLSXODWLRQV RI FDUERQ DQG S+ KDYH UHVXOWHG LQ VKLIWV LQ VSHFLHV GRPLQDQFH WRF\DQREDFWHULDXQGHUFRQGLWLRQVRILQRUJDQLFFDUERQ OLPLWDWLRQ 6KDSLUR VXJJHVW WKDW D ORZ DYDLODELOLW\ RI &2 FDQ DGYDQWDJH WKH JDVYDFXRODWH F\DQREDFWHULD 7KLV DGYDQWDJH ZRXOG EH UHHQIRUFHG GXULQJ SHULRGV RI ODUJH F\DQREDFWHULDO EORRPV DVVLVWLQJ WKHVH RUJDQLVPV WR PDLQWDLQ GRPLQDQFH +RZHYHU WKH EORRPIRUPLQJ F\DQREDFWHULD DUH QRW UHVWULFWHG WR HXWURSKLF ZDWHUV DQG FDQ GRPLQDWH WKH FRPPXQLWLHVRIOHVVSURGXFWLYHODNHVHYHQWKRXJKWKH WRWDO SK\WRSODQNWRQ ELRPDVV LV ORZ &RQYHUVHO\ F\DQREDFWHULDFDQ DOVR EHIRXQG WRGRPLQDWHLQVRPH ZHOOPL[HG VKDOORZ ZDWHUV ZKHUH LQRUJDQLF FDUERQ OLPLWDWLRQ LV FRQVLGHUHG XQOLNHO\ 6WHLQEHUJ DQG *UXKO 6FKHIIHU HW DO 7KHVH REVHUYDWLRQV ZRXOG DUJXH DJDLQVW &2 OLPLWDWLRQ EHLQJ HVVHQWLDO IRU WKH IRUPDWLRQ RI F\DQREDFWHULDO EORRPV EXW ZKDWHYHU WKH ILQDO RXWFRPH UHJDUGLQJ GLUHFW FRPSHWLWLRQ IRU LQRUJDQLF FDUERQ DQ\ OLPLWDWLRQZLOOVWLOOKDYHDPDMRULQIOXHQFHWKURXJKLWV LPSDFW RQ EXR\DQF\ UHJXODWLRQ 8QGRXEWHGO\ PRUH LQIRUPDWLRQ LV UHTXLUHG RQ WKH LQRUJDQLF FDUERQ UHTXLUHPHQWVRIWKHSK\WRSODQNWRQ
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aphnia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aphnia galeata mendotae ZKLFK FRQWLQXHG LQWR HDUO\ VXPPHU EHIRUH EHFRPLQJ VXEGRPLQDQWWR FDODQRLGV DQG F\OFRSRLGV Daphnia KDG WKH ODUJHVW LPSDFW RQ WKH VSULQJ SK\WRSODQNWRQ DQG ZDV FRQVLGHUHG PDLQO\ UHVSRQVLEOH IRU WKH FOHDU ZDWHU SKDVH ,W ZDV OHVV HIIHFWLYH RQ WKH ODUJH LQHGLEOHSK\WRSODQNWRQ VSHFLHV WKDWZHUHGRPLQDQWLQ VXPPHU SDUWLFXODUO\ F\DQREDFWHULD DQG WKH GLQRIODJHOODWH Ceratium. ,Q /DNHV +XPH DQG 'DUWPRXWK ERWK PDQPDGH ODNHV ORFDWHG LQ VRXWKHDVWHUQ $XVWUDOLD PDQLSXODWLRQV RIWKH ODUJH FODGRFHUDQV Daphnia DQG Diaphanosoma DQG WKH ODUJH FRSHSRG Boeckella KDG QHJDWLYH HIIHFWV RQ WKH SK\WRSODQNWRQ ELRPDVV
ZKLOH VPDOOHU FRSHSRGV Mesocyclops DQG Calamoecia KDG OLWWOH LPSDFW RU RFFDVLRQDOO\ D SRVLWLYH HIIHFW 0DWYHHY DQG 0DWYHHYD ,Q ERWK UHVHUYRLUV WKH YDULDWLRQ LQ Daphnia carinata DORQH FRXOG DFFRXQWIRU RIWKH YDULDQFH LQ WRWDO SK\WRSODQNWRQ ELRYROXPH &RORQLHV RI Microcystis P GLDPHWHU ZHUH JUD]HG LQ IHHGLQJ WULDOV E\ ERWK Daphnia DQG Boeckella ZLWKRXW DQ\ VXJJHVWLRQ RI VHOHFWLYLW\ :KHQ Microcystis GRPLQDWHG LQ WKH ODNHV HQFORVXUH H[SHULPHQWV VKRZHG D QHJDWLYH UHVSRQVH RI ELRPDVV WR Daphnia DQG Boeckella FRQFHQWUDWLRQV 7KHVH VWXGLHV DUH LQ JHQHUDO DFFRUG ZLWK PDQ\ RWKHUV WKDW KDYH IRXQG FRUUHODWLRQV EHWZHHQ WKH DEXQGDQFH RI ]RRSODQNWRQ JUD]HUV DQG WKH WLPLQJ RI F\DQREDFWHULDO EORRPV +DQH\ ,Q JHQHUDO WKH UHGXFHG JUD]LQJ RQ F\DQREDFWHULD LV DVVRFLDWHG ZLWK ODUJH VL]H KLJK GHQVLW\ DOOHORSDWKLF FRPSRXQGV DQG SRRU DVVLPLODELOLW\ 7KH PD[LPXP VL]H RI IRRG SDUWLFOHV\ WDNHQLQE\Daphnia VSHFLHVLVDIXQFWLRQ RI WKH DQLPDO¶V OHQJWK / DV GHVFULEHG E\ %XUQV y
L
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phanizomenon flos-aquae ZKLFK RFFXUV DV D ODUJH IODNH DQG GRPLQDWHV LQ WKH SUHVHQFH RI Daphnia pulex, ZLWK ILODPHQWRXV Aphanizomenon elenkinii ZKLFK GRPLQDWHG RQO\ ZKHUH D. pulex
 Â
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scillatoria WR FRPSOH[ ODUJH FRORQLHV LQ Microcystis. 7KHVPDOO VL]H DQG VORZ VLQNLQJIORDWLQJ YHORFLWLHV RI Oscillatoria SHUPLWV LW WR SRVLWLRQ LWVHOI DW LWV SUHIHUUHG ORZ OLJKW HQYLURQPHQW ZKHUH LW RIWHQ IRUPV GHHS ZDWHU PD[LPD LQ VWUDWLILHG ODNHV 7KH ODUJH FRORQLDO IRUPVRIMicrocystis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icrocystis PD\ WDNH WZR GD\V WR DGDSW WR KLJK LUUDGLDQFHV EXW JUHHQ DOJDH HJ Scenedesmus UHVSRQG PRUH TXLFNO\ ZKLFK SURYLGHV DQ H[SODQDWLRQRIZK\Scenedesmus PD\GRPLQDWHLQ WXUELG FRQWLQXRXVO\PL[HG VKDOORZ ZDWHUERGLHV +RZHYHU WKH HQHUJ\ UHTXLUHG WR V\QWKHVLVH JDV YHVLFOHV LQ WKH FRPSHWLQJ F\DQREDFWHULD PD\ EH VXIILFLHQW WR SXVK WKH HQHUJ\ EDODQFH LQ IDYRXU RI JUHHQ DOJDH XQGHU ORZ OLJKW FRQGLWLRQV 7KH SK\FRHU\WKULQ ULFK VSHFLHV VXFK DV Oscillatoria agardhii DQG Gloeotrichia echinulata DUH ORZ OLJKW DGDSWHG KDYLQJ D ZLGH VSHFWUDO DFWLYLW\ DQG KDYH DQ DGYDQWDJH RYHU ERWK PLFURDOJDH DQG SK\FRHU\WKULQ SRRU F\DQREDFWHULD HVSHFLDOO\ LQ ZDWHU ERGLHV ZKHUH WKH PRVW SHQHWUDWLQJ ZDYHEDQGLVQP 7KHUH LV QR HYLGHQFH WR VXSSRUW WKH K\SRWKHVLV WKDW EORRPIRUPLQJ F\DQREDFWHULD SUHIHU KLJKHU +RZHYHU LQ WHPSHUDWXUHV WKDQ PLFURDOJDH Microcystis FDUERK\GUDWH PHWDEROLVP LV GHSUHVVHG
&KDSWHU )UHVKZDWHU%ORRPV DW WHPSHUDWXUHV EHORZ & ZKLFK LQIOXHQFHV EXR\DQF\UHJXODWLRQ 7KHUH GRHV QRW DSSHDU WR EH DQ\ FRQVLVWHQW SK\ORJHQHWLF GLIIHUHQFHV LQ SKRVSKRUXV NLQHWLFV EHWZHHQ WKH F\DQREDFWHULD DQG WKH PLFURDOJDH 6RPH F\DQREDFWHULD DUH DEOH WR VWRUH VLJQLILFDQW TXDQWLWLHV RI SKRVSKRUXV EXW LQ WKH FDVH RI Microcystis DQG Gloeotrichia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per se LQIOXHQFH WKH RFFXUUHQFH RI SODQNWRQLF F\DQREDFWHULD +RZHYHU QLWURJHQ IL[HUVHJAnabaena VSS PD\RFFXUif WKH7173 UDWLR LV ORZ DQG FRUUHFWO\ SUHGLFWV WKDW LQRUJDQLF QLWURJHQ LV OLPLWLQJ 2Q WKH RWKHU KDQG Gloeotrichia echinulata PD\ GRPLQDWH ZKHQ 7173 UDWLRV DUH KLJK EXW WKH 3FRQWHQW RI WKH ZDWHUFROXPQLVLQVXIILFLHQWWRVXSSRUWJURZWK 9HU\ IHZ GDWD DUH DYDLODEOH WR WHVW FULWLFDOO\ WKH SURSRVDO WKDW F\DQREDFWHULD DUH EHWWHU DEOH WR
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
5/ 2OLYHU DQG ** *DQI
Fig. 10 )ORZ FKDUW VXPPDULVLQJ SURPLQHQW HQYLURQPHQWDO FKDUDFWHULVWLFV VXSSRUWLQJ WKH GHYHORSPHQW RI F\DQREDFWHULDO EORRPV DQG VHOHFWLQJ IRU SDUWLFXODU JHQHUD 7KH WH[W SURYLGHV IXUWKHU GHWDLO RQ VRPH RIWKHVH FRPSRQHQWV .H\ %&/$' ELRPDVV RIFODGRFHUDQV %3+<72 ELRPDVVRISK\WRSODQNWRQ ]HX HXSKRWLFGHSWK ]PL[ GHSWKRIPL[LQJX VKHDUYHORFLW\ 9IORDWLQJRUVLQNLQJYHORFLW\RIF\DQREDFWHULD WZ WLPH WKDWWKHZLQGEORZV/ODNHIHWFKFV VXUIDFHFXUUHQWVSHHG
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scillatoria agardhii YDU isothrix. 0LWW9HUK ,QW9HUHLQ/LPQRO %URRNHV -' *DQI**DQG %XUFK 0' 6HSDUDWLRQ RIIRUPV RI Microcystis IURP Anabaena LQ PL[HG SRSXODWLRQV E\ WKH DSSOLFDWLRQRISUHVVXUH $XVW-0DU)UHVKZDWHU5HV %XUFK0' %DNHU3' 6WHIIHQVHQ'$ %XUVLOO'% %DLQ'% *DQI ** DQG %URRNHV -' &ULWLFDO IORZ DQG EORRPV RI WKH F\DQREDFWHULXP Anaebaena circinalis LQ WKH5LYHU0XUUD\ 6RXWK $XVWUDOLD Proceedings of the Environmental Flows Seminar. $XVWUDOLDQ:DWHUDQG:DVWHZDWHU$VVRFLDWLRQ&DQEHUUD %XPV &: 7KH UHODWLRQVKLS EHWZHHQ ERG\ VL]H RI ILOWHU IHHGLQJ FODGRFHUD DQG WKH PD[LPXP VL]H RI SDUWLFOH LQJHVWHG /LPQRO2FHDQRJU %XPV &: ,QVLJKWV LQWR ]RRSODQNWRQF\DQREDFWHULD LQWHUDFWLRQV GHULYHG IURP HQFORVXUH VWXGLHV 1 = - 0DU )UHVKZDWHU5HV &DQWHU +0 +HDQH\ 6, DQG /XQG -:* 7KH HFRORJLFDO VLJQLILFDQFH RI JUD]LQJ RQ SODQNWRQLF SRSXODWLRQV RI F\DQREDFWHULDE\WKHFLOLDWHNassula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nacysris nidulans. 3ODQWD &XOOHQ3 5HSRUWIURPH[SHUWVHPLQDURQDOJDOEORRPVLQWKH +DZNHVEXU\ 5LYHU $ UHSRUW WR WKH +DZNHVEXU\1HSHDQ &DWFKPHQW0DQDJHPHQW7UXVW16:$XVWUDOLD SS
&XOOHQ 3 DQG )RUVEHUJ & ([SHULHQFHV ZLWK UHGXFLQJ SRLQW VRXUFHVRISKRVSKRUXVWRODNHV+\URELRORJLD 'DPHUYDO7 &DVWHWV $0+RXPDUG - DQG7DQGHDX GH0DUVDF 1 *DV YHVLFOH V\QWKHVLV LQ WKH F\DQREDFWHULXP Pseudanabaena VS RFFXUUHQFHRIDVLQJOHSKRWRUHJXODWHGJHQH 0RO 0LFURELRO 'DYH\0& DQG:DOVE\ $( 7KHIRUPUHVLVWDQFHRIVLQNLQJ DOJDOFKDLQV%U3K\FRO- 'HDFRQ & DQG :DOVE\ $( *DV YHVLFOH IRUPDWLRQ LQ WKH GDUNDQGLQOLJKWRIGLIIHUHQWLUUDGLDQFHVE\ WKHF\DQREDFWHULXP Microcystis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cenedesmus protuberans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scillatoria VSHFLHV %U3K\FRO- )UDQFLV* 3RLVRQRXV$XVWUDOLDQ/DNH 1DWXUH/RQGRQ *DQI ** 'LXUQDO PL[LQJ DQG WKH YHUWLFDO GLVWULEXWLRQ RI SK\WRSODQNWRQ LQ D VKDOORZ HTXDWRULDO ODNH /DNH *HRUJH 8JDQGD -(FRO *DQI ** $Q HFRORJLFDO UHODWLRQVKLS EHWZHHQ Aphanizomenon DQG Daphnia pulex. $XVW- 0DU)UHVKZDWHU5HV *DQI** DQG 2OLYHU 5/ 9HUWLFDO VHSDUDWLRQ RI OLJKW DQG DYDLODEOH QXWULHQWV DV D IDFWRU FDXVLQJ UHSODFHPHQW RI JUHHQ
DOJDH E\ EOXHJUHHQ DOJDH LQ WKH SODQNWRQ RID VWUDWLILHG ODNH - (FRO *DQI ** 2OLYHU5/ DQG:DOVE\ $( 2SWLFDO SURSHUWLHV RI JDVYDFXRODWH FHOOV DQG FRORQLHV RI Microcystis LQ UHODWLRQ WR OLJKW DWWHQXDWLRQ LQ D WXUELG VWUDWLILHG UHVHUYRLU 0RXQW %ROG 5HVHUYRLU 6RXWK$XVWUDOLD $XVW-0DU)UHVKZDWHU5HV *DQWW ( 2KNL . DQG )XMLWD < Trichodesmium thiebautii; VWUXFWXUH RI D QLWURJHQIL[LQJ PDULQH EOXHJUHHQ DOJD &\DQRSK\WD 3URWRSODVPD *DUFLD*RQ]DOH] 0 6LYDN 01 *XHUUHUR 0* 3UHLVV - DQG/DUD& 'HSUHVVLRQ RI FDUERQ IORZ WR WKHJO\FRJHQ SRRO LQGXFHG E\ QLWURJHQ DVVLPLODWLRQ LQ LQWDFW FHOOV RI Anacystis nidulans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cenedesmus protuberans )ULWVFK LQ OLJKW OLPLWHG FRQWLQXRXV FXOWXUHV ZLWK D OLJKWGDUN F\FOH $UFKLY Hydrobiol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nabaena flos-aquae. -([S%RW *URVVPDQ $5 %KD\D ' $SW .( DQG .HKRH '0 /LJKW KDUYHVWLQJ FRPSOH[HV LQ R[\JHQLF SKRWRV\QWKHVLV'LYHUVLW\ FRQWURO DQGHYROXWLRQ $QQXDO5HYLHZ RI*HQHWLFV *XHUUHUR 0* DQG /DUD & $VVLPLODWLRQ RI LQRUJDQLF QLWURJHQ ,Q )D\ 3 DQG YDQ %DDOHQ & HGV 7KH&\DQREDFWHULD SS (OVHYLHU $PVWHUGDP +DQH\ -) $QLQsitu H[DPLQDWLRQ RI WKHJUD]LQJDFWLYLWLHVRI QDWXUDO]RRSODQNWRQFRPPXQLWLHV $UFK +\GURELRO
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icrocystis VS VFXPV )(06 0LFURE (FRO ,EHOLQJV %: 0XU /5 .LQVPDQ 5 DQG :DOVE\ $( D 0LFURF\VWLV FKDQJHV LWV EXR\DQF\ LQ UHVSRQVH WR WKH DYHUDJH LUUDGLDQFH LQ WKH VXUIDFH PL[HG OD\HU $UFK +\GURELRO ,EHOLQJV%:0XU/5 DQG:DOVE\$(E 'LXUQDOFKDQJHVLQ WKH EXR\DQF\ UHJXODWLRQ DQG YHUWLFDO GLVWULEXWLRQ LQ SRSXODWLRQV RI0LFURF\VWLVLQ WZRVKDOORZODNHV- 3ODQNWRQ 5HV ,PEHUJHU - 7KH GLXUQDO PL[HG OD\HU /LPQRO 2FHDQRJU
&KDSWHU )UHVKZDWHU%ORRPV ,PEHUJHU - DQG +DPEOLQ 3) '\QDPLFV RI ODNHV UHVHUYRLUV DQG FRROLQJ SRQGV $QQXDO 5HYLHZ RI )OXLG 0HFKDQLFV ,VWYiQRYLFV 9 3HWWHUVVRQ . 5RGULJR 0$ 3LHUVRQ ' 3DGLViN DQG &RORP : Gloeotrichia echinulata, D FRORQLDO F\DQREDFWHULXP ZLWK D XQLTXH SKRVSKRUXV XSWDNH DQG OLIH VWUDWHJ\ - 3ODQNWRQ5HV -HQVHQ -3 -HSSHVHQ ( 2OULN . DQG .ULVWHQVHQ 3 ,PSDFW RI QXWULHQWV DQG SK\VLFDO IDFWRUV RQ WKH VKLIW IURP F\DQREDFWHULDO WR FKORURSK\WH GRPLQDQFH LQ VKDOORZ 'DQLVK ODNHV &DQ - )LVK $TXDW 6FL .LQJ '/ 7KHUROHRI FDUERQLQHXWURSKLFDWLRQ - :DWHU3ROO &RQWU )HG .LQVPDQ 5 OEHOLQJV %: DQG :DOVE\ $( *DV YHVLFOH FROODSVH E\ WXUJRU SUHVVXUH DQG LWV UROH LQ EXR\DQF\ UHJXODWLRQ E\ Anabaenaflos-aquae. - *HQ 0LFURELRO .LUN -72 /LJKW DQG SKRWRV\QWKHVLV LQ DTXDWLF HFRV\VWHPV &DPEULGJH 8QLYHUVLW\ 3UHVV &DPEULGJH .LUN -72 DQG 2OLYHU 5/ 2SWLFDO FORVXUH LQ DQ XOWUDWXUELG ODNH -RXUQDO RI *HRSK\VLFDO 5HVHDUFK .OHEDKQ + *DVYDNXROHQ HLQ %HVWDQGWLHO GHU =HOOHQ GHU :DVVHUEOXWHELOGHQGHQ 3K\FRFKURPDFHHQ )ORUD -HQD .OHPHU $5 7KH YHUWLFDO GLVWULEXWLRQ RI Oscillatoria aqardhii YDU isothrix. $UFK +\GURELRO .OHPHU $5 1LWURJHQ OLPLWDWLRQ RI JURZWK DQG JDV YDFXRODWLRQ LQ Oscillatoria rubescens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scillatoria. 6FLHQFH .RKOHU - ,QIOXHQFH RI WXUEXOHQW PL[LQJ RQ JURZWK DQG SULPDF\ SURGXFWLRQ RI Microcystis aeruginosa LQ WKH K\SHUWURSKLF %XDW]HQ 5HVHUYRLU $UFK +\GURELRO .RQRSND $ (IIHFW RI OLJKWQXWULHQW LQWHUDFWLRQV RQ EXR\DQF\ UHJXODWLRQ E\ SODQNWRQLF F\DQREDFWHULD ,Q 5HGG\ &$ DQG .OXJ 0- HGV &XUUHQW 3HUVSHFWLYHV LQ 0LFURELDO (FRORJ\ 3URFHHGLQJV RI WKH 7KLUG ,QWHUQDWLRQDO 6\PSRVLXP RQ 0LFURELDO (FRORJ\ 0LFKLJDQ 6WDWH 8QLYHUVLW\ $XJ $XJ SS $PHULFDQ 6RFLHW\ RI 0LFURELRORJ\ :DVKLQJWRQ .RQRSND $ 0HWDOLPQHWLF F\DQREDFWHULD LQ KDUGZDWHU ODNHV %XR\DQF\ UHJXODWLRQ DQG SK\VLRORJLFDO VWDWH /LPQRO 2FHDQRJU ² .RQRSND $ .URPNDPS - DQG 0XU /5 5HJXODWLRQ RI JDV YHVLFOH FRQWHQW DQG EXR\DQF\ LQ OLJKW RU SKRVSKRUXVOLPLWHG FXOWXUHV RI Aphanizomenon flos-aquae &\DQRSK\WD - 3K\FRO .RQRSND $ DQG 6FKQXU 0 (IIHFW RI OLJKW LQWHQVLW\ RQ PDFURPROHFXODU V\QWKHVLV LQ F\DQREDFWHULD 0LFURE (FRO .RQRSND $( .OHPHU $5 :DOVE\ $( DQG OEHOLQJV %: (IIHFWV RI PDFURQXWULHQWV XSRQ EXR\DQF\ UHJXODWLRQ E\ PHWDOLPQHWLF Oscillatoria agardhii LQ 'HPLQJ /DNH 0LQQHVRWD - 3ODQNWRQ 5HV
.URPNDPS - )RUPDWLRQ DQG IXQFWLRQDO VLJQLILFDQFH RI VWRUDJHSURGXFWVLQF\DQREDFWHULD 1 = - 0DU)UHVKZDWHU5HV .URPNDPS - .RQRSND $ DQG 0XU /5 %XR\DQF\ UHJXODWLRQ LQ D VWUDLQ RI Aphanizomenon flos-aquae &\DQRSK\FHDH WKH LPSRUWDQFH RI FDUERK\GUDWH DFFXPXODWLRQ DQGJDVYHVLFOHFROODSVH -*HQ0LFURELRO .URPNDPS - .RQRSND $ DQG 0XU /5 %XR\DQF\ UHJXODWLRQ LQ OLJKWOLPLWHG FRQWLQXRXV FXOWXUHV RI Microcystis aeruginosa. - 3ODQNWRQ 5HV .URPNDPS - 9DQ 'HQ +HXYHO $ DQG 0XU/5 3KRVSKRUXV XSWDNH DQG SKRWRV\QWKHVLV E\ SKRVSKDWHOLPLWHG FXOWXUHV RI WKH F\DQREDFWHULXP Microcystis aeruginosa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pulex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
0HIIHUW 0( DQG .UDPEHFN +- 3ODQNWRQLF EOXHJUHHQ DOJDH RI WKH 2VFLOODWRULD redekei JURXS $UFK +\URELRO 0HQGD\ '& DQG %XFN $$ $ VKRFNZDYH WHFKQLTXH WR FROODSVHWKHYDFXROHVRIEOXHJUHHQDOJDH:DWHU5HV 0RQLVPLWK 6* ,PEHUJHU - DQG 0RULVRQ 0/ &RQYHFWLYH PRWLRQV LQ WKH VLGHDUP RI D VPDOO UHVHUYRLU /LPQRO 2FHDQRJU 0XU /5 6RPH DVSHFWV RI WKH HFRSK\VLRORJ\ RI F\DQREDFWHULD$QQDOHV0LFURELRORJ\% 2FKRD GH $OGD-$* 7DSLD 0, )DEULFH ) /ODPD0-DQG 6HUUD-/ &KDQJHV LQ QLWURJHQ VRXUFH PRGLI\ GLVWULEXWLRQ RI H[FLWDWLRQ HQHUJ\ LQ WKH F\DQREDFWHULXP Phormidium laminosum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nabaena flos-aquae F\DQREDFWHULD /LPQRO 2FHDQRJU 2UPHURG -* 3K\VLRORJ\RI WKHSKRWRV\QWKHWLF SURNDU\RWHV ,Q 0DQQ 1+ DQG&DUU1* HGV 3KRWRV\QWKHWLF3URNDU\RWHVSS 3OHQXP3UHVV1HZ
5/2OLYHUDQG***DQI 3HUDNLV 66 :HOFK (% DQG -DFRE\ -0 6HGLPHQWWRZDWHU EOXHJUHHQ DOJDO UHFUXLWPHQW LQ UHVSRQVH WR DOXP DQG HQYLURQPHQWDOIDFWRUV +\GURELRORJLD 3HWWHUVVRQ . +HUOLW] ( DQG ,VWYiQRYLFV 9 7KH UROH RI Gloeotrichia echinulata LQ WKH WUDQVIHU RI SKRVSKRUXV IURP VHGLPHQWVWRZDWHULQ /DNH(UNHQ +\GURELRORJLD 3LFN )5 DQG /HDQ '56 7KHUROH RI PDFURQXWULHQWV & 1 3 LQ FRQWUROOLQJ F\DQREDFWHULDO GRPLQDQFH LQ WHPSHUDWH ODNHV 1 = - 0DU)UHVKZDWHU5HV 3RVW $) 'H :LW 5 DQG 0XU /5 D ,QWHUDFWLRQV EHWZHHQ WHPSHUDWXUHDQG OLJKWLQWHQVLW\ RQ JURZWK DQG SKRWRV\QWKHVLV RI WKH F\DQREDFWHULXP 2VFLOODWRULD agardhii. - 3ODQNWRQ 5HV 3RVW $) (LMJHQUDDP ) DQG 0XU /5 E ,QIOXHQFH RI OLJKW SHULRG OHQJWK RQ SKRWRV\QWKHVLV DQG V\QFKURQRXV JURZWK RIWKH JUHHQDOJDScenedesmus protuberans. %U3K\FRO- 3RVW $) /RRJPDQ -* DQG 0XU /5 3KRWRV\QWKHVLV FDUERQ IORZV DQG JURZWK RI 2VFLOODWRULD agardhi *RPRQW LQ HQYLURQPHQWV ZLWK D SHULRGLF VXSSO\ RIOLJKW - *HQ 0LFURELRO 3UHVWRQ , 6WHZDUW:'3 DQG 5H\QROGV &6 %ORRPIRUPLQJ F\DQREDFWHULXP Microcystis aeruginosa RYHUZLQWHUV RQ VHGLPHQWVXUIDFH 1DWXUH 5H\QROGV &6 7KH HFRORJ\ RI WKH SODQNWRQLF EOXHJUHHQ DOJDH LQ WKH1RUWK6KURSVKLUHPHUHV )LHOG6WXGLHV ² 5H\QROGV &6 6HVWRQ VHGLPHQWDWLRQ H[SHULPHQWV ZLWK Lycopodium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¶WWHOO XV ,Q6XWFOLIIH ': DQG -RQHV -* HGV (XWURSKLFDWLRQ 5HVHDUFK DQG $SSOLFDWLRQ WR :DWHU 6XSSO\ SS )UHVKZDWHU %LRORJLFDO $VVRFLDWLRQ$PEOHVLGH8. 5H\QROGV &6 6ZLQJV DQG URXQGDERXWV (QJLQHHULQJ WKH HQYLURQPHQW RI DOJDO JURZWK ,Q :KLWH .1 DQG RWKHUV HGV 8UEDQ :DWHUVLGH 5HJHQHUDWLRQ 3UREOHPV DQG 3URVSHFWV (OOLV+RUZRRG 1HZ
&KDSWHU )UHVKZDWHU %ORRPV 5H\QROGV &6 -DZRUVNL *+0 &PLHFK +$ DQG /HHGDOH*) 2Q WKH DQQXDO F\FOH RI WKH EOXHJUHHQ DOJD Microcystis aeruginosa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scillatoria agardhii. $UFK 0LFURELRO 5LHJPDQ 5 5XWJHUV 0 DQG 0XU /5 (IIHFWV RI SKRWRSHULRGLFLW\ DQG OLJKW LUUDGLDQFH RQ SKRVSKDWHOLPLWHG Oscillatoria agardhii LQ FKHPRVWDW FXOWXUHV $UFK 0LFURELRO 5REDUWV 5' DQG =RKDU\ 7 Microcystis aeruginosa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ulacoseira DQG Anabaena GRPLQDQFH LQ D WXUELGULYHUZHLUSRRO /LPQRO2FHDQRJU 6LPRQ 5' ,QFOXVLRQ ERGLHV LQ WKH F\DQREDFWHULD F\DQRSK\FLQ SRO\SKRVSKDWH SRO\KHGUDO ERGLHV ,Q )D\ 3 DQG YDQ %DDOHQ & HGV 7KH &\DQREDFWHULD SS (OVHYLHU $PVWHUGDP 6PLWK ,5 $ VLPSOH WKHRU\ RI DOJDO GHSRVLWLRQ )UHVKZDWHU %LRO
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nabaena DQG Synechococcus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hormidium laminosum. 3ODQWD 7KRPDV 5+ DQG :DOVE\ $( %XR\DQF\ UHJXODWLRQ LQ D VWUDLQ RI Microcystis. - *HQ 0LFURELRO 7KRPDV 5+ DQG :DOVE\ $( 7KH HIIHFW RI WHPSHUDWXUH RQ UHFRYHU\ RI EXR\DQF\ E\ Microcystis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scillatoria XQDEOHWRFROODSVHJDV YDFXROHV E\WXUJRU SUHVVXUH $UFK+\GURELRO YDQ%DDOHQ& 1LWURJHQIL[DWLRQ ,Q )D\3DQGYDQ%DDOHQ &HGV 7KH&\DQREDFWHULD SS (OVHYLHU$PVWHUGDP YDQ /LHUH / DQG 0XU /5 2FFXUUHQFH RI Oscillatoria agardhii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
5/2OLYHUDQG***DQI :DOVE\ $( DQG 5H\QROGV &6 6LQNLQJ DQG IORDWLQJ 0RPV ,* HG 7KH 3K\VLRORJLFDO (FRORJ\ RI 3K\WRSODQNWRQ SS %ODFNZHOO 6FLHQWLILF 2[IRUG :DOVE\ $( 5H\QROGV &6 2OLYHU 5/ DQG .URPNDPS- 7KH UROH RI JDV YDFXROHV DQG FDUERK\GUDWH FRQWHQW LQ WKH EXR\DQF\ DQG YHUWLFDO GLVWULEXWLRQ RI Anabaena minutissima LQ /DNH 5RWRQJDLR 1HZ =HDODQG (UJHE /LPQRO :DOVE\ $( 8WNLOHQ +& DQG -RKQVHQ ,- %XR\DQF\FKDQJHV RI D UHG FRORXUHG Oscillatoria agardhii LQ /DNH *MHUVM¡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scillatoria redekei YDQ *RRU $UFK 0LFURELRO :RON &3 (UQVW $ DQG (OKDL - +HWHURF\VW PHWDEROLVP DQG GHYHORSPHQW ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 'RUGUHFKW 7KH 1HWKHUODQGV :RRG 0' DQG 2OLYHU 5/ )OXRUHVFHQFH WUDQVLHQWV LQ UHVSRQVH WR QXWULHQW HQULFKPHQW RI QLWURJHQ DQG SKRVSKRUXV OLPLWHG Microcystis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icrocystis aeruginosa K\SHUVFXPV IURP D K\SHUWURSKLF ODNH )UHVKZDWHU %LRO =RKDU\ 7 DQG 5REDUWV 5' 'LXPDO PL[HG OD\HUV DQG WKH ORQJWHUP GRPLQDQFH RI Microcystis aeruginosa. - 3ODQNWRQ 5HV
&KDSWHU 3LFRSODQNWRQ DQG 2WKHU 1RQ%ORRP)RUPLQJ &\DQREDFWHULD LQ /DNHV -RKQ6WRFNQHU&ULVWLDQD&DOOLHUL DQG*HUWUXG&URQEHUJ (FR/RJLF /WG0DWKHUV $YHQXH :HVW 9DQFRXYHU%ULWLVK&ROXPELD &DQDGD 99 - &15 OVWLWXWROWDOLDQRGLOGURELRORJLD/DUJR9 7RQROOL 3DOODQ]D,WDO\ 'HSDUWPHQW RI (FRORJ\?/LPQRORJ\ 8QLYHUVLW\ RI /XQG 6ZHGHQ
6XPPDU\ , ,QWURGXFWLRQ ,, 6DPSOLQJ3UHVHUYDWLRQDQG(QXPHUDWLRQ ,OO 7KH1RQ%ORRP)RUPHUV:KDWDUH7KH\" $ 6SHFLHV % 3F\&RPPXQLW\ ,9 $&RPPRQ(FRORJ\" $ 3F\6HDVRQDO'LVWULEXWLRQ % 3F\9HUWLFDO'LVWULEXWLRQ & &3F\'LVWULEXWLRQ ' &RORQ\)RUPDWLRQ ( *URZWK ) &RQWULEXWLRQWR7RWDO3URGXFWLRQDQG%LRPDVV * )DFWRUV$IIHFWLQJ*URZWK'LVWULEXWLRQDQG3URGXFWLRQ /LJKW 89% S+ 1XWULHQWV *UD]LQJ + 0LFURELDO)RRG:HEV 9 &RQFOXVLRQV
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hitton and M. Potts (eds), The Ecology of Cyanobacteria, © Kluwer Academic Publishers. Printed in the Netherlands.
- 6WRFNQHU&&DOOLHULDQG* &URQEHUJ
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µVLQNV¶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
, ,QWURGXFWLRQ 7KLV FKDSWHU GLVFXVVHV D JURXS RIF\DQREDFWHULD WKH ³QRQEORRP IRUPHUV´ WKDW DUH FRPPRQ LQ ODNHV WKURXJKRXW WKH ZRUOG DQG DEXQGDQW DFURVV D ZLGH VSHFWUXP RI WURSKLF FRQGLWLRQV
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iUHN &URQEHUJ .RPiUNRYi/HJQHURYi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
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV ODUJH DQG GLYHUVH JURXS RI F\DQREDFWHULD VKDUH D FRPPRQ HFRORJ\ :H FRQFOXGH WKH UHYLHZ ZLWK D SOHD IRU OLPQRORJLVWV WR SD\ PRUH DWWHQWLRQ WR WKHVH RUJDQLVPV EHFDXVH WKHH[WHQWRIWKHLU DEXQGDQFH DQG GLVWULEXWLRQ LQ ODNHV PD\ SURYLGH DQ LPSRUWDQW PHVVDJH DERXWWKH FKDQJHV OLNHO\ WR RFFXU LQ SHODJLF FRPPXQLW\VWUXFWXUHLQWKHZDUPHUZRUOGH[SHFWHGLQ WKHVWFHQWXU\0DQQ 6WRFNQHU
,, 6DPSOLQJ 3UHVHUYDWLRQ DQG (QXPHUDWLRQ ,QIRUPHG FRPPHQW RQ WKH HFRORJ\ RI QRQEORRP IRUPHUV UHOLHV RQ WKH LQYHVWLJDWRU¶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nopore® UHVXOWV LQ WKH EHVW UHWHQWLRQ RI VPDOO SDUWLFOHV 6WRFNQHU HW DO %HUWRQL DQG WKH PRVW KRPRJHQHRXV GLVWULEXWLRQ RI FHOOV RQ WKH ILOWHU VXUIDFH 5 %HUWRQL XQSXEOLVKHG Anopores® GR QRW QHHG WR EH EODFNHQHG RU SUHVWDLQHG IRU
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¶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
,,, 7KH1RQ%ORRP)RUPHUV:KDWDUH 7KH\" $ 6SHFLHV 6RPHRIWKHPRVWLPSRUWDQWVSHFLHV RI3F\ DQG &3F\ DUH OLVWHG LQ 7DEOH WRJHWKHU ZLWK VDOLHQW PRUSKRORJLFDO DQG HFRORJLFDO LQIRUPDWLRQ 7KH VLQJOHFHOOHG 3F\ ZHUH PLVVHG E\ PRVW HDUO\ LQYHVWLJDWRUV EHFDXVH RIWKHLU PLQXWH VL]H RU LIWKH\ ZHUH VHHQ WKH\ ZHUH HLWKHU SRRUO\ GHVFULEHG RU VLPSO\LJQRUHG6WRFNQHUDQG$QWLD 6WRFNQHU 7KH\KDYHEHHQFDOOHG µDOJDH¶ /XQG 2YHUEHFN GHVFULEHG DV µOLWWOH URXQG JUHHQ WKLQJV¶ /5*7 3HDUO RUDV VPDOO FRFFRLG RU &KORUHOODOLNH FHOOV 5HFHQWO\ .RPiUHN KDV SURSRVHG D QHZ FODVVLILFDWLRQ IRU WKH VROLWDU\ OLYLQJ SLFRF\DQRSURNDU\RWHV 3F\ EDVHG RQ F\WR PRUSKRORJLFDO PROHFXODU DQG ELRFKHPLFDO DSSURDFKHV +H VHSDUDWHG WKUHHWD[RQRPLF JURXSV DW WKH JHQHULF OHYHO Cyanobium, Synechococcus DQG Cyanothece dianae/cedrorum W\SHV +H LQGLFDWHV WKDW WKH ILUVW FOXVWHU LV WKH RQH FRPSULVLQJ WKH PDMRULW\ RISLFRSODQNWRQ RU 3F\ RUJDQLVPV WKDWPRVW DXWKRUV IRUODFNRIDEHWWHUQDPHFDOO Synechococcus. :KLOH PRVW SHODJLF 3F\ DUH \HW EH GHVFULEHG 6WRFNQHU DQG $QWLD VRPH Synechococcus VSHFLHV KDYH EHHQ ZHOO VWXGLHG %DLOH\:DWWV DQG .RPi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yanonephron, Snowella ,Q WKRVH ODNHV ZKHUH &3F\ DUH FRPPRQ WKHUH LV XVXDOO\ D PL[WXUH RI VSHFLHV HDFK ZLWK GLVWLQFWLYH FRORQ\VWUXFWXUHVZKLFKDUHTXLWHUHDGLO\LGHQWLILDEOH &3F\ DUH IRXQG WKURXJKRXW WKH HQWLUH VSHFWUXP RI ODNH WURSKLF FRQGLWLRQV KRZHYHU PRVWWHQG WR RFFXU LQPRUHSURGXFWLYHPHVRHXWURSKLFODNHV6RPHRIWKH PRVW FRPPRQ &3F\ LQ IUHVK ZDWHUV DUH VSHFLHV EHORQJLQJ WR WKH JHQHUDAphanocapsa, Aphanothece, Chroococcus, Coelosphaerium, Cyanodictyon, Merismopedia, Snowella DQG Tetrarcus 7DEOH
- 6WRFNQHU&&DOOLHULDQG* &URQEHUJ % 3F\&RPPXQLW\ ,W LV LPSRUWDQW WR LGHQWLI\ WKH FRPSRVLWLRQ RI DOO FRPSRQHQWV RI $33 FRPPXQLWLHV WR JHW D EHWWHU XQGHUVWDQGLQJ RI VSHFLHV LQWHUDFWLRQV VSDWLDO DQG WHPSRUDO VXFFHVVLRQ DQG VXUYLYDO VWUDWHJLHV RIPL[HG $33SRSXODWLRQV 6WRFNQHU %HFDXVH3F\ DUH QXPHULFDOO\ WKH SUHGRPLQDQW VSHFLHV LQ $33 FRPPXQLWLHV DQG KDYHUHFHLYHG WKH PRVW DWWHQWLRQ LQ FRPPXQLW\ VWXGLHV WKHUH KDV EHHQ D WHQGHQF\ ¶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¡QGHUJDDUG 6WRFNQHU DQG 6KRUWUHHG )LQGOH\DQG.DVLDQ +RZHYHU LQ ODUJH GHHS ROLJRWURSKLF DQG PHVRWURSKLF ODNHV 3F\ SRSXODWLRQV DUH DOZD\V GRPLQDQW DQG UHSUHVHQWWKH PDMRU VSHFLHV ZLWKLQWKH$33 FRPPXQLW\ 7DEOH :HLVVH &DOOLHULDQG3LQROLQL 3DGLViNHWDO
,9 $&RPPRQ (FRORJ\"
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µVWUDLQV¶ RI 3F\ HDFK UHVSRQVLEOH IRU RQH RI WKH SHDNV 7KLV QHZ SHUVSHFWLYH YDOLGDWHG E\ UHFHQW ILQGLQJV RQ WKH JHQHWLF GLYHUVLW\ RI PRUSKRORJLFDOO\ VLPLODU
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV Table 1. &RPPRQ SLFRF\DQREDFWHULD3F\ IURPGLIIHUHQWKDELWDWV
Table 1 FRQWLQXHG &RPPRQSLFRF\DQREDFWHULD3F\ IURPGLIIHUHQWKDELWDWV
-6WRFNQHU&&DOOLHULDQG* &URQEHUJ
&KDSWHU 3LFRSODQNWRQDQGRWKHUQRQEORRP IRUPHUV Table 1 FRQWLQXHG &RPPRQSLFRF\DQREDFWHULD3F\ IURPGLIIHUHQWKDELWDWV
Table FRQWLQXHG &RPPRQ EDFWHULD 3F\ IURP GLIIHUHQW KDELWDWV
- 6WRFNQHU & &DOOLHUL DQG * &URQEHUJ
&KDSWHU 3LFRSODQNWRQDQG RWKHUQRQEORRPIRUPHUV
Table 1 FRQWLQXHG &RPPRQSLFRF\DQREDFWHULD3F\ IURPGLIIHUHQWKDELWDWV
Table 2. $YHUDJH RU PD[LPXP ZKHQ VSHFLILHG QXPEHU RISLFRSODQNWRQLF FHOOV GLYLHG LQWR SURNDU\RWLF 3F\ HXNDU\RWLF DQG FRORQLDO &3F\ LQ ODNHV RIGLIIHUHQW WURSKLF FRQGLWLRQV 7KH SHUFHQWDJH FRQWULEXWLRQ RI$33 WR WRWDO DOJDO ELRPDVV LV UHSRUWHG DV D PHDQ RU DV D UDQJH
Synechococcus LQ /DNH &RQVWDQFH (UQVW HW DO 3RVWLXV HW DO VXJJHVWV D SRVVLEOH DGDSWLYH UHVSRQVH E\ 3F\ VWUDLQV WR WKH VHDVRQDOO\ YDULDEOH SK\VLFRFKHPLFDO IDFWRUV LQ WKH HXSKRWLF ]RQH ,Q D \HDU VWXG\ RI $33 FRPPXQLWLHV LQ /DNH 0DJJLRUHWKH DEXQGDQFH RI3F\ JUDGXDOO\ LQFUHDVHG DV WKH ODNH¶V QXWULHQW ORDGV GHFOLQHG DQG WKH ELPRGDO DEXQGDQFHSDWWHUQZLWK DODUJH VSULQJSHDN LVDSURPLQHQWIHDWXUHRIWKH VHDVRQDOF\FOH )LJ &DOOLHULDQG6WRFNQHU LQSUHVV /DUJHVSULQJSHDNV DUH DOVR FRPPRQ LQ HXWURSKLF DQG K\SHUHXWURSKLF ODNHV 6LPH1JDQGR 9RURVHWDO EXW QRW LQ ZDUPPRQRPLFWLF ODNHV 6WRFNQHU DQG 6KRUWUHHG 0DOLQVN\5XVKDQVN\HWDO 7KH VHDVRQDO SDWWHUQV IRXQG LQ VHYHQ 'DQLVK ODNHV 6¡QGHUJDDUG VHYHUDO &DQDGLDQ ODNHV 3LFN
- 6WRFNQHU&&DOOLHULDQG*&URQEHUJ DQG$JEHWL /DNH%LZD -DSDQ0DHGDHWDO DQG LQ WKH QRUWKHUQ (QJOLVK ODNHV +DZOH\ DQG :KLWWRQ E DOO ODFNWKH VSULQJSHDN WKHUH EHLQJ RQO\ D VXPPHU RU DXWXPQ PD[LPXP 6XFK ILQGLQJV REWDLQHGIURPDEXQGDQFHHVWLPDWHVDWRQO\ RQH VHOHFWHG GHSWK DUH QRW UHDOO\ UHSUHVHQWDWLYH RI WKH HXSKRWLF ]RQH DQG SURYLGH DW EHVW RQO\ D µVQDSVKRW¶ GHVFULSWLRQ RI VHDVRQDO YDULDWLRQ LQ VXUIDFH ZDWHUV $V ZHOO GHPRQVWUDWHG E\ D \HDU WLPH VHULHV IURP /DNH 0DJJLRUH RQO\ E\ XVLQJ LQWHJUDWHGVDPSOHVLQORQJHUWHUPVWXGLHVFDQDFOHDU SLFWXUH RI 3F\ VHDVRQDO DQG GHSWK GLVWULEXWLRQ SDWWHUQV HPHUJH )LJ 7KLV NLQG RI VDPSOLQJ VKRXOG EH DFFRPSDQLHG E\ D VWXG\ RI WKH YHUWLFDO GLVWULEXWLRQ ZLWK IUHTXHQW VDPSOHV GRZQ WKH ZDWHU FROXPQ
Fig. 1. /DNH0DJJLRUH$33DEXQGDQFHIURP WR (XNDU\RWHVDQGSURNDU\RWHV3F\ ZHUHFRXQWHGVHSDUDWHO\&DOOLHULDQG 6WRFNHULQSUHVV
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a PG 5XL]HWDO EXW EHFDXVH RI WKH ORZ VLQNLQJ YHORFLW\ RI 3F\ WKHLU GLVWULEXWLRQ ZLWKLQ WKH ZDWHU FROXPQ LV QRW OLNHO\ WR EH DIIHFWHG E\ WKH ODNH V WKHUPDO VWUXFWXUH 'XULQJ WKHUPDO VWUDWLILFDWLRQ SHDN 3F\ DEXQGDQFHKDVEHHQQRWHGLQWKHORZHUPHWDOLPQLRQ RUXSSHUK\SROLPQLRQLQ/DNH+XURQ DQG 0LFKLJDQ
)DKQHQVWLHODQG&DUULFN LQWKHPHWDOLPQLRQ EHORZ WKH WKHUPRFOLQH LQ /DNHV &RQVWDQFH 0DJJLRUHDQG6WHFKOLQ:HLVVH 6FKZHL]HU &DOOLHULDQG3LQROLQL 3DGLVi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ynechococcus FXOWXUHV WR VSHFLILF OLJKW LUUDGLDQFHV :DWHUEXU\ HW DO +RZHYHU 3F\ KDYH EHHQ FODVVLILHG DV HXU\SKRWLF DQG DUH
Fig. 2. ,VRSOHWKHVRI3F\DEXQGDQFH LQ/DNH0DJJLRUHGXULQJ 3DVVRQLHWDO
- 6WRFNQHU & &DOOLHUL DQG * &URQEHUJ
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¶V FRQFOXVLRQV WKDW3F\DUHH[WUHPHO\DGDSWDEOHRUJDQLVPV DQGFDQ JURZ LQ D YDULHW\ RI OLJKW FRQGLWLRQV /LJKW OLNH WHPSHUDWXUHLV DQLPSRUWDQWGHWHUPLQLVWLFIDFWRUIRU 3F\ VXUYLYDO EXW LWV LQIOXHQFH RQ YHUWLFDO GLVWULEXWLRQ LV GLIILFXOWWR LVRODWH IURPWKRVHUHODWHG WRELRORJLFDOLQWHUDFWLRQVLHJUD]LQJ ,Q /DNH 0DJJLRUH WKH FLOLDWH DQG IODJHOODWH YHUWLFDO GLVWULEXWLRQV GXULQJ VXPPHU VWUDWLILFDWLRQ DUH LQYHUVHO\ UHODWHG ZLWK 3F\ DEXQGDQFH )LJ ZKLFK VXJJHVWV WKDW µWRS GRZQ¶ IDFWRUV JUD]LQJ H[HUW PRUH FRQWURO RI3F\ SRSXODWLRQV DQG YHUWLFDO GLVWULEXWLRQ SDWWHUQV WKDQ SK\VLFDO IDFWRUV LQ ODUJH ROLJR PHVRWURSKLF ODNHV :HLVVH DQG 6FKZHL]HU 6WRFNQHUDQG6KRUWUHHG DQGWKHVDPH VHHPVDOVRWREHWUXHLQVPDOOPHVRWURSKLFODNHVDQG UHVHUYRLUV 6LPHN HW DO +RZHYHU WKHUH LV FRPSHOOLQJ HYLGHQFH IURP ZKROHODNH IHUWLOLVDWLRQ H[SHULPHQWV WKDW LQ XOWUDROLJRWURSKLF %ULWLVK &ROXPELD ODNHV QXWULHQW OLPLWDWLRQ RU µERWWRP XS¶ IDFWRUVPD\EHPRUHLPSRUWDQWWKDQOLJKWRUJUD]LQJ LQ HVWDEOLVKLQJ 3F\ YHUWLFDO GLVWULEXWLRQ SDWWHUQV ZLWKLQ WKH HXSKRWLF ]RQH 6XWWOH HW DO 6WRFNQHUDQG6KRUWUHHG
& &3F\'LVWULEXWLRQ 0DQ\ RI WKH &3F\ VKRZ D FRVPRSROLWDQ GLVWULEXWLRQ ZKLOH RWKHUV DUH DSSDUHQWO\ PRUH UHVWULFWLYHDQGIRXQG RQO\LQWKHWHPSHUDWHODWLWXGHV RI WKH QRUWKHUQ KHPLVSKHUH HJ Aphanothece minutissima, $ bachmannii, Lemmermanniella pallida. 0RVW FRORQLDO &3F\ KDYH EHHQ UHFRUGHG IURP PHVR WR HXWURSKLF ODNHV RU SRQGV ZLWK YHU\ IHZ VSHFLHV IRXQG LQ ROLJRWURSKLF ODNHV 7DEOH 2QO\ WKH WD[RQ Tetrarcus ilsteri KDV EHHQ IRXQG H[FOXVLYHO\LQROLJRWURSKLF DQGSRO\KXPLFODNHV LQ )LQODQG6ZHGHQDQGWKH%DOWLF6WDWHV6NXMD * &URQEHUJ XQSXE EXW LQ VXFK ORFDOLWLHV ZKHQ SUHVHQW LW FDQ EH YHU\ DEXQGDQW 0DQ\ &3F\ FR
)LJ 9HUWLFDO GLVWULEXWLRQ RI 3F\ FHOO QXPEHUV DQG RI LPSRUWDQW JUD]HUV GXULQJ VXPPHU VWUDWLILFDWLRQ LQ ROLJRWURSKLF /DNH 0DJJLRUH ,Q -XQH FLOLDWHV JUD]H RQ SLFR DQG +1) DQG LQ $XJXVW +1) DUH WKH PRVW LPSRUWDQW 3F\ JUD]HU
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV
)LJ 3F\DEXQGDQFHLQDVLQJOHDQGDJJUHJDWHGFRORQ\IRUPLQ/DNH0DJJLRUH
RFFXU ZLWK ODUJHU EOXHJUHHQV HJ Anabaena, Aphanizomenon, Microcystis, Planktothrix DQG Woronichinia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phanocapsa nubilum, A. elachista DQG Coelosphaerium punctiferum. Aphanocapsa, Aphanothece DQG Chroococcus DUH FRPPRQ LQ HXWURSKLF ODNHV LQ 6FDQGLQDYLD LQ )LQODQG WKH %DOWLF VWDWHV DQG QRUWKHUQ *HUPDQ\ .RPiUNRYi/HJQHURYi DQG &URQEHUJ ,Q IDFW PRVW RIWKH FRPPRQ &3F\ VSHFLHV KDYH EHHQ GHVFULEHG IURP WKLV SDUW RI WKH ZRUOG 7DEOH 2QO\ DIHZ RIWKHVH WD[DKDYHEHHQ FXOWXUHG LQWKH ODERUDWRU\ XQGHU FRQWUROOHG FRQGLWLRQV WKXV WKH YDULDELOLW\ZLWKLQRUDPRQJWKHVHVSHFLHVLVQRWZHOO NQRZQ 7KH 6ZHGLVK 6FDQLDQ ODNHV ZKHUH PHPEHUV RI PRVW RI WKH ZHOONQRZQ JHQHUD KDYH EHHQIRXQG DUH HXWURSKLF WRK\SHUWURSKLF ZLWK73
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
' 3F\&RORQ\)RUPDWLRQ 6PDOO 3F\ FRORQLHV DUH FRPPRQ LQ PRVW ODNHV DQG XVXDOO\ UHDFK KLJKHVW DEXQGDQFH GXULQJ SHULRGV RI VHYHUH QXWULHQW OLPLWDWLRQ LQ VXPPHU DQG DXWXPQ )LJ 3ODWH 6WRFNQHU .OXW DQG 6WRFNQHU +RZHYHU WKHUH LV D SDXFLW\ RI OLWHUDWXUH RQ WKH VXEMHFW DQG ZLWKRXW D EHWWHU FRPSUHKHQVLRQ RI WKH SURFHVV VWXGLHV RI 3F\ VHDVRQDO VXFFHVVLRQ ZLOO UHPDLQ HTXLYRFDO 6WRFNQHU:HLVVH 7KH SUHVHQFH RI ORRVH RU WLJKWO\ FOXVWHUHG FRORQLHV DSSHDUV WR KDYH DGDSWLYH VLJQLILFDQFH
SHUKDSV DV D VWUDWHJ\ IRU PRUH HIILFLHQW QXWULHQW UHF\FOLQJ GXULQJ SHULRGV RI VFDUFLW\ .OXW DQG 6WRFNQHU 3HGUyV$OLy DQG %URFN RU SRVVLEO\ DV DQ DQWLSUHGDWRU PHFKDQLVP 6WRFNQHU /LWWOH LV NQRZQ DV WR ZKHWKHU FRORQLHV DUH PRUH UHDGLO\ JUD]HG WKDQ VLQJOH FHOOV EXW LW LV WKRXJKWWKDWWKH ODUJH H[WHUQDO SRO\VDFFKDULGH OD\HU LV DYRLGHG E\ PLFUR DQG PDFUR]RRSODQNWRQ .OXW DQG6WRFNQHU 6WXGLHV RQ PDULQH Synechococcus VWUDLQV KDYH VKRZQDQRYHOIRUPRIµVZLPPLQJ¶ PRWLOLW\ZLWKRXW IODJHOOD ZKRVH HFRORJLFDO VLJQLILFDQFH PD\ EH UHODWHG WR DJJUHJDWH IRUPDWLRQ DQG WKHLU DELOLW\ WR H[KLELW FKHPRWD[LV DQG PRYH WR KLJKHU QXWULHQW ]RQHV HJ PDULQH VQRZ QLWULFOLQH ERXQGDULHV :DWHUEXU\ HW DO :LOOH\ DQG :DWHUEXU\ ,WLVTXLWHOLNHO\WKDWVLPLODUµPRWLOLW\¶ RFFXUV LQ IUHVKZDWHU Synechococcus VWUDLQV ZKLFK FRXOG H[SODLQ ZK\ FRORQLHV DUH PRUH SUHYDOHQW GXULQJ SHULRGV RI QXWULHQW GHSOHWLRQ LQ XOWUD ROLJRWURSKLF V\VWHPV .OXW DQG 6WRFNQHU 7KH DELOLW\ RI 3F\ WR PRYH DQG FRQJUHJDWH DURXQG DJJUHJDWHV ZRXOG EH D SODXVLEOH PHFKDQLVP RI FRORQ\ IRUPDWLRQ 7KH SURGXFWLRQ RI VWLFN\ H[RSRO\PHULF VXEVWDQFHVRUPXFXVZKHQWKHFHOOQXPEHULQFUHDVHV FRXOG IDFLOLWDWH WKH FOXPS IRUPDWLRQ .L¡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
( *URZWK &HOOV RI Synechococcus KDYH EHHQ ZLGHO\ XVHG IRU ODERUDWRU\ H[SHULPHQWV RQ JURZWK UDWH DQG FHOO F\FOHV DQG UHVXOWV IURP WKHVH VWXGLHV KDYH EHHQ DSSOLHG WR QDWXUDO SRSXODWLRQV :DWHUEXU\ HW DO
-6WRFNQHU&&DOOLHULDQG*&URQEHUJ &KLVKROP HW DO &DPSEHOO DQG &DUSHQWHU D 6ZHHQH\ DQG %RUJHVH $UPEUXVW HW DO )DKQHQVWLHO HW DO E %LQGHU DQG &KLVKROP &DOOLHUL HW DO 7KH SURFHVVHV RI FHOO JURZWK DQG GLYLVLRQ DUH DV WLJKWO\ FRXSOHG DV SKRWRV\QWKHVLV DQG JURZWK UDWH DQG DUH OLJKW GHSHQGHQW .DQD DQG *OLEHUW D E &KLVKROPHW DO 7KHUH LV OLWWOH GLIIHUHQFH EHWZHHQ PDULQH DQG IUHVK ZDWHU VWUDLQV RI Synechococcus LQ ERWK FHOO GLYLVLRQ DQG JURZWK ZLWK FHOO GLYLVLRQ UHDFKLQJ D PD[LPXP LQ WKH DIWHUQRRQWULJJHULQJDQLQFUHDVHLQFHOOQXPEHUWKDW SURFHHGVLQWKHGDUNF\FOH)LJ &KLVKROPHWDO 7KHVH OLJKWGDUN F\FOHV SURGXFH UK\WKPLF FHOO GLYLVLRQVDQGUHODWHGJURZWKUDWHDQGSKRWRV\QWKHWLF DFWLYLW\ WKDW DUH DOO GULYHQ E\ SUHYDLOLQJ OLJKW FRQGLWLRQV ([SHULPHQWDO ODERUDWRU\ HYLGHQFH RI WKH LQIOXHQFH RI OLJKW RQ JURZWK UDWH LV ZHOO GRFXPHQWHG E\ )DKQHQVWLHO HW DO E IRU IUHVKZDWHU Synechococcus VWUDLQV DQG E\ &DPSEHOO DQG &DUSHQWHU D IRU PDULQH VWUDLQV 7KHVH LQYHVWLJDWRUV KDYH PHDVXUHG JURZWK UDWHV DW OLJKW LQWHQVLWLHV XS WR DQG PRO SKRWRQ P V UHVSHFWLYHO\ VLPXODWLQJ QDWXUDO LUUDGLDQFH OHYHOV .DQD DQG *OLEHUW E KDYH H[WHQGHG WKH OLJKW LQWHQVLW\ OLPLW XS WR PRO P V GHPRQVWUDWLQJ WKDW JURZWK UDWH EHFRPHV OLJKW VDWXUDWHG DW PRO P V EXW Synechococcus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iNHWDO ZKLOHLQ WKH /DNH %DODWRQ LW ZDV G 0DVWDOD HW DO 3F\ JURZWK LQ /DNH 0DJJLRUH OLHV EHWZHHQ G DV DQHW JURZWK UDWH DQG G DV D SRWHQWLDO JURZWK XVLQJ WKH )'& PHWKRG &DOOLHUL HW DO /RVV UDWHV HVWLPDWHG DV D GLIIHUHQFH EHWZHHQ SRWHQWLDO DQG QHW JURZWK UDWH VKRZ WKH GLVDSSHDUDQFH RI! RIWKH QHZ $33 SURGXFWLRQ 7KLV UHVXOW LV REWDLQHG E\ WDNLQJ LQWR
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV DFFRXQW SRWHQWLDO JURZWK UDWH DQG WKHUHIRUH WKH FDOFXODWHG ORVVHV DUH WKRVH GXH WR ELRORJLFDO SURFHVVHV VXFK JUD]LQJ O\VLV SDUDVLWLVP DJJUHJDWLRQDQG VHGLPHQWDWLRQ 6LPLODUUHVXOWV IURP in VLWX H[SHULPHQWV KDYH DOVR GHPRQVWUDWHG D JRRG EDODQFH EHWZHHQ JURZWK DQG JUD]LQJ UDWHV IRU 3F\ SRSXODWLRQV :HLVVH $ KLJK DEXQGDQFH RI YLUXVOLNH SDUWLFOHV ZHUH QRWHG LQ DQ XOWUD ROLJRWURSKLF %ULWLVK &ROXPELD ODNH DQG ZHUH DOVR IRXQGZLWKLQ3F\FHOOVLQYDULRXVVWDJHVRILQIHFWLRQ DQGO\VLV .OXWDQG6WRFNQHU 6XWWOH KDV LPSOLFDWHG F\DQRSKDJHV DV SRWHQWLDOO\ UHVSRQVLEOH IRU RI Synechococcus O\VLV LQ PDULQH V\VWHPV DQG WKLV SURSRUWLRQ FDQ LQFUHDVH PDUNHGO\ GHSHQGLQJ RQ KRVW DEXQGDQFH WHPSHUDWXUH DQG SKRVSKDWH VWDWXV RI WKH FHOO :LOVRQHWDO
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ynechococcus +XDQJHWDO .RQGRHWDO
/DERUDWRU\ ([SHULPHQW 0D\
Fig. 5. 'LHO SDWWHUQ RIWKH FHOO QXPEHUV DQG RI WKH SHUFHQWDJH RI GLYLGLQJ FHOOV )'& RI D Synechococcus FXOWXUH LVRODWHG IURP /DNH 0DJJLRUH&DOOLHULHWDO UHSURGXFHGZLWKSHUPLVVLRQ
7KH VWURQJ FRUUHODWLRQ EHWZHHQ WKH IUHTXHQF\ RI GLYLGLQJ FHOOV )'& DQG ERWK QHW JURZWK UDWH LQ ODERUDWRU\ FXOWXUHV )DKQHQVWLHO HW DO E DQG SRWHQWLDO JURZWK UDWH )'& G LQ /DNH 0DJJLRUH )LJ &DOOLHUL HW DO GHPRQVWUDWHV WKH JRRG SUHGLFWLYH YDOXH RI WKH )'& PHWKRG LQ HVWLPDWLQJ in situ JURZWKUDWHV 6RPHFRQVLGHUWKDW WKHSUHVHQFHRIWZR JDSVGXULQJWKHFHOOF\FOHWREH OLJKWGHSHQGHQW EORFNLQJ WKH SRSXODWLRQ LQ D GRXEOHW VWDJH GXULQJ WKH QLJKW DW D FRQVWDQW )'& $UPEUXVW HW DO EXW %LQGHU DQG &KLVKROP KDYH VXJJHVWHGWKH SRVVLEOH XVH RIWKH )'& WHFKQLTXH IRU ILHOG VWXGLHV GHVSLWH VRPH XQFHUWDLQWLHV UHJDUGLQJ WKH FHOO F\FOH 'DWD RQ Synechococcus JURZWK UDWH IURP )'& ZKHQ UHJUHVVHG DJDLQVW PD[LPXP )'& IURP ODNHV 0DJJLRUH +XURQDQG 0LFKLJDQ VKRZV D VLJQLILFDQW FRUUHODWLRQ )LJ ZKLFK DOORZV RQH WR REWDLQ DQ
- 6WRFNQHU & &DOOLHUL DQG * &URQEHUJ HVWLPDWH RI 3F\ JURZWK UDWH IURP FDOFXODWLRQV RI )'&IURPQDWXUDOSRSXODWLRQV %DVHG RQ SUHVHQWNQRZOHGJH RI3F\ JURZWK UDWHV LQODNHVXVLQJDYDULHW\RIDYDLODEOHPHWKRGVZHFDQ FRQFOXGH • GRXEOLQJ WLPHV RI Synechococcus QDWXUDO SRSXODWLRQVUDQJHIURPKWRG • OLJKW LQIOXHQFHV JURZWK ZKLFK LQ WXUQ LV WLJKWO\ FRXSOHGZLWKWKHGLHOF\FOH • WKH GLHO F\FOH FRXOG EH WULJJHUHG E\ DQ HQGRJHQRXVFORFN • JURZWK UDWHV DUH GLVFRQWLQXRXV DQG ODUJHO\ FRQWUROOHGE\SUHGDWLRQ FHOOO\VLV DQGRWKHU ORVV SURFHVVHV
Fig. 6. 0D[LPXPSHUFHQWRIGLYLGLQJFHOOVRIQDWXUDOSRSXODWLRQRI6\QHFKRFRFFXVIURP/DNHV0DJJLRUH0LFKLJDQDQG+XURQDQGWKHLUin situ JURZWKUDWHV )'& G &&DOOLHULXQSXEOLVKHGGDWD
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in situ PHDVXUHPHQWV IURP GLIIHUHQWODNHV DUH RIWHQ LQGLVDJUHHPHQW 7KHPRGHO RXWOLQHG E\ 6WRFNQHU RI DQ LQFUHDVH LQ WKH FRQWULEXWLRQRI$33SURGXFWLRQDQGELRPDVV ZLWKWKH GHFUHDVH RI SKRVSKRUXV FRQFHQWUDWLRQ LQ ODNHV KDV EHHQ ZLGHO\ DFFHSWHG EXW LQ WKH OLJKW RIUHFHQW GDWD QHHGV IXUWKHU VSHFLILFDWLRQV DQG UHILQHPHQW 7KH RULJLQ RIWKH PRGHO DSSOLHG WR ODNH WURSKLF JUDGLHQW OLHV LQ WKH REVHUYDWLRQV WKDW PDULQH Synechococcus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|U|VHWDO 6RPH RIWKHVHUHVXOWVGHPRQVWUDWHWKDWWKH 6WRFNQHU PRGHO FDQQRW H[SODLQ WKH ZLGH YDULDELOLW\ RI $33 RU 3F\ FRQWULEXWLRQV WR WRWDO SURGXFWLRQ LQ DOO ODNHV DQG OHDYHV TXHVWLRQV XQDQVZHUHG WKDW UHTXLUH IXUWKHUUHVHDUFK 7ZRLQWHUSUHWDWLRQV RIWKHPRGHO DUH SRVVLEOHHLWKHU WKDW $33 FRPPXQLWLHV FDQ DGDSW VR TXLFNO\ WR FKDQJLQJ SK\VLFRFKHPLFDO UHJLPHV HJ OLJKW WHPSHUDWXUH QXWULHQW FRQFHQWUDWLRQV WXUEXOHQFH WKDW LW LV LPSRVVLEOH WR SUHGLFWWKH UHODWLRQVKLS RQ D ORQJHUWHUPPRUHJHQHUDOLVHGEDVLVRU WKH PHWKRGRORJLHV DQG GDWD HODERUDWLRQ XVHG E\ YDULRXVUHVHDUFKHUV DUH VR GLIIHUHQW WKDW GDWD FDQQRW EHHDVLO\FRPSDUHG )XUWKHUPRUH IHZ ODUJH GDWD VHWV H[LVW WKDW FOHDUO\ GRFXPHQWWKHVHDVRQDOYDULDWLRQV RIIUHVKZDWHU3F\ RU $33 SULPDU\ SURGXFWLRQ )DKQHQVWLHO DQG &DUULFN :HLVVHDQG6FKZHL]HU DQGWKHLUUHODWLYH FRQWULEXWLRQV WR WRWDOSK\WRSODQNWRQ SURGXFWLRQ LV QRW RIWHQ H[SUHVVHG DV WKH \HDUO\ PHDQ FDOFXODWHG IURP WKH HXSKRWLF ]RQH $V SUHYLRXVO\ GLVFXVVHG LIYDOXHV DUH QRW H[SUHVVHG RQ DQ DQQXDO EDVLV WKH SHUFHQWDJH FRQWULEXWLRQV FDQ YDU\ ZLGHO\ GHSHQGLQJ RQ WKH G\QDPLFVRIVKRUWWHUPYDULDWLRQV $ JRRG H[DPSOH RI KRZ WKH FKRLFH RI GDWD H[SUHVVLRQ RU SUHVHQWDWLRQ FDQ LQIOXHQFH WKH LQWHUSUHWDWLRQ LV SURYLGHG E\ /DNH 0DJJLRUH $33 SURGXFWLRQPHDVXUHGLQWZRVXFFHVVLYH\HDUV )LJ ,I H[SUHVVHG DV D \HDUO\ PHDQ FDOFXODWHG XVLQJ WKH 3ODWW PRGHO WKH YDOXHV DUH YHU\ VLPLODU LQ J&P \ RIWRWDO 33 DQG LQ J&P \ RIWRWDO 33 6LPLODU UHVXOWV DUH REWDLQHG LIZH FRPSDUH WKH VHDVRQDO YDULDWLRQ RIWKH SKRWRV\QWKHWLF UDWH FDOFXODWHG RQ D GDLO\ EDVLV DV PJ &P G RUH[SUHVVHG RQ DQKRXUO\ EDVLV DV PJ&P K )LJ +RZHYHU E\ H[DPLQLQJ WKH YHUWLFDO SURILOH RIFDUERQ WKH $33 SHUFHQWDJH FRQWULEXWLRQ WR WRWDO SURGXFWLRQ UDQJHV IURP WR LQ DQG
- 6WRFNQHU&&DOOLHULDQG*&URQEHUJ
Fig. 7. +RXUO\UDWHRIFDUERQIL[DWLRQE\$33 FRPPXQLW\LQ/DNH0DJJLRUHGXULQJ
IURP WR LQ 7KLV H[DPSOH LOOXVWUDWHV WZR SRLQWV VKRUWWHUP HYHQWV FDQ JR XQUHFRJQLVHG DQG FRQWULEXWHIRUPDQ\\HDUVEHIRUHUHVXOWLQJLQHYLGHQW WUDQVIRUPDWLRQV RIWKHV\VWHP WKHUHIRUH LQIRUPDWLRQ REWDLQHG IURP VKRUWWHUP GDWD VHWV FDQQRW H[SODLQ WKHHYROXWLRQRIWKHV\VWHP JHQHUDO PRGHOV RI WKH FRQWULEXWLRQ WR FDUERQ SURGXFWLRQ DFURVV WURSKLF JUDGLHQWV VKRXOG EH REWDLQHG XVLQJ GDWD IURP PRUH WKDQ D VLQJOH \HDU GDWDVHW ,W DSSHDUV WKDW 3F\ SRSXODWLRQV DUH LQFUHDVLQJ WKHLU LPSRUWDQFH LQ WKLV ODNH DV DQQXDO QXWULHQW ORDGV GHFOLQH %HUWRQL DQG &DOOLHUL &DOOLHUL DQG 6WRFNQHULQSUHVV :LWK GLIIHUHQWLDO ILOWUDWLRQ DQG WKH WUDGLWLRQDO & XSWDNH PHWKRGV LW LV QRW SRVVLEOH WR HVWLPDWH WKH DFWXDO 3F\ SURGXFWLRQ EHFDXVH RIWKH LQWHUIHUHQFH RI WKH SLFRHXNDU\RWHV )RU WKLV UHDVRQ IUHTXHQW PLFURVFRSLF H[DPLQDWLRQ IRU WKH SUHVHQFH RI SLFR HXNDU\RWHVZRXOG EHKHOSIXOWRFULWLFDOO\HYDOXDWHWKH UHVXOWV ,Q PRVW ODNHV WKH PDMRULW\ RI $33 DUH Synechococcus 3F\ FHOOV 6WRFNQHU EXW LQ VRPH FDVHV UHG IOXRUHVFLQJ SLFRHXNDU\RWHV FDQ UHSUHVHQW D ODUJH IUDFWLRQ RI SLFRSODQNWRQ )DKQHQVWLHO HW DO D ,Q DGGLWLRQ TXHVWLRQV
FRQFHUQLQJ WKH DFFXUDF\ RI SRVWLQFXEDWLRQ GLIIHUHQWLDO ILOWUDWLRQ WR PHDVXUH $33 SULPDU\ SURGXFWLRQ )DKQHQVWLHO HWDO JLYHULVH WRWKH QHHG IRU D FULWLFDO UHHYDOXDWLRQ RI WKH HVWLPDWHV REWDLQHG 7KH )'& PHWKRG KDV EHHQ SURSRVHG DV DQ DOWHUQDWLYHWRUDGLRFKHPLFDO WHFKQLTXHV EHFDXVH RILWV VHOHFWLYLW\3F\DUHUHFRJQLVHGXQGHUWKHPLFURVFRSH DQG EHFDXVH VDPSOH LQFXEDWLRQ LV QRW UHTXLUHG &DUSHQWHU DQG &DPSEHOO $IIURQWL DQG 0DUVKDOO 7KH in situ SKRWRV\QWKHWLF UDWH RI 3F\ HVWLPDWHG E\ )'& DQG E\ WKH & LQFRUSRUDWLRQ PHWKRG KDYH UHFHQWO\ EHHQ FRPSDUHG LQ IUHVK ZDWHU &DOOLHULHWDO DQGWKHUHVXOWVDUHYHU\VLPLODU GHVSLWH WKHSUHVHQFH RIPDUNHG IOXFWXDWLRQV 7KH XVH RI WKH )'& PHWKRG IRU in situ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
&KDSWHU 3LFRSODQWRQ DQG RWKHU QRQEORRP IRUPHUV
&KORURSK\OO D J, Fig. 8. 7KHUHODWLRQEHWZHHQ$33&+/&RQWULEXWLRQ DQG WRWDO&+/DIURP9|U|VHWDO
UROHV RI $33 SRSXODWLRQV LQ ODFXVWULQH SODQNWRQ FRPPXQLWLHV 7KLVKDVUHFHQWO\EHHQVKRZQWREHWKH FDVH LQ PDULQH HFRV\VWHPV /LQGHOO DQG 3RVW &KDUS\DQG%ODQFKRW 6WRFNQHU V PRGHO DSSOLHV TXLWH ZHOO WR WKH REVHUYHG $33 FRQWULEXWLRQ WR WRWDO SK\WRSODQNWRQ ELRPDVVDORQJDWURSKLFJUDGLHQW 6¡QGHUJDDUG IRXQGWKDWDOWKRXJKWKHDFWXDOYDOXHVRI$33ELRPDVV ZHUH UDWKHU VLPLODU DPRQJ ODNHV WKH UHODWLYH LPSRUWDQFH ZDV KLJKHVW LQ PRUH ROLJRWURSKLF ODNHV DQGORZHULQHXWURSKLFODNHV 7KHVHYHQ'DQLVKODNHV KH VWXGLHG RQH DFLGLF WKH RWKHUV VKDOORZ K\SHUWURSKLF DQG GHHS PHVRHXWURSKLF H[KLELW &KO FRQFHQWUDWLRQVUDQJLQJIURP J/ ,WLV LQWHUHVWLQJWRFRPSDUH 6¡QGHUJDDUG VGDWDZLWK VLPLODU GDWD REWDLQHG LQ D VWXG\ RI ODNHV WKDW LQFOXGHG KLJK DOWLWXGH FOHDU ODNHV GHHS ODUJH VXEDOSLQHODNHV ODUJH VKDOORZODNHV VPDOOUHVHUYRLUV DQG ILVKSRQGV )LJ 9|U|V HWDO $FURVV VXFK D WURSKLF JUDGLHQW &KO FRQFHQWUDWLRQV UDQJHG EHWZHHQDQG J/7KHUHODWLYHFRQWULEXWLRQ RI 3F\ FDQ EH PRUH WKDQ RI WKH WRWDO
SK\WRSODQNWRQELRPDVVLQODNHVZLWK&KOEHORZ J / ZKHUHDV DERYH WKH WKUHVKROG RI J &KO / WKH FRQWULEXWLRQ RIWKH 3F\ IUDFWLRQ GRHV QRW H[FHHG 9|U|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|U|VHWDO 8VLQJWKH PRUH H[WHQVLYH GDWDEDVH RI9|U|V HW DO ZHKDYH DWWHPSWHGWR LQGLFDWH D JHQHUDO WUHQG LQ D VFKHPDWLF FRQFHSW LOOXVWUDWLQJ WKH DUHDV RI SRVVLEOH RFFXUUHQFH RI 3F\ QXPEHUV DQG WKHLU
-6WRFNQHU& &DOOLHULDQG* &URQEHUJ
&KORURSK\OO a J , Fig. 9. 3F\DEXQGDQFHDQGHXSKRWLF]RQH&+/DFRQFHQWUDWLRQVLQODNHVRILQFUHDVLQJWURSKLFVWDWHIURP9|U|VHWDO
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
* )DFWRUV$IIHFWLQJ3F\*URZWK 3URGXFWLRQ DQG'LVWULEXWLRQ /LJKW 8QGHUZDWHU OLJKW FRQGLWLRQV DUH H[WUHPHO\ LPSRUWDQW IRU WKH JURZWK DQG SURGXFWLRQ RI SHODJLF SKRWRDXWRWURSKV DQGWKHFKDQJH LQOLJKWTXDQWLW\DQG TXDOLW\ PRGXODWHV WKH DOJDO JURZWK UHVSRQVH 7KH JUHDW DEXQGDQFH RI 3F\ FHOOV LQ PRVW ODNHV GHPRQVWUDWHV WKH HIILFLHQF\ RI WKHLU DFFOLPDWLRQ PHFKDQLVPV WR FKDQJLQJ OLJKW FRQGLWLRQV DQG
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
&KDSWHU 3LFRSODQNWRQDQGRWKHU QRQEORRP IRUPHUV
Fig.10. 6FKHPDWLF GUDIW RI WKH DUHDV RI RFFXUUHQFH RI 3F\ DEXQGDQFH DQG WKHLU UHODWLYH FRQWULEXWLRQ WR WRWDO SK\WRSODQNWRQ ELRPDVV
ZLWK D JURZWK VDWXUDWLRQ DW PRO SKRWRQ PV )LJ 7KLV ODWWHU YDOXH FRUUHVSRQGV WR WKH OLJKW LUUDGLDQFHZKHUHLQQDWXUDOV\VWHPVWKHPD[LPXPFHOO DEXQGDQFH DQGSKRWRV\QWKHWLF DFWLYLW\ZDV IRXQG & &DOOLHULXQSXEOLVKHG 7KH.DQDDQG*OLEHUWD E UHVXOWV HYLGHQWO\ FRQWUDGLFW WKRVH RI PDQ\ RI WKH SUHYLRXV VWXGLHV DQG WKHSUHYDOHQWEHOLHIWKDW3F\ DUH VKDGHWROHUDQWRUJDQLVPV +RZHYHULWLVLPSRUWDQWWR QRWH WKDW .DQD DQG *OLEHUW XVHG Synechococcus FXOWXUHV DOUHDG\ DGDSWHG WR KLJK LUUDGLDQFH LQ WKHLU H[SHULPHQWV JLYLQJ WKHP WKH QHFHVVDU\ WLPH WR DFFOLPDWHLQWKLVFDVHORQJHUWKDQDJHQHUDWLRQWLPH 7KH VXFFHVV RI 3F\ XQGHU ORZ OLJKW FRQGLWLRQV LV WLJKWO\ FRXSOHG ZLWK FRPSHWLWLRQ IRU OLPLWLQJ QXWULHQWV DQGORZOLJKWVHHPV WR JLYHSynechococcus WKHDGYDQWDJHEXWRQO\XQGHUORZ3FRQGLWLRQV:HKU $QRWKHU W\SLFDO UHVSRQVH WR ORZ OLJKW LV DQ LQFUHDVH LQ DFFHVVRU\ SLJPHQWV UHODWLYH WR &KO D )DONRZVNL DQG /D5RFKH DQG QLWURJHQ OLPLWDWLRQPD\EHUHVSRQVLEOHIRUDORZ3(&KODUDWLR 0F0XUWHU DQG 3LFN (YHQ WKRXJK PDQ\ JHQHWLFDOO\ GLVWLQFW Synechococcus VWUDLQV KDYH EHHQ IRXQG(UQVWHWDO LWLVVWLOOKHOSIXOWREURDGO\
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ynechococcus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ynechococcus FXOWXUH WKHUH LV D UDSLG LQWHUFKDQJH IURP D OLJKW VHQVLWLYH IRUP RI WKH ' SURWHLQ RI 36,, FHQWUHV )RUP , WR D IRUP OHVV VXVFHSWLEOHWRSKRWRLQKLELWLRQ)RUP &ODUNHWDO ,WKDV DOVR EHHQ VKRZQ WKDWGXULQJ DGDSWDWLRQ WR KLJK OLJKW LQWHQVLW\ WKHUH LV JHQH UHJXODWLRQ WR HQFRGH WKH )RUP ,, RI ' SURWHLQ .XONDUQL DQG *ROGHQ 7KHUHIRUHWKHSKRWRLQKLELWLRQFDQEH HIILFLHQWO\ FRXQWHUDFWHG DQG HYHQWXDOO\ RYHUFRPH &ODUN HW DO ,W LV LPSRUWDQW WR NHHS LQ PLQG WKDW WKHVH ODERUDWRU\ H[SHULPHQWV ZHUH UXQ XVLQJ D VLQJOH Synechococcus VWUDLQ 3&& DQG VLPLODU ZRUN PXVW EH H[WHQGHG WR RWKHU VWUDLQV EHIRUH JHQHUDOLVLQJWKHVHUHVXOWVWRQDWXUDOV\VWHPV ,WDSSHDUV WKDWLQ3F\WKH IOXFWXDWLRQ RIWKHUHODWLYH SLJPHQW FRQWHQW PD\ GHSHQG RQ OLJKW FOLPDWH :\PDQDQG)D\ +DXVFKLOGHWDO DQG WKDW RQO\ F\DQREDFWHULD FDSDEOH RI 3( V\QWKHVLV FDQ XQGHUJR FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ 7DQGHDX GH 0DUVDF 5HFHQW VWXGLHV E\ +DXVFKLOG HW DO GLVWLQJXLVKHG WKUHH SK\VLRORJLFDO JURXSV RI 3( FHOOV *URXS , ZLWK QR
-6WRFNQHU&&DOOLHULDQG*&URQEHUJ
Fig. 11. 7KHHIIHFWRILUUDGLDQFHRQJURZWKUDWHRISynechococcus :+IURP.DQD *OLEHUW D
DGDSWDWLRQ *URXS ,, ZLWK UHJXODWLRQ RI 3( DQG *URXS ,,, ZLWK UHJXODWLRQ RI 3( DQG 3& SLJPHQW 0DULQH Synechococcus VWUDLQV H[KLELWHG FKURPDWLF DGDSWDWLRQRIWKH*URXS,,W\SHZLWKDQLQFUHDVHLQ3( FRQFHQWUDWLRQ DQG DFFHOHUDWHG JURZWK 7KH FRPSDULVRQ RIWKH JURZWKUHVSRQVH RIWZR IUHVKZDWHU
VWUDLQV RI Synechococcus, RQH RI 3( W\SHWKHRWKHURI 3& W\SH GHPRQVWUDWHVWKHVHOHFWLYHYDOXHRI UHG OLJKW LQ VWLPXODWLQJ 3& DQG LQ VXSSUHVVLQJ 3( )LJ &DOOLHUL HW DO &DOOLHUL 7KH LPSRUWDQFH RI UHG OLJKWIRU3&DQG ELRPDVV SURGXFWLRQ KDVEHHQ
Fig. 12. ([SRQHQWLDOJURZWKUDWH.GLYG DVDIXQFWLRQRI OLJKWLQWHQVLWLHVIRU3(DQGIRU3&3F\FXOWXUHVIURP&DOOLHULHW DO
Fig.13. *URZWK UDWHV RI WZR VWUDLQV RI 6\QHFKRFRFFXV ZLWK 3( DQG 3& SLJPHQWV H[SRVHG WR JUHHQ DQG UHG OLJKW RI DQ HTXDO TXDQWXPIOX[ PRO SKRWRQ PV IURP&DOOLHULHW DO
&KDSWHU 3LFRSODQNWRQ DQG RWKHUQRQEORRPIRUPHUV
Fig. 14. 5HODWLYHFRQWULEXWLRQRI3(FHOOV WRWRWDO3F\DEXQGDQFHLQODNHV ZLWK XQGHUZDWHUOLJKWFOLPDWHH[SUHVVHG DV . 5(' . *5((1 IURP 9RURVHWDO
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
89% 7KH HIIHFW RI XOWUDYLROHW UDGLDWLRQ RQ DTXDWLF RUJDQLVPV KDV EHHQ ZLGHO\ UHYLHZHG +ROP+DQVHQ HWDO +l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
&KDSWHU 3LFRSODQNWRQ DQG RWKHU QRQEORRP IRUPHUV
Fig.15. 3KRWRV\QWKHWLF UHVSRQVH WR 89% SKRWRLQKLELWLRQ PHDVXUHG LQ WZR FHOO VL]H IUDFWLRQV P DQG!P LQ /DNH 0DJJLRUH 1RUWKHUQ,WDO\ 7KHUHODWLYHSKRWRLQKLELWLRQLVFDOFXODWHGE\WKHUDWLR89%H[SRVHG89%SURWHFWHGFDUERQXSWDNH%HUWRQLDQG&DOOLHUL VXEPLWWHG
6KRUWUHHG DQG LQ VHYHUDO ORZ S+ KXPLF 'DQLVK ODNHV 6¡QGHUJDDUG 7KH HIIHFWRIODNH DFLGLILFDWLRQ RQ WKH PLFURELDO FRPPXQLW\ FDQ EH LQGLUHFW WKURXJK DOWHUDWLRQ RI FRPPXQLW\ VWUXFWXUH DQG KHQFH FDUERQ IORZV WR KLJKHU WURSKLF OHYHOV RU GLUHFWE\ LQGXFWLRQ RISK\VLRORJLFDO VWUHVV ,Q UHFHQW VWXGLHV RQ D 6ZHGLVK DFLGLILHG ODNH EHIRUH DQG DIWHU OLPLQJ DQRQHGLEOH&3F\ Merismopedia tenuissima ZDV WKH GRPLQDQW VSHFLHV LQ WKH ODWH VXPPHU SK\WRSODQNWRQFRPPXQLW\LQWKHQDWXUDOO\DFLGLFODNH +RZHYHU WKLVSRSXODWLRQZDV UHPRYHGE\WKHOLPLQJ %HOO DQG 7UDQYLN %ORPTYLVW 8QIRUWXQDWHO\ WKH DXWKRUV SUHVHQW QR GDWD RQ 3F\ DEXQGDQFH LQ WKLV ODNH KRZHYHU WKH\ VXJJHVW D SUREDEOH DOOHORSDWKLF PHFKDQLVP WR H[SODLQ WKH SRSXODWLRQ G\QDPLFV RI Merismopedia tenuissimu %ORRPTYLVW 9UHGH ,QDQ,WDOLDQDFLGLF DOSLQH ODNH 3F\ SRSXODWLRQV DUH YHU\ ORZ DQG WKHLU FRQWULEXWLRQ WR WKH PLFURELDO IRRG ZHE DSSHDUV WR EH QHJOLJLEOH & &DOOLHUL XQSXEOLVKHG 7KH SUHVHQFH RI DVKLIWIURP3F\WRQHWSODQNWRQKDVEHHQGHVFULEHGLQ PHVRFRVP H[SHULPHQWV +DYHQV DQG +HDWK
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
1XWULHQWV 'DWD IURP ERWK ILHOG DQG ODERUDWRU\ KDYH GHPRQVWUDWHG WKDW 3F\ DUH VHOGRP 3OLPLWHG EXW GR VKRZ LQFUHDVHG JURZWK DQG SURGXFWLRQ UHVSRQVHV ZKHQ 1 LV DGGHG 6XWWOH HW DO :HKU 3F\ DQG VRPH &3F\ SUHIHU DPPRQLXP1 IRU JURZWK 0DVWDOD HW DO DQG KDYH VKRZQ D ORZHU FRPSHWLWLYHQHVV IRU QLWUDWH1 QLWUDWHUHGXFWDVH
-6WRFNQHU&&DOOLHULDQG*&URQEHUJ
Fig. 16. ,QVLWXSKRWRV\QWKHWLFUDWHV RI3F\PHDVXUHGZLWK& WHFKQLTXHDQGLQFXEDWHGIRUKLQTXDUW]DQGP\ODUWXEHV LQ/DNH 0DJJLRUH 1RUWKHUQ,WDO\&&DOOLHULXQSXEOLVKHGGDWD
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ynechococcus FHOO F\FOH LQ PDULQH V\VWHPV 9DXORW HW DO REWDLQHG UHVXOWV VXJJHVWLQJ WKH LPSRUWDQFH RI 3 DV D OLPLWLQJ QXWULHQW RQO\ GXULQJ FHUWDLQ WLPHV RI WKH \HDU ZKLFK DUH FRQWUDU\ WR
:HKU¶V UHVXOWV DQG WR WKH JHQHUDO FRQVHQVXV IRU 3F\ LQ IUHVKZDWHU 6WRFNQHU +RZHYHU FDXWLRQ PXVW EH H[HUFLVHG LQ LQWHUSUHWLQJ WKHUHVXOWV RI9DXORWHWDO EHFDXVH ZKHQ3 LV DGGHG WKH FHOO F\FOH YDULDEOHV UHVSRQG YHU\ TXLFNO\ VR WKH DSSOLFDELOLW\ RI WKHLU ILQGLQJV WR in situ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
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV
VHOHFWLRQ RI UHDOLVWLF SDUDPHWHUV WR PRQLWRU ZKLFKUHVSRQGUDSLGO\LQQXWULHQWELRDVVD\V WKH SURSHU HYDOXDWLRQ RI LQWHUDFWLRQV DPRQJ GLIIHUHQW OLPLWLQJ UHVRXUFHV DQG ELRWLF IDFWRUV HJ JUD]LQJ
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
Fig.17. 3HUFHQWDJH89% SKRWRLQKLELWLRQRIQDWXUDO DXWRWURSKLF DQG KHWHURWURSKLF PLFURRUJDQLVPV H[SRVHG WR QDWXUDO DQG DUWLILFLDOO\ LQFUHDVHG 89% UDGLDWLRQ & &DOOLHUL XQSXEOLVKHG GDWD
Fig.18. 3URJUHVVLYH GLVDSSHDUDQFH RI3F\ LQ WKH $33 FRPPXQLW\ LQWHPSHUDWH ODNHV ZLWK DGHFUHDVLQJS+ 6WRFNQHUDQG 6KRUWUHHG
KDV EHHQ XVHG WR VWRS SURWR]RDQ JUD]LQJ DFWLYLW\ RQ 3F\ &DPSEHOO DQG &DUSHQWHU E &DURQ HW DO 5HFHQWO\/LXHW DO FKRVH NDQDPLFLQ DV DQ HIIHFWLYH JURZWK LQKLELWRU RI Synechococcus DQG 3URFKORURFRFFXV WR HVWLPDWH JURZWK DQG JUD]LQJ UDWHV 8VLQJWKLV QHZ DSSURDFK3F\ JUD]LQJ PRUWDOLW\ KDVEHHQHVWLPDWHGWR UDQJHIURP RI JURZWK UDWH LQ PDULQH V\VWHPV /LX HW DO /DQGU\ HW DO KDYH SURSRVHG D UHILQHG GLOXWLRQ WHFKQLTXH WR RYHUFRPH WKH DVVXPSWLRQ WKDW JUD]LQJ LPSDFW YDULHV LQ GLUHFW SURSRUWLRQ WR WKH GLOXWLRQ RI JUD]HU SRSXODWLRQ GHQVLW\ ,Q WKH QHZ SURWRFRO WKH\ LQWURGXFH IOXRUHVFHQW ODEHOOHG FHOOV DQG IORZ F\WRPHWU\ WR PHDVXUH WKH UHODWLYH JUD]LQJ DFWLYLW\ LQ HDFKGLOXWLRQWUHDWPHQW $\XNDL KDV VXJJHVWHG WKH XVH RI)'& WR FRQWURO DQG PRQLWRUWKHJURZWK UDWH LQ GLOXWLRQPHDVXUHPHQWV D WHFKQLTXHWKDWKDV QRW \HW EHHQ DSSOLHG LQ IUHVK ZDWHU V\VWHPV 7KH PHWKRG RI IOXRUHVFHQWO\ODEHOOHGSDUWLFOHVLQ JUD]LQJH[SHULPHQWV JDLQHG QHZ LPSHWXV ZKHQ LQHUW DQG OHVV HIIHFWLYH PLFURVSKHUHV ZHUH VXEVWLWXWHG IRU IOXRUHVFHQWO\ PDUNHG EDFWHULD DQG 3F\ )/% )/& 6KHUU HW DO 6KHUU DQG 6KHUU
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
-6WRFNQHU&&DOOLHULDQG* &URQEHUJ
ã
LQSUHVV LPHN HWDO KDYHVXPPDULVHGWKUHH HFRORJLFDO FDWHJRULHV RI IUHVKZDWHU FLOLDWHV ZLWK GLIIHUHQW IHHGLQJ VWUDWHJLHV DQG D GHFUHDVLQJ LPSRUWDQFH RISLFRVL]HSUH\LQ WKHLUGLHW $PRQJWKH PRVWHIILFLHQWVXVSHQVLRQIHHGHUVWKHUH DUHVRPHYHU\ DFWLYH 3F\ JUD]HUV HJ Vorticella aquadulcis, Halteria grandinella, Cyclidium DQG Strobilidium hexachinetum. 7KHVHSURWR]RDDUH DEOHWR JUD]H FHOOVKUHVSHFWLYHO\ ZLWKFOHDUDQFHUDWHV UHDFKLQJ YDOXHV RI Q/ FHOOV K 6LPHN HW DO 6RPHUDSWRULDO IHHGHUV VXFK DV Urotricha DQG Balanion planktonicum KDYH JUD]LQJ UDWHV RI DQG $33 FHOOV KUHVSHFWLYHO\ DQG FOHDUDQFH UDWHV RI DERXWQ/FHOOVK 6LPHNHWDO 7KHUHDUHPDQ\OLQNRUJDQLVPVJUD]LQJRQ3F\DQG EDFWHULD WKDW SURYLGH LPSRUWDQW FRQQHFWLRQV EHWZHHQ WKHFODVVLFSHODJLFIRRGZHEDQGPLFURELDOIRRGZHEV 3ODWH &XUUHQW PRGHOV RIWKH LPSDFWV RISHODJLF PLFURELDOIRRGZHEVRQPDWHULDODQGHQHUJ\IORZVDUH EDVHG RQ WKH SUHVHQFH RU DEVHQFH RI Daphnia, RU RWKHU ODUJH NH\VWRQH JUD]HUV WKDW FDQ QRW RQO\ LQWHUIHUH ZLWK RWKHU 3F\ SUHGDWRUV IODJHOODWHV FLOLDWHV PLFUR]RRSODQNWRQ EXW DOVR KDYH WKH FDSDELOLW\ WR XVH WKHP DV SUH\ 6WRFNQHU 3RUWHU :HLVVH DQG 6WRFNQHU -XUJHQV &DOOLHUL HW DO LQ SUHVV 7RJHWKHU ZLWK Daphnia VHYHUDO RWKHU FODGRFHUDQ JHQHUD VXFK DV Bosmina, Eubosmina DQG Ceriodaphnia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
&KDSWHU 3LFRSODQNWRQDQGRWKHUQRQEORRP IRUPHUV $WSUHVHQWWKHUHDUHIHZVWXGLHV DQGWKHVHDUHDOOLQ PDULQH V\VWHPV RQ WKH UROH RIELYDOYH DQG JDVWURSRG ODUYDH DV SRVVLEOH SLFRSODQNWLYRUHV ,Q RQH RI WKHVH %HOO KDVGHPRQVWUDWHGWKDWCrepidula aculeata DQG Littoraria scabra, WZR JDVWURSRG ODUYDH FDQ WKULYH RQ D 3F\ DQG EDFWHULD GLHW )XUWKHUPRUH WKH ODUYDH RI D ELYDOYH Mercenaria mercenaria, KDYH UHFHQWO\ EHHQ VKRZQ WR LQJHVW DQG JURZ RQ D Synechococcus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in situ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erismopedia tenuissima WR EH UHVLVWDQW WR JUD]LQJ E\ WKH GRPLQDQW JUD]HU Eudiaptomus LQ D FOHDUZDWHUROLJRWURSKLFODNHLQFHQWUDO6ZHGHQ7KXV IURP ZKDW OLWWOH LV NQRZQ DW WKLV WLPH ZH FDQ FRQFOXGH WKDW LW LV OLNHO\ WKDW WKH PXFLODJLQRXV FRORQLDO PRUSKRORJ\ DV IRXQG LQ ODUJHU FRORQLDO F\DQREDFWHULD LV DQ HIIHFWLYH DQWLJUD]LQJ DGDSWDWLRQ WKDW SHUKDSV ZKHQ FRXSOHG ZLWK WKH SRVVLELOLW\ RI DOOHORSDWKLF VWUDWHJLHV LQ &3F\ FRORQLHV FUHDWH WKH SHUIHFW FRXSOHG DQWLSUHGDWLRQGHYLFH IRU WKHVH &3F\ VSHFLHV 7KLV PD\ YHU\ ZHOO EH WKH UHDVRQ IRU WKHLU XELTXLW\DQGDEXQGDQFHLQODNHV
+ 0LFURELDO)RRG:HEV $ JOLPSVH DW WKH $33 DQG 3F\ OLWHUDWXUH IURP WKH HDUO\ V VKRZV DPDUNHG HVFDODWLRQ RILQWHUHVWE\
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µVLQNV¶ ZLWK ODUJH PDLQWHQDQFH FRVWV IRU FRDVWDO PDULQH SHODJLF HFRV\VWHPV 6HYHUDO VWXGLHV KDYH VXJJHVWHG WKDW PDWHULDO DQG HQHUJ\ IOX[ ZLWKLQ 0): LQ ODNHV FDQ EH HQKDQFHG E\ KDYLQJ D LQWHUPHGLDWH µOLQN¶ RU µNH\VWRQH¶ JUD]HU OLNH Daphnia SUHVHQW LQ WKH V\VWHP WR PRYH FDUERQ IURP 0): WR ILVK 6WRFNQHU 3RUWHU HW DO 3DFH HW DO +RZHYHU DV 6WRFNQHU DQG 0DF,VDDF KDYH VKRZQ HYHQ LQ WKH DEVHQFH RI WKH NH\VWRQH JUD]HU 0): FDQ IXQQHO FDUERQ DOEHLW LQHIILFLHQWO\ WR DOO WURSKLF OHYHOV 7KXV ZH XUJH OLPQRORJLVWV QRW WR LJQRUH WKH FRQWULEXWLRQV RI 3F\ DQG DOVR &3F\ WR HQHUJ\ IORZV LQ SODQNWRQLF 0): RI ODNHV RI DOO WURSKLF W\SHV
9 &RQFOXVLRQV 1HZ SHUVSHFWLYHV RQ 3F\ DQG &3F\ HFRORJ\ ZLOO XOWLPDWHO\UHO\RQWKHLPSURYHPHQWDQGLQFRUSRUDWLRQ RIQHZ VDPSOLQJ DQG HQXPHUDWLRQ SURWRFROV DV ZHOO DVWKHDSSOLFDWLRQRIQHZPROHFXODUPHWKRGV WRVWXG\ JHQHWLF GLYHUVLW\ LQ 3F\ DQG &3F\ FRPPXQLWLHV 7KHVH DGYDQFHV VKRXOG LPSURYH RXU FRPSUHKHQVLRQ DQG XQGHUVWDQGLQJ RI 3F\ VSDWLDO DQG WHPSRUDO YDULDWLRQ ZLWKLQ WKH IUDPHZRUN RI D QHZ FORQDO
-6WRFNQHU&&DOOLHULDQG*&URQEHUJ VXFFHVVLRQ WKHRU\ 5HFHQW ILQGLQJV VXJJHVW WKDW WKH VHDVRQDO FKDQJHV RI3F\DVVHPEODJHV LQ ODNHV DUH WKH UHVXOW RIWKH GHYHORSPHQW RID VXFFHVVLRQ RIGLIIHUHQW VWUDLQV RUFORQHV LQWHUDFWLQJ ZLWK PXOWLSOHELRWLF DQG DELRWLF YDULDEOHV (UQVW (UQVW HW DO /RRNLQJ DW WKH FRQFHSW RI SK\WRSODQNWRQ VSHFLHV VXFFHVVLRQ LQ ODNHV 3F\ FRPPXQLWLHV PD\ QRZ DVVXPH KHLJKWHQHG VLJQLILFDQFH IRU WKH FRPSUHKHQVLRQRI³VSHFLHV DVVRFLDWLRQV´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³FDQDULHV LQ WKH FRDO PLQH´ VLJQDOOLQJERWKLPSHQGLQJFKDQJHV LQ WKH ZD\QXWULHQWV DUHUHF\FOHG DQGHQHUJ\IORZV WKURXJK SHODJLF IRRG ZHEV DV SK\WRSODQNWRQ FRPPXQLWLHV PRYHIURPUHOLDQFHRQH[RJHQRXVQXWULHQWVXSSOLHVWR VXSSRUWQHZSURGXFWLRQ WRUHF\FOHG QXWULHQWV :HLVVH DQG 6WRFNQHU :LOO WKH JUDGXDO GHFOLQH RI GLDWRPEDVHGIRRGFKDLQVRU µFODVVLF¶ IRRG ZHEV DQG WKH LQFUHDVHG LPSRUWDQFH RI 3F\ EDVHG µPLFURELDO¶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
&KDSWHU 3LFRSODQNWRQ DQG RWKHUQRQEORRP IRUPHUV DUHIDLUTXHVWLRQVEHFDXVH3F\ DQG&3F\DUHRXUPRVW DQFLHQW H[WDQW VXUYLYRUV RISULPRUGLDO VHDV FDSDEOH RI DGDSWLQJ WR H[WUHPH HQYLURQPHQWV 7KHVH DUH FUXFLDO TXHVWLRQV WKDWQHHG DQVZHUV LIZH DUHWR SUHVHUYH WKH TXDOLW\ DQG LQWHJULW\ RI IUHVK ZDWHU DQG JOREDO 2FHDQ ILVKHULHV OHW DORQHWKH YLDELOLW\ RIWKH KXPDQ VSHFLHV RQHDUWK
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
5HIHUHQFHV $IIURQWL /)DQG0DUVKDOO+* 8VLQJ IUHTXHQF\RIGLYLGLQJ FHOOV LQ HVWLPDWLQJ DXWRWURSKLF SLFRSODQNWRQ JURZWK DQG SURGXFWLYLW\ LQ WKH &KHVDSHDNH %D\ +\GURELRORJLD cOYLN * 3ODQNWRQ$OJHQ QRUZHJLVFKHU $XVWHUQ ROOHQ , 6\VWHPDWLNXQG 9RUNRPPHQGHU$UWHQ%HUJHQV 0XV c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iUHN - 7RZDUGV D IRUPDO GHVFULSWLRQ RI D QHZ VSHFLHV RI 6\QHFKRFRFFXV &\DQREDFWHULD&\DQRSK\FHDH IURP IUHVKZDWHU SLFRSODQNWRQ $UFKLY+\GURELRO6XSSO$OJRORJLFDO6WXGLHV %HOO -/ 3DWFKHV DQG SLFRSODQNWRQ (IIHFWV RQ ODUYDO OLIH VSDQVRQJDVWURSRGODUYDH$P=RRO $ %HOO 57DQG 7UDQYLN/ ,PSDFWRIDFLGLILFDWLRQ DQG OLPLQJ RQWKHPLFURELDOHFRORJ\RIODNHV$PELR %HUWRQL 5 7KH URXWLQH XVH 2I $QRGLVF ILOWHUV :LWK DXWRPDWLF &+1 DQDO\VHUV0HP,VW,WDO,GURELRO
%HUWRQL 5 DQG&DOOLHUL & 2UJDQLF PDWWHU DQG GHFRPSRVHUV LQ/DJR0DJJLRUHDSOXULDQQXDOVWXG\ 0HP,VW,WDO,GURELRO %HUWRQL 5 DQG &DOOLHUL & 0LFURELDO ORRS DQG RUJDQLF FDUERQ LQ ODUJH VRXWKHUQ $OSLQH ODNHV ,Q 0RVHOOR 5 DQG *LXVVDQL*HGV 'RFXPHQWD,VWLWDO,GURELRO %HUWRQL 5 DQG &DOOLHUL & VXEPLWWHG 7KH HIIHFWV RI 89% UDGLDWLRQ RQ IUHVKZDWHU DXWRWURSKLF DQG KHWHURWURSKLF SLFRSODQNWRQLQDVXEDOSLQHODNH %HUWRQL 5 3XJQHWWL $ DQG %DUEDUHZLF] & /D UDGLD]LRQH 89% QHLODJKLHGLVXRLHIIHWWL VXOOHFRPXQLWj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p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
&DOOLHUL & DQG 6WRFNQHU -* LQ SUHVV 3LFREOXHV VXFFHVV DFURVV IUHVKZDWHUWURSKLFJUDGLHQW/DNHV 5HVHUYRLUV &DPSEHOO/DQG&DUSHQWHU(- D 'LHOSDWWHUQRIFHOOGLYLVLRQ LQPDULQHSynechococcus VSS &\DQREDFWHULD XVHRIIUHTXHQF\ RI GLYLGLQJ FHOO WHFKQLTXH WR PHDVXUH JURZWK UDWH 0DU (FRO 3URJU6HU &DPSEHOO / DQG &DUSHQWHU (- E (VWLPDWLQJ WKH JUD]LQJ SUHVVXUH RI KHWHURWURSKLF QDQRSODQNWRQ RQ Synechococcus VSS XVLQJ WKH VHDZDWHU GLOXWLRQ DQG VHOHFWLYH LQKLELWRU WHFKQLTXHV 0DU(FRO3URJU6HU &DURQ '$ /LP (/ 0LFHOL * :DWHUEXU\ -% DQG 9DORLV ): *UD]LQJ DQG XWLOL]DWLRQ RI FKURRFRFFRLG F\DQREDFWHULD DQGKHWHURWURSKLFEDFWHULDE\SURWR]RDLQODERUDWRU\FXOWXUHVDQG FRDVWDOSODQNWRQFRPPXQLW\0DU(FRO3URJU6HU &DUSHQWHU(- DQG &DPSEHOO / 'LHO SDWWHUQ RIFHOO GLYLVLRQ DQG JURZWK UDWH RI Synechococcus VSS LQ /RQJ ,VODQG 6RXQG 0DU(FRO3URJU6HU &DUUHWR -, &DULJQDQ 02 'DOHR * DQG 'H 0DUFR 6* 2FFXUUHQFH RI P\FRVSRULQHOLNH DPLQR DFLGV LQ WKH UHG WLGH GLQRIODJHOODWH Alexandrium escavutum: 89SKRWRSURWHFWLYH FRPSRXQGV"-3ODQNWRQ5HV &KDUS\ / DQG %ODQFKRW - Prochlorococcus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ynechococcus SFF 3ODQWD &OHYHQ (- ,QGLUHFWO\ IOXRUHVFHQWO\ ODEHOOHG IODJHOODWHV (IFLF): D WRRO WR HVWLPDWH WKH SUHGDWLRQ RQ IUHHOLYLQJ KHWHURWURSKLFIODJHOODWHV-3ODQNWRQ5HV &UDLJ65 7KHGLVWULEXWLRQDQGFRQWULEXWLRQRISLFRSODQNWRQ WR GHHS SKRWRV\QWKHWLF OD\HUV LQ VRPH PHURPLFWLF ODNHV $FWD $FDG$ER &ULVWRIIHUVHQ . 9DULDWLRQ RI IHHGLQJ DFWLYLWLHV RI KHWHURWURSKLF QDQRIODJHOODWHV RQSLFRSODQNWRQ ,Q $PGW+ HG 3URWR]RRSODQNWRQ (FRORJ\ 0HWKRGV DQG 6HDVRQDO 6XFFHVVLRQV SS9RO &URQEHUJ * Cyanodictyon tubiforme D QHZ FKURRFRFFDO EOXHJUHHQ DOJDIURP/DNH%|UULQJHVM|Q 6FDQLD 6ZHGHQ $UFK +\GURELRO6XSSO $OJRORJLFDO6WXGLHV &URQEHUJ * Cyanothamnos plancticus JHQ HW VS QRY D QHZ FRORQLDO F\DQRSK\WH IURP DQ HXWURSKLF 6FDQLDQ ODNH 6ZHGHQ$UFK+\GURELRO6XSSO $OJRORJLFDO 6WXGLHV &URQEHUJ * DQG .RPiUHN - 3ODQNWLF &\DQRSURNDU\RWHV IRXQG LQ 6RXWK 6ZHGLVK ODNHV GXULQJ WKH ;,,WK ,QWHUQDWLRQDO 6\PSRVLXPRQ&\DQRSK\WH5HVHDUFK $OJRORJLFDO6WXGLHV &URQEHUJ * DQG :HLEXOO & Cyanodictyon imperfectum, D QHZ FKURRFRFFDO DOJD IURP /DNH 7UXPPHQ 6ZHGHQ $UFK +\GURELRO6XSSO$OJRORJLFDO6WXGLHV &XOOHQ -- 1HDOH 3- DQG /HVVHU 30 %LRORJLFDO ZHLJKWLQJ IXQFWLRQ IRU WKH LQKLELWLRQ RI SK\WRSODQNWRQ SKRWRV\QWKHVLV E\ XOWUDYLROHW UDGLDWLRQ 6FLHQFH
- 6WRFNQHU&&DOOLHULDQG* &URQEHUJ 'UHZV * 3UDXVHU + DQG 8KOPDQQ ' 0DVVHQYRUNRPPHQ YRQ Synechococcus pluncticus QRY VSHF HLQHU VROLWl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ynechococcus VSS F\DQREDFWHULD LVRODWHG IURP WKH SHODJLDO RI/DNH &RQVWDQFH )(060LFURELRO (FRO )DKQHQVWLHO */DQG &DUULFN +- 3KRWRWURSKLF SLFRSODQNWRQ LQ ODNHV +XURQ DQG 0LFKLJDQ DEXQGDQFH GLVWULEXWLRQ FRPSRVLWLRQ DQG FRQWULEXWLRQ WR ELRPDVV DQGSURGXFWLRQ &DQ - )LVK$TXDW6FL )DKQHQVWLHO*/&DUULFN+-5RJHUV&(DQG6LFNR*RDG/D 5HG IOXRUHVFLQJ SKRWRWURSKLF SLFRSODQNWRQ LQ WKH /DXUHQWLDQ *UHDW/DNHV ZKDWDUH WKH\DQGZKDW DUHWKH\ GRLQJ" ,QW5HYXH JHV +\GURELRO )DKQHQVWLHO */ 3DWWRQ 75 &DUULFN +- DQG 0F&RUPLFN 0- E 'LHO GLYLVLRQ F\FOH DQG JURZWK UDWHV RI Synechococcus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ynechococcus E\ ODUYDHRIWKHELYDOYHMercenaria mercenaria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
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV *RSKHQ0DQG*HOOHU: )LOWHU PHVK VL]HDQGIRRGSDUWLFOH XSWDNHE\Daphnia. 2HFRORJLD +l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ncorhynchus nerka *HR-RXUQDO +HVVHQ '2 9DQ'RQN ( DQG $QGHUVHQ 7 *URZWK UHVSRQVHV 3XSWDNH DQG ORVV RI IODJHOODH LQ Chlamydomonas reinhardfii H[SRVHGWR89%-3ODQNWRQ5HV +LFNHO % Cyanonephron styloides JHQ HW VS QRY D QHZ FKURRFRFFDO EOXHJUHHQ DOJDH &\DQRSK\WD IURP D EUDFNLVK ODNH $UFK +\GURELRO6XSSO $OJRORJLFDO 6WXGLHV +LFNHO % 7ZR QHZ FKURRFRFFDO F\DQRSK\WHV IURP D EUDFNLVK HQYLURQPHQW 6FKOHL)MRUG *HUPDQ\ $OJRORJLFDO 6WXGLHV +LQG ) (LQLJH QHXH XQG LQWHUHVVDQWH 3ODQNWRQEODXDOJHQ DXV GHU :HVWVORZDNH $UFK +\GURELRO6XSSO $OJRORJLFDO 6WXGLHV +LQGiN ) 2Q VRPH SODQNWRQLF FRFFRLG EOXHJUHHQ DOJDH FKDUDFWHULVWLF RI )HSUHFLSLWDWHV $UFK +\GURELRO 6XSSO $OJRORJLFDO6WXGLHV +LQG ) 7KH F\DQRSK\FHDQ JHQXV Lemmermanniella *HLWOHU $UFK +\GURELRO 6XSSO $OJRORJLFDO 6WXGLHV +ROP+DQVHQ 2 /XELQ ' DQG +HOEOLQJ (: 8OWUDYLROHW UDGLDWLRQ DQG LWV HIIHFWV RQ RUJDQLVPV LQ DTXDWLF HQYLURQPHQWV ,Q
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yV$OLy& &RQWURO RI KHWHURWURSKLF EDFWHULD DQG SURWR]RDQV E\ Daphnia pulex LQ WKHHSLOLPQLRQRI/DNH&LVz$UFK+\GURELRO .DQD70DQG*OLEHUW30 D (IIHFWRILUUDGLDQFHVXSWR ( PV RQ PDULQH Synechococcus :+ , *URZWK SLJPHQWDWLRQDQGFHOOFRPSRVLWLRQ'HHS6HD5HV .DQD70DQG*OLEHUW30 E (IIHFWRILUUDGLDQFHVXSWR (P V RQ PDULQH Synechococcus :+,, 3KRWRV\QWKHWLF UHVSRQVHVDQGPHFKDQLVPV 'HHS6HD5HV .DUHQW]'0F(XHQ)6 /DQG0&DQG'XQODS&: 6XUYH\ RIP\FRVSRULQHOLNHDPLQR DFLG FRPSRXQGV LQ $QWDUFWLF PDULQH RUJDQLVPV SRWHQWLDO SURWHFWLRQ IURP XOWUDYLROHW H[SRVXUH 0DU %LRO .L¡UERH 7 DQG +DQVHQ - 3K\WRSODQNWRQ DJJUHJDWH IRUPDWLRQ REVHUYDWLRQV RI SDWWHUQV DQG PHFKDQLVPV RI FHOO VWLFNLQJ DQG WKH VLJQLILFDQFH RI H[RSRO\PHULF PDWHULDO - 3ODQNWRQ5HV .LUFKPDQ '/ 6WDWLVWLFDO DQDO\VLV RI GLUHFW FRXQWV RI PLFURELDODEXQGDQFH,Q.HPS3) 6KHUU%)6KHUU(%DQG&ROH -- HGV +DQGERRN RI 0HWKRGV LQ $TXDWLF 0LFURELDO (FRORJ\ SS/HZLV3XEO .OXW (0 DQG6WRFNQHU-* 3LFRSODQNWRQDVVRFLDWLRQV LQDQ XOWUDROLJRWURSKLF ODNH RQ 9DQFRXYHU ,VODQG %ULWLVK &ROXPELD &DQ-)LVK$TXDW6FL .RPILUHN - 'LH WD[RQRPLVFKH 5HYLVLRQ GHU SODQNWLVFKHQ %ODXDOJHQ GHU WVFKHFKRVORYDNHL .RPILUHN - (WWO + $OJRORJLFDO 6WXGLHVSS .RPILUHN- 7D[RQRPLF UHYLHZ RIWKH JHQHUD Synechocystis 6$89 Synechococcus 1DJ DQG Cyanothece JHQ QRY&\DQRSK\FHDH $UFK3URWLVWHQN .RPILUHN - 7RZDUGV D FRPELQHG DSSURDFK IRU WKH WD[RQRPLF DQG VSHFLHV GHOLPLWDWLRQ RI SLFRSODQNWLF F\DQRSURNDU\RWHV$OJRORJLFDO6WXGLHV .RPILUHN - DQG .OLQJ + 9DULDWLRQ LQ VL[ SODQNWRQLF F\DQRSK\WH JHQHUD LQ /DNH 9LFWRULD (DVW $IULFD $OJRORJLFDO 6WXGLHV .RPILUHN - DQG .RPiUNRYi/HJQHURYi - 9DULDELOLW\ RI VRPH SODQNWLF JRPSKRVSKDHULRG F\DQRSURFDU\RWHV LQ QRUWKHUQ ODNHV1RUG-%RW .RPiUNRYi - DQG &URQEHUJ * Lemmermanniella pallida /(00 *(,7/ IURP 6RXWK 6ZHGLVK ODNHV $UFK +\GURELRO 6XSSO $OJRORJLFDO6WXGLHV .RPiUNRYi/HJQHURYi - DQG &URQEHUJ * 3ODQNWLF EOXH JUHHQ DOJDH IURP ODNHV LQ 6RXWK 6FDQLD 6ZHGHQ 3DUW , &KURRFRFFDOHV$OJRORJLFDO6WXGLHV .RQGR7 6WUD\HU&$ .XONDUQL5' 7D\ORU: ,VKLXU0 *ROGHQ 66 DQG -RKQVRQ &+ &LUFDGLDQ UK\WKPV LQ SURNDU\RWHV OXFLIHUDVH DV D UHSRUWHU RI FLUFDGLDQ JHQH H[SUHVVLRQ LQ F\DQREDFWHULD3URF1DWO$FDG6FL86
.XONDUQL 5' DQG *ROGHQ 66 )RUP ,, RI' LV LPSRUWDQW GXULQJ WUDQVLWLRQ IURP VWDQGDUG WR KLJK OLJKW LQWHQVLW\ LQ Synechococcus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gIY.JO9HWHQVN $NDG)|UKDQGO /HPPHUPDQQ ( 'DV 3ODQNWRQ VFKZHGLVFKHU *HZDVVHU $UN%RW /LQGHO ' DQG 3RVW $) 8OWUDSK\WRSODQNWRQ VXFFHVVLRQ LV WULJJHUHG E\ GHHS ZLQWHU PL[LQJ LQ WKH *XOI RI $TDED (LODW 5HG6HD/LPQRO2FHDQRJU /LX = (VWLPDWLQJ WKH JUD]LQJ SUHVVXUH RI KHWHURWURSKLF QDQRSODQNWRQ RQ 6\QHFKRFRFFXV VSS LQ WKH VXUIDFH FRDVWDO ZDWHU XVLQJ VHOHFWLYH LQKLELWRU WHFKQLTXHV 'RQJD\0DU 6FL /LX+ &DPSEHOO /DQG /DQGU\05 *URZWK DQG PRUWDOLW\ UDWHV RI Prochlorococcus DQG Synechococcus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pO\ * 6KDILN +0 DQG 9RURV / 1XWULHQW UHTXLUHPHQW DQG JURZWK RI D Synechococcus VSHFLHV LVRODWHG IURP /DNH %DODWRQ ,QW 5HYXH JHV+\GURELRO 0F0XUWKHU+-&DQG3LFN) )OXRUHVFHQFHFKDUDFWHULVWLFV RI D QDWXUDO DVVHPEODJH RI IUHVKZDWHU SLFRF\DQREDFWHULD - 3ODQNWRQ5HV 0H\HU% $ QHZ VSHFLHV RICyanodictyon &\DQRSK\FHDH &KURRFRFFDOHV SODQNWLF LQ HXWURSKLF ODNHV $OJRORJLFDO 6WXGLHV 0RHOOHU 5( &RQWULEXWLRQ RI XOWUDYLROHW UDGLDWLRQ 89$ 89% WR SKRWRLQKLELWLRQ RIHSLOLPQHWLF SK\WRSODQNWRQ LQ ODNHV RIGLIIHULQJ89WUDQVSDUHQF\$UFK+\GURELRO %HLK 0RQJHU %& DQG /DQGU\ 05 6L]H VHOHFWLYLW\ JUD]LQJ E\ KHWHURWURSKLF QDQRIODJHOODWHV DQ DQO\VLV XVLQJ OLYHVWDLQHG EDFWHULD DQG GXDOEHDP IORZ F\WRPHWU\ ,Q %MRHUVHQ 3. DQG 5LHPDQQ % HGV 0LFURELDO (FRORJ\ RI 3HODJLF (QYLURQPHQWV SS
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ynechococcus VSS JURZWKUDWHVDQGJUD]LQJSUHVVXUHE\KHWHURWURSKLFQDQRSODQNWRQ LQWKH(QJOLVKFKDQQHODQGWKH&HOWLF 6HD $FWD2FHDQRO6LQ 2YHUEHFN- 'DV1DQQRSODQNWRQ $OJHQ GHU5XJHQVFKHQ %UDFNZDVVHU DOV +DXSWSURGX]HQW LQ $EKDQJLJNHLW YRP 6DO]JHKDOW.LHOHU0HHUHVIRUVFKXQJHQ 3DFH 0/ 0F0DQXV *% DQG )LQGOD\ 6(* 3ODQNWRQ FRPPXQLW\ VWUXFWXUH GHWHUPLQHV WKH IDWH RIEDFWHULDO SURGXFWLRQ LQDWHPSHUDWHODNH/LPQRO2FHDQRJU 3DGLViN- .ULHQLW]/ .RVFKHO5DQG1HGRPD- 'HHSOD\HU DXWRWURSKLF SLFRSODQNWRQ PD[LPXP LQ WKH ROLJRWURSKLF /DNH 6WHFKOLQ *HUPDQ\ RULJLQ DFWLYLW\ GHYHORSPHQW DQG HURVLRQ (XU- 3K\FRO 3DGLVi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yV$OLy & DQG %URFN 7' 7KH LPSRUWDQFH RI DWWDFKPHQW WR SDUWLFOHV IRU SODQNWRQLF EDFWHULD $UFK +\GURELRO 3HUQWKDOHU- âLPHN . 6DWWOHU% 6FKZDU]HQEDFKHU$ %RENRYD - DQG3VHQQHU5 D 6KRUWWHUPFKDQJHV RISURWR]RDQFRQWURO RQ DXWRWURSKLF SLFRSODQNWRQ LQ DQ ROLJRPHVRWURSKLF ODNH - 3ODQNWRQ5HV 3HPWKDOHU- 6DWWOHU% âLPHN. 6FKZDU]HQEDFKHU% DQG 3VHQQHU 5 E 7RSGRZQHIIHFWV RQ WKHVL]HELRPDVVGLVWULEXWLRQ RI D IUHVKZDWHU EDFWHULRSODQNWRQ FRPPXQLW\ $TXDW 0LFURE (FRO 3HWHUVHQ 5 &DUERQ XSWDNH E\ SLFRSODQNWRQ DQG WRWDO SK\WRSODQNWRQ LQ HLJKW 1HZ =HDODQG ODNHV ,QW 5HYXH JHV +\GURELRO
&KDSWHU 3LFRSODQNWRQ DQG RWKHU QRQEORRP IRUPHUV 3LFN )5 7KH DEXQGDQFH DQG FRPSRVLWLRQ RI IUHVKZDWHU SLFRF\DQREDFWHULD LQ UHODWLRQ WR OLJKW SHQHWUDWLRQ /LPQRO 2FHDQRJU 3LFN )5 DQG $JEHWL '0 7KH VHDVRQDO G\QDPLF DQG FRPSRVLWLRQ RI SKRWRV\QWKHWLF SLFRSODQNWRQ FRPPXQLWLHV LQ WHPSHUDWHODNHVLQ2QWDULR&DQDGD ,QW5HYXHJHV+\GURELRO 3LFN )5 DQG %pUXEp & 'LHO F\FOHV LQ WKH IUHTXHQF\ RI GLYLGLQJ FHOOV RI IUHVKZDWHU SLFRF\DQREDFWHULD - 3ODQNWRQ 5HV 3ODWW 7 7KH DQQXDO SURGXFWLRQ E\ SK\WRSODQNWRQ LQ 6W 0DUJDUHW V%D\1RYD6FRWLD -&RQV LQW([SORU0HU 3ODWW 7 DQG /L :.: 3KRWRV\QWKHWLF SLFRSODQNWRQ &DQ %XOO)LVK$TXDW6FL 3RUWHU .* 3HDUO + +RGVRQ 5 3DFH 0 3ULVFX - 5LHPDQQ % 6FDYLD ' DQG 6WRFNQHU - 0LFURELDO LQWHUDFWLRQV LQ ODNH IRRGZHEV ,Q&DUSHQWHU65HG &RPSOH[,QWHUDFWLRQVLQ/DNH &RPPXQLWLHVSS6SULQJHU9HUODJ1HZ
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âLPHN. %RENRYD- 0DFHN0 1HGRPD- DQG 3VHQQHU5 &LOLDWH JUD]LQJ RQ SLFRSODQNWRQ LQ D HXWURSKLF UHVHUYRLU GXULQJ WKHVXPPHUSK\WRSODQNWRQPD[LPXP DVWXG\DWWKHVSHFLHVDQG FRPPXQLW\OHYHO/LPQRO2FHDQRJU âLPHN . 0DFHN 0 3HUQWKDOHU- 6WUDVNUDERYi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¡QGHUJDDUG 0 3KRWRWURSKLF SLFRSODQNWRQ LQ WHPSHUDWH ODNHV VHDVRQDO DEXQGDQFH DQG LPSRUWDQFH DORQJ D WURSKLF JUDGLHQW ,QW5HYXHJHV +\GURELRO 6WDQLHU5< .XQLVZDZD5 0DQGHO5DQG&RKHQ%D]LUH* 3XULILFDWLRQ DQG SURSHUWLHV RI XQLFHOOXODU EOXHJUHHQ DOJDH 2UGHU&KURRFRFFDOHV %DFWHULRO5HY 6WRFNQHU -* /DNH IHUWLOL]DWLRQ WKH HQULFKPHQW F\FOH DQG ODNH VRFNH\H VDOPRQ Oncorhynchus nerka SURGXFWLRQ ,Q 6PLWK +' 0DUJROLV / DQG :RRG && HGV 6RFNH\H 6DOPRQ Oncorhynchus nerka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
6WRFNQHU -* DQG 0DF,VDDF (0 %ULWLVK &ROXPELD ODNH HQULFKPHQWSURJUDPPH WZR GHFDGHV RIKDELWDWHQKDQFHPHQWIRU VRFNH\HVDOPRQ 5HJ5LYHUV 6WRFNQHU-*DQG3RUWHU.* 0LFURELDO IRRG ZHEV LQ IUHVK ZDWHU SODQNWRQLF HFRV\VWHPV &DUSHQWHU 65 HG &RPSOH[ ,QWHUDFWLRQV LQ /DNH &RPPXQLWLHV SS 6SULQJHU9HUODJ 1HZ
-6WRFNQHU&&DOOLHULDQG *&URQEHUJ 9|U|V / *XO\DV 3 DQG 1HPHWK - 2FFXUUHQFH G\QDPLFV DQG SURGXFWLRQ RISLFRSODQNWRQ LQ +XQJDULDQ VKDOORZ ODNHV ,QW 5HYXH JHV +\GURELRO 9UHGH . 5HJXODWLRQ RI EDFWHULRSODQNWRQ SURGXFWLRQ DQG ELRPDVV LQ DQ ROLJRWURSKLF FOHDUZDWHU ODNH WKH LPSRUWDQFH RI WKHSK\WRSODQNWRQFRPPXQLW\-3ODQNWRQ5HV :DQEHUJ6$6HOPHU-6 DQG*XVWDYVRQ. (IIHFWV RI89% UDGLDWLRQ RQ ELRPDVV DQG FRPSRVLWLRQ LQ PDULQH SK\WRSODQNWRQ FRPPXQLWLHV 6FHQWLD0DULQD :DQJ.6 DQG&KDL7- 5HGXFWLRQLQRPHJDIDWW\DFLGVE\ 89%LUUDGLDWLRQLQPLFURDOJDH-$SSO3K\FRO :DWHUEXU\ -% :DWVRQ 6: *XLOODUG 55 DQG %UDQG /( :LGHVSUHDG RFFXUUHQFH RI D XQLFHOOXODU PDULQH SODQNWRQLF F\DQREDFWHULXP 1DWXUH -% :DWVRQ 6: 9DORLV FW DQG )UDQNV '* %LRORJLFDO DQG HFRORJLFDO FKDUDFWHULVDWLRQ RI WKH PDULQH XQLFHOOXODU F\DQREDFWHULXP Synechococcus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
&KDSWHU 3LFRSODQNWRQ DQGRWKHUQRQEORRP IRUPHUV VSHFWURVFRS\ HSLIOXRUHVFHQFH PLFURVFRS\ DQG IORZ F\WRPHWU\ /LPQRO2FHDQRJU :\PDQ 0 DQG )D\ 3 8QGHUZDWHU OLJKW FOLPDWH DQG WKH JURZWK DQG SLJPHQWDWLRQ RI SODQNWRQLF EOXHJUHHQ DOJDH &\DQREDFWHULD , WKH LQIOXHQFH RI OLJKW TXDOLW\ 3URF 5 62& /RQG =DUHW0DQG6XIIHP./ 9HUWLFDOPLJUDWLRQLQ]RRSODQNWRQ DV DSUHGDWRUDYRLGDQFHPHFKDQLVP /LPQRO2FHDQRJU
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
6XPPDU\ &\DQREDFWHULD DUH DQ LPSRUWDQW FRPSRQHQW RI PDQ\ VRLOV LQFOXGLQJ WKH VXUIDFH FUXVWV WKDW VRPHWLPHV FRYHU H[WHQVLYHDUHDVLQ VHPLDULGUHJLRQV DQG PLQH VSRLO ZDVWHV 7KH\ DUHDOVR DEXQGDQWLQ PDQ\ DUHDV ZKLFK DUH ZHW RU VXEPHUJHG IRU SDUW RI WKH \HDU HVSHFLDOO\ ULFHILHOGV 0RVW VRLOV IRUPV KDYH VKHDWKV RU PXFLODJH DQG WKLV SRO\VDFFKDULGH KDV LPSRUWDQW HIIHFWV RQ WKH VRLO PRVWO\ EHQHILFLDO VXFK DV LPSURYHG VRLO VWUXFWXUH EXW VRPHWLPHV DGYHUVH ZKHUH DGHQVH VXUIDFHOD\HU LPSHGHV GUDLQDJH 1LWURJHQIL[LQJ VSHFLHV RIWHQ FRQVWLWXWH KDOI RU PRUH RI WKH VSHFLHVSUHVHQWLQ VRLOV QRW HQULFKHG ZLWK QLWURJHQRXV IHUWLOL]HU DQG WKHVH FDQ FRQWULEXWH FRPELQHG QLWURJHQ LQ VHYHUDO ZD\V WRDGMDFHQWYDVFXODU SODQWV $WWHPSWV WRHQKDQFHFURS\LHOG E\ DGGLQJ F\DQREDFWHULD WR VRLOVKDYHPRVWO\ IRFXVVHG RQ SDGG\ULFH $OWKRXJK PDQ\ VWXGLHV KDYH UHSRUWHG SRVLWLYH HIIHFWV RI VXFK DOJDOL]DWLRQ WKH QXPEHU RI ORFDWLRQV ZKHUH LW KDV EHHQ DGRSWHG DV URXWLQH SUDFWLFH DSSHDU WR EH IHZ LQ FRQWUDVW WR WKH UHODWLYHO\ ZLGHVSUHDG XVH RI Azolla ZLWK ULFH FXOWXUH $OJDOL]DWLRQ LV PRVW VXFFHVVIXO ZKHUH ORFDO VSHFLHV DUH XVHG WR SUHSDUH WKH LQRFXOXP EXW WKHUH LV FRQVLGHUDEOH VFRSH IRU RWKHU LPSURYHPHQWV ,W LV LPSRUWDQW WR REWDLQ D PXFK PRUH GHWDLOHG XQGHUVWDQGLQJ RI F\DQREDFWHULDO SRSXODWLRQ G\QDPLFV RYHUWKH ZKROH DQQXDO F\FOH LQ DJULFXOWXUDO V\VWHPV ZKHUH ULFH LV JURZQ IRU RQO\SDUWRI WKH\HDU B A Whitton and M Potts (eds). The Ecology of Cyanobacteria. Kluwer Academic Publishers Printed in the Netherlands
%$:KLWWRQ
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
,, 6RLOV $ 7D[RQRPLF&RPSRVLWLRQ $ GHWDLOHG UHYLHZ RI VRLO DOJDH E\ 0HWWLQJ JLYHV PDQ\ UHIHUHQFHV WR F\DQREDFWHULD LQFOXGLQJ D OLVW RI WKRVH JHQHUD ZLWK UHFRUGV IRU VRLO VSHFLHV %RWK XQLFHOOXODU DQG ILODPHQWRXV IRUPV DUH UHSUHVHQWHG EXW WKH ELRPDVV XVXDOO\ FRQVLVWV ODUJHO\ RI ILODPHQWRXV IRUPV ([DPSOHV RI VWXGLHV WKDW FRQWDLQ GHVFULSWLRQV DQG ILJXUHV RI VRLO VSHFLHV DUH JLYHQLQ7DEOH $QDQG V WD[RQRPLF DFFRXQW RI ULFHILHOG VSHFLHV LQ VRXWK ,QGLD SURYLGHV D SDUWLFXODUO\ XVHIXO JXLGH WR VSHFLHV OLNHO\ WREH IRXQG LQ ULFHILHOGV HYHU\ZKHUH
% (QYLURQPHQWDO)DFWRUV /LJKW &\DQREDFWHULDJURZLQJRQWKHVRLOVXUIDFHRIWHQVKRZ GDUN FRORUDWLRQV EOXHEODFN EURZQ UHGEURZQ RU UHG 7KH GDUNHU FRORXUV XVXDOO\ UHVXOW IURP WKH SUHVHQFH RI EURZQ VKHDWKV VXUURXQGLQJ WKH W\SLFDO SKRWRV\QWKHWLFWULFKRPHDQGWKLVFRORXULVPXFKPRUH SURQRXQFHG LQ SRSXODWLRQV LQ RSHQ WKDQ LQ VKDGHG SRVLWLRQV 7KH SLJPHQW UHSRQVLEOH IRU WKH EURZQ FRORXUDWLRQ VF\WRQHPLQ DEVRUEV VWURQJO\ LQWKH QHDU XOWUDYLROHWUHJLRQRIWKH VSHFWUXPF QP in vivo: *DUFLD3LFKHO DQG &DVWHQKRO] DQG WKH HYLGHQFH VWURQJO\ VXJJHVWV WKDW VF\WRQHPLQ SURGXFWLRQLV DQDGDSWLYHVWUDWHJ\IRUSKRWRSURWHFWLRQ DJDLQVW VKRUWZDYHOHQJWK VRODU LUUDGLDWLRQ &KDSWHU
&KDSWHU6RLOVDQG 5LFH)LHOGV
Table 1. 6RPH H[DPSOHV RI VWXGLHV ZLWK GHWDLOHG WD[RQRPLF DQGRU IORULVWLF LQIRUPDWLRQ RQ VRLO F\DQREDFWHULD 6HH DOVR &KDSWHU 7DEOH
Region
Subject
Reference
$UJHQWLQD $XVWUDOLD %DQJODGHVK &]HFK5HSXEOLF 5XVVLD *UHHFH ,QGLD
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
GH+DOSHULQHWDO (OGULGJH *UHHQH .KDQHWDO 'HVHUWRYi (FRQRPRXHWDO 3UDVDG 6ULYDVWDYD *XSWD 7LZDUL 7LZDUL 3DQGH\ -KDHWDO $QDQG +RSSHU $QDQG 6LQJKHWDOD 6LQJKHWDO E $O.DLVL *UXLD )RUHVW %ULVWRO $QGHUVRQ 5XVKIRUWK $VKOH\HWDO -RKDQVHQ
,UDT 5RPDQLD 5XVVLDDQGRWKHUIRUPHU8665 8. 86$8WDK
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icrocoleus vaginatus, Nostoc DQG Scytonema, ZKHUH VWURQJ DWWHQXDWLRQ PHDQW WKDW WKH HXSKRWLF ]RQH RI UHFHQWO\ ZHWWHG FUXVWV ZDV RQO\ DERXW PPEHORZ WKH VXUIDFH RI WKH FUXVW ,QIURVW VRUWHG SRO\JRQV RQ 6LJQ\ ,VODQG 6RXWK 2UNQH\ ,VODQGV DODUJH SURSRUWLRQ RI WKH PLFURIORUD RFFXUUHG LQ WKH ]RQH PPEHORZWKHVXUIDFHDQG IHZ DOJDH
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alothrix marchica ZDV WKH RQO\ RQH RI IRXU 6SDQLVK ULFHILHOG VRLO LVRODWHVFDSDEOHRIGRLQJVR3URVSHULHWDO 7KH ILODPHQWRXV IRUPV DUH SUREDEO\ DOO FDSDEOH RI PRYHPHQWXQGHU VRPH FRQGLWLRQV DQG SKRWRWD[LV KDV EHHQGHPRQVWUDWHGLQPDQ\F\DQREDFWHULD LQFOXGLQJ VWUDLQVLVRODWHGIURPVRLO&DVWHQKRO] 6SHFLHV GLIIHU KRZHYHU LQ WKH H[WHQW WR ZKLFK ILODPHQWV
H[KLELW PRWLOLW\LQQDWXUH $FWLYHO\ JURZLQJILODPHQWV RI Phormidium autumnale DQG VRPH RWKHU 2VFLOODWRULDFHDHDUHDSSDUHQWO\PRWLOHIRUPXFK RI WKH WLPH ZKHUHDV PRWLOLW\ LQ PDQ\ W\SLFDOO\ KHWHURF\VWRXV IRUPV HJ Nostoc DQG Calothrix LV UHVWULFWHG WR KRUPRJRQLD ZKLFK ODFN KHWHURF\VWV DQG DUH IRUPHGXQGHUVSHFLILFFRQGLWLRQV VXFK DVDGGLWLRQ RI SKRVSKDWH WR 3OLPLWHG ILODPHQWV :KLWWRQ 7KH DELOLW\ RI VRPH VRLO F\DQREDFWHULD WR PRYH SHUPLWV WKHP WR EH YLHZHG LQ D VHPLQDWXUDO IRUPE\ SODFLQJ FRYHUVOLSV LQ FORVH FRQWDFW ZLWK WKH VXUIDFH RI PRLVWHQHG VRLO DQGWKHQLQFXEDWLQJ LQ WKHOLJKWHJ %URDG\ 'DYH\ 0RWLOH RUJDQLVPV IUHTXHQWO\ DWWDFKWKHPVHOYHVWRWKHFRYHUVOLS ZLWKLQ D IHZ KRXUV ZKLOH QHZ FRORQLHV RI WKHVH DQG QRQ PRWLOH IRUPV PD\ GHYHORS ZLWKLQ D ZHHN 0RVW VSHFLHV RFFXUULQJ EHORZ WKH VRLO VXUIDFH DUH SUREDEO\ DWWDFKHG WR VRLO SDUWLFOHV 'DYH\ HW DO LQYHVWLJDWHG WKH LQIOXHQFH RI PRUSKRORJ\ PXFLODJH SURGXFWLRQ DQG VRLO WH[WXUH DV IDFWRUV LQIOXHQFLQJ DWWDFKPHQW RI F\DQREDFWHULD DQG HXNDU\RWLF DOJDH LVRODWHG IURP $QWDUFWLF IHOOILHOG VRLOV Phormidium autumnale VKRZHG WKH KLJKHVW DWWDFKPHQW RI DQ\ WD[RQ XQGHU VLPXODWHG IORZ FRQGLWLRQV LQFRQWUDVWWRWKHHXNDU\RWLFDOJDZygnema VS Pseudanabaena catenata VKRZHG JUHDWHU UHWHQWLRQ WRWKHVXEVWUDWXPZLWK WKHOHDVWUXJRVLW\
:DWHUDQG'HVLFFDWLRQ $LUGULHG WHUUHVWULDO DQG VRLO F\DQREDFWHULD FDQ VXUYLYH SURORQJHG GU\ SHULRGV DV KDV EHHQ VKRZQ LQ PDQ\ VWXGLHV 6RPH RI WKHVH KDYH EHHQ EDVHG RQ PDWHULDO VWRUHG GHOLEHUDWHO\ VXFK DV 7UDLQRU V XVH RI \U ROG PDWHULDO EXW PRVW KDYH EHHQ PDGH RQ VDPSOHV NHSW IRU RWKHU SXUSRVHV :LGHO\ TXRWHG VDPSOHV DUH DQ \U UHFRUG IRU Nostoc commune /LSPDQ DQG WKH JHQHUD RI F\DQREDFWHULD ZKLFK JUHZ IURP VDPSOHV RI SDUWLDOO\ DLUGULHG VRLOV IURP(QJODQG NHSW LQ VHDOHG ERWWOHV IRU \U%ULVWRO 3DUNHU HW DO UHYLHZHG ROG OLWHUDWXUH DQG FDUULHG RXWIXUWKHU VWXGLHV RQ PDWHULDOV IURP WKH 0LVVRXUL %RWDQLFDO *DUGHQ 6RPH RI WKH ROGHU VWXGLHVZHUHQRW ULJRURXVHQRXJKWRUXOHRXWWKH SRVVLELOLW\ RI DHULDO FRQWDPLQDQWV EXW LW LV FOHDU WKDW VRPH DNLQHWH DQG QRQDNLQHWHIRUPLQJ F\DQREDFWHULD FDQ VXUYLYH IRU YHU\ ORQJ SHULRGV 7KH PRVW IUHTXHQWO\ PHQWLRQHG JHQHUD DUHAnabaena, Aulosira, Cylindrospermum, Fischerella, Lyngbya, Nostoc, Plectonema DQG Stigonema. $ QXPEHU RI JUHHQ DOJDH DQG D IHZ GLDWRPV DUH DOVR PHQWLRQHG VR WKLV SURSHUW\ LV QRWXQLTXHWR F\DQREDFWHULD
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ynechococcus VWUDLQ VXUYLYHG IRUDW OHDVW RQH \HDU 0RVW VRLO F\DQREDFWHULD VXUYLYH SHULRGV RI GHVLFFDWLRQ ZLWK OLWWOH REYLRXV PRUSKRORJLFDO FKDQJH RWKHU WKDQ WKDW UHVXOWLQJ IURP ORVV RI ZDWHU 7KRVH JHQHUD ZKLFK IRUP DNLQHWHV HJ Anabaena GR VR ODUJHO\ DV D UHVSRQVH WR QXWULHQW RU RWKHU OLPLWDWLRQ UDWKHU WKDQ GHVLFFDWLRQ :KLWWRQ 3K\VLRORJLFDO IHDWXUHV RI GHVLFFDWLRQWROHUDQW F\DQREDFWHULD KDYH EHHQ UHSRUWHG LQ D QXPEHW RI VWXGLHV )RU LQVWDQFH WZR GURXJKWUHVLVWDQW VWUDLQV DFFXPXODWHG VXJDUV WR KLJK FRQFHQWUDWLRQV ZKHQ PDWULF ZDWHU VWUHVV ZDV DSSOLHG ZKHUHDV WZR VWUDLQV QRW VKRZLQJ GURXJKW UHVLVWDQFH GLG QRW GR VR +HUVKNRYLW] HW DO 7KH PRVW GHWDLOHG VWXGLHV KDYHEHHQGRQHRQ Nostoc &KDSWHU $OWKRXJK PDQ\ VRLO DQG GHVHUW F\DQREDFWHULD DUH KLJKO\ WROHUDQW RI GHVLFFDWLRQ WKH\ UHTXLUH WR EH UHZHWWHG WKRURXJKO\ IRU IXOO PHWDEROLF SURFHVVHV WR UHVXPH $ VWXG\ 3DOPHU DQG )ULHGPDQQ RI FU\SWRHQGROLWKLF FRPPXQLWLHV LQ GHVHUWV VKRZHG WKDW F\DQREDFWHULD SKRWRV\QWKHVL]H RQO\ DW KLJK PDWULF ZDWHU SRWHQWLDOV ! 03D UHODWLYH KXPLGLW\ DW & UHODWLYH WR JUHHQ DOJDOFRQWDLQLQJ OLFKHQV ZKLFK EHJDQ WR SKRWRV\QWKHVL]H DW D PXFK ORZHU SRWHQWLDO 03D UHODWLYH KXPLGLW\ DW & 7KH UHZHWWLQJ RI GULHG Nostoc commune OHDGV WR SK\VLRORJLFDO UHVSRQVHV LQ WKH VHTXHQFH UHVSLUDWLRQ SKRWRV\QWKHVLV DQG ODVWO\ QLWURJHQ IL[DWLRQ 6FKHUHU HW DO $ QRQFRORQLDOFXOWXUHRI N. commune, DIWHU VWRUDJH DW 03D VKRZHG LQFUHDVHG
&KDSWHU6RLOVDQG 5LFH)LHOGV QLWURJHQDVH DFWLYLW\ DQG VL]H RI LQWUDFHOOXODU $73 SRRO ZKHQ UHZHWWHG WKH XSVKLIW LQ QLWURJHQDVH DFWLYLW\ ZDV SUHFHGHG E\ D ODJ 3RWWV DQG %RZPDQ
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icrocoleus DSSHDUVWREHWKHPRVWZLGHVSUHDG F\DQREDFWHULXP RQ VDOLQHVXUIDFHVWKURXJKRXWWKHZRUOG :KLWWRQ ZKHUH LW FDQ SOD\ DQLPSRUWDQWSDUW LQ VWDELOL]DWLRQ RI WKH XQGHUO\LQJ VRLO $QGHUVRQ DQG 5XVKIRUWK %XWWDUV HW DO KDYH GHYHORSHG D PHWKRG IRU LQRFXODWLQJ VWHULOL]HG VRLO FUXVWV ZKLFK LQYROYHV WKH HVFDSH RI Microcoleus IURP DOJLQDWH EHDGV DQG LWV VXEVHTXHQW JURZWK RQWKH VRLO ,I WKLV PHWKRG FDQ EH DSSOLHG RQ D ODUJH VFDOH LW PLJKW EH SRVVLEOH WR DFFHOHUDWH WKH UDWH RI UHFRYHU\ RI KLJKO\ GLVWXUEHG VRLOV DQG DGG WR WKH PRUH FRQYHQWLRQDO PHWKRGV VXJJHVWHG E\ 6LQJK
S+ $V PHQWLRQHG DERYH F\DQREDFWHULD DUH LQIUHTXHQW EHORZ S+ LQ PRVW WHPSHUDWH VRLOV WKH\ ZHUH IRU LQVWDQFH HQWLUHO\ DEVHQW EHORZ S+ LQ VHYHUDO VDPSOHV IURP ,UHODQG 'RROH\ DQG +RXJKWRQ +RZHYHU WKHUH DUH VLWXDWLRQV ZKHUH WKH\ RFFXU DW VXEVWDQWLDOO\ ORZHUS+ YDOXHV 7KHVH LQFOXGHWURSLFDO VRLOV 0RRUH DQG WKH HGJHV RI VPDOO SRROV 6HFWLRQ ,,& 9DOXHV GRZQ WR S+ RU HYHQ VOLJKWO\ ORZHU KDYH EHHQ UHFRUGHG LQ ERWK FDVHV DXWKRU V REVHUYDWLRQV ,W VHHPV SUREDEOH WKDW WKH VRLOV DW S+ YDOXHV MXVW DERYH ZKLFK LQFOXGH F\DQREDFWHULD DUH DOO RQHV ZLWK ZHDN EXIIHULQJ FDSDFLWL\ 7KHUH DUH DOVR VFDWWHUHG UHFRUGV RI F\DQREDFWHULD LQ KLJKO\ DFLGLF HQYLURQPHQWV EXW LQ WKRVH FDVHV ZKHUH WKH VLWHV KDYH EHHQ UHYLVLWHG WKH RUJDQLVPV KDYH SURYHG WR EH Cyanidium RU SHUKDSV RQH RI WKH RWKHU VPDOO SK\FRELOLQFRQWDLQLQJ
HXNDU\RWHV 1HYHUWKHOHVV WKH SRVVLEOH RFFXUUHQFH RI F\DQREDFWHULD DW KLJKO\ DFLGLF VLWHV VWLOO UHTXLUHV WKRURXJK UHHYDOXDWLRQ &\DQREDFWHULD PD\ DOVR LQIOXHQFH WKH S+ GXH WR WKHLU PHWDEROLF DFWLYLW\ ,Q DGGLWLRQ WR WKH PDUNHG FKDQJHV ZKLFK FDQ WDNH SODFH LQ SDGG\ ILHOGV DQG WHPSRUDU\ SRROV RYHUO\LQJ VRLO WKH\ FDQ DOVR RFFXU ZLWKLQ VXUIDFH FUXVWV /RFDOL]HG YDOXHV RI XQLWV DERYH WKH VRLO S+ KDYH EHHQ IRXQG LQ GHVHUW FUXVWV DV D UHVXOW RI SKRWRV\QWKHWLF DFWLYLW\ *DUFLD 3LFKHODQG%HOQDS
+HUELFLGHVDQG3HVWLFLGHV 7KH LQIOXHQFH RI KHUELFLGHV DQG SHVWLFLGHV RQ F\DQREDFWHULD KDV EHHQ LQYHVWLJDWHG LQ PDQ\ VWXGLHV 3DGK\ 5RJHU E 9DLVKDPSD\DQ HW DO WKRXJKPRVWKDYH EHHQ UHVWULFWHG WR ODERUDWRU\ FXOWXUHV &\DQREDFWHULDDUHLQ JHQHUDOTXLWH VHQVLWLYH WR KHUELFLGHV EHFDXVH WKH\ VKDUH PDQ\ RI WKH SK\VLRORJLFDO IHDWXUHV RI KLJKHU SODQWV /HJDQpV DQG )HUQDQGH]9DOLHQWH ZKLFK IRUP WKH VLWH RI KHUELFLGH DFWLRQ 7KH SUHHPHUJHQFH KHUELFLGH EHQWKLRFDUE ZDV KLJKO\ WR[LFWR VWUDLQV RINostoc DQG Anabaena =DLWVHYD +RZHYHU 1IL[LQJ F\DQREDFWHULD DUH PRVWO\ UHODWLYHO\ WROHUDQW WR ' GLFKORURSKHQR[\ DFHWLF DFLG DWOHDVWXQGHU ILHOG FRQGLWLRQV 6WUDWWRQ /HJDQpV DQG )HUQDQGH] 9DOLHQWH 'LIIHUHQFHV KDYH EHHQ IRXQG EHWZHHQ WKH WROHUDQFH WR KHUELFLGHV RI F\DQREDFWHULD DQG WKDW RI RWKHU RUJDQLVPV Anabaena, Nostoc DQG Oscillatoria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
%$ :KLWWRQ ILQG HYLGHQFH IRU D nif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
&KDSWHU 6RLOV DQG 5LFH)LHOGV
Fig. 1 . &RORQLHV RIF\DQREDFWHULDAnabaena, Cylindrospermum, Lyngbya, Porphyrosiphon notarisii,Scytonema mirabile DQG Tolypothrix byssoidea GHYHORSLQJRQPRLVWVRLOLQDGHHSZDWHUULFHILHOGDW6RQDUJDRQ %DQJODGHVK VHYHUDOZHHNVEHIRUHWKHDUULYDORIWKHIORRGZDWHU 9LVXDOO\REYLRXVFRPPXQLWLHVZLOOFRYHUDERXWKDOIWKHVXUIDFHRIWKHILHOGE\WKHWLPHWKHIORRGDUULYHV :LGWKRIILJXUHLVFP
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hormidium autumnale KDG D YHU\ PDUNHG FORJJLQJFDSDFLW\ 7KLVZDV DVFULEHGWR WKH UHOHDVH RI FRSLRXV PXFXV WZR RWKHU F\DQREDFWHULD ZKLFK IRUPHG OHVVSRO\VDFFKDULGHKDG D PXFK ORZHU FORJJLQJ FDSDFLW\
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nabaena ZLWK URRW
&KDSWHU6RLOVDQG 5LFH)LHOGV KDLUV RI ZKHDW *DQWDU HW DO D E IRXQG WKDW WKHUH ZHUH PXFK WLJKWHU DVVRFLDWLRQV ZLWK VRPH Nostoc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estuca octojlora JURZLQJ RQ VRLOV
KHDYLO\ HQFUXVWHG ZLWK F\DQREDFWHULD DQG F\DQROLFKHQVWKDQLQSODQWVRQWKHVDPHVRLOZKHUHWKH FUXVW KDG EHHQ GHVWUR\HG ,Q WKH FDVH RI Mentzelia multiflora, 1 0JDQG)HZHUHSUHVHQWDWVLJQLILFDQWO\ KLJKHU FRQFHQWUDWLRQV EXW 3 ZDV SUHVHQW DW VLJQLILFDQWO\ ORZHU FRQFHQWUDWLRQV 7KH DXWKRUV VXJJHVWHG D YDULHW\ RI SRVVLEOH UHDVRQV IRU WKHVH GLIIHUHQFHV $V GHVFULEHG DERYH F\DQREDFWHULDO 1 IL[DWLRQ LV OLNHO\ WR LQFUHDVH WKH DYDLODELOLW\ RI1 WR DGMDFHQW SODQWV 7KH UHGXFHG XSWDNH RI 3 E\ Mentzelia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nabaena VWLPXODWLQJ JURZWK RQO\ RQH UHPDLQHGHIIHFWLYHDIWHULWKDGEHHQPDGHD[HQLF
,,, 6XEDHULDO +DELWDWV 'HYHORSPHQWRI6XEDHULDO&RPPXQLWLHV 6XEDHULDO FRPPXQLWLHV RI F\DQREDFWHULD DQG DOJDH ZHUHILUVWGHVFULEHGLQGHWDLOIURP6UL/DQND&H\ORQ E\ )ULWVFK DF DQG KLV JHQHUDO DFFRXQW RIVLWHVLQDQXPEHURIFRXQWULHVLVVWLOOZRUWKUHDGLQJ +HFRQFOXGHGWKDWWKHILUVWFRORQL]HUV RIURFNVXUIDFHV DUH FRORQLDO &KURRFRFFDOHV Gloeocapsa, Gloeothece DQG Aphanocapsa RIWHQ DFFRPSDQLHG E\ Nostoc. 7KHVWUDWXPIRUPHGE\WKHVHJHQHUDLV VRRQHURUODWHU FRORQL]HG E\ ILODPHQWRXV IRUPV Lyngbya, Scytonema, Stigonema DQG Hapalosiphon XVXDOO\ SRVVHVVLQJ VKHDWKV ZKLFK DUH ILUPHU DQG OHVV
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olypothrix, Anabaena, Symploca DQG Lyngbya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olypothrix ZKLFK ZDV REVHUYHG GLUHFWO\ DPRQJ YROFDQLF DVK 7KLV VWXG\ ZRXOG EH ZHOO ZRUWK UHSHDWLQJ WKLV WLPH LQFOXGLQJ FRQWUROV WR HVWLPDWHWKH FRQWULEXWLRQ IURP WKHDLU VSRUDDQG XVLQJ D PXFK OHVVQXWULHQWULFK PHGLXP
'LVSHUVDOLQ WKH$LU 6LQFH F\DQREDFWHULD DUH IUHTXHQW FRPSRQHQWV RI WKH DLU VSRUD 6FKOLFKWLQJ 7LEHUJ HW DO WKH\ DUHOLNHO\ RIWHQWR EH UHFRYHUHGIURPVRLO HQULFKPHQW FXOWXUHV %RWK WKH VSHFLHV RI
%$:KLWWRQ F\DQREDFWHULD Phormidium ambiguum, 3 laminosum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
&UXVW&RPPXQLWLHVLQ 6HPLDULG5HJLRQV 7KH PRVW GHWDLOHG DFFRXQWV RI VXEDHULDO FRPPXQLWLHV ZLWK DEXQGDQW F\DQREDFWHULD FRPH QRW IURP WKH ZHW WURSLFV EXW IURP DULG UHJLRQV VXFK DV WKH YROFDQLF VRLOV LQ 6: 86$ ZKHUH Microcoleus DQG Porphyrosiphon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ostoc commune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icrocoleus vaginatus RQ VDQG\ VRLOV ZDV PRUH VXVFHSWLEOH WKDQ FUXVWV ZLWK WKH OLFKHQ Collema tenax. $OWKRXJK GLVWXUEDQFH PD\ DFWXDOO\ OHDG WR DQ LQFUHDVH LQ WKH DEXQGDQFH RI IUHHOLYLQJ F\DQREDFWHULD DV VKRZQ IRU VLWHV RQ WKH &RORUDGR 3ODWHDX (YDQV DQG %HOQDS LW OHDGV WR D GHFUHDVH LQ WKH DEXQGDQFH RI OLFKHQV VRPH RI ZKLFK
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
+HDY\0HWDODQG2WKHU&RQWDPLQDWLRQ &\DQREDFWHULD DUH VRPHWLPHV DEXQGDQW RQ DQG QHDU WKH VXUIDFH RI VRLOV FRQWDPLQDWHG E\ YDULRXV SROOXWDQWV RU QDWXUDOO\ HQULFKHG E\ KHDY\ PHWDOV 0RVW UHFRUGV IRU KHDY\ PHWDOV DUH IRU ZDVWHV DVVRFLDWHG ZLWK OHDG]LQF PLQLQJ :KLWWRQ :KLWWRQ HW DO EXW WKHUH DUH DOVR UHFRUGV IRU VRLOV ULFK LQ FRSSHU DQG RWKHU PHWDOV (UQVW 5DQD HW DO 7KH GRPLQDQWV DUHXVXDOO\ QDUURZ IRUPV RI 2VFLOODWRULDFHDH VXFK DV Plectonema DQG Schizothrix DQG WHQG WR RFFXU PDLQO\ DW WKH VXUIDFH 3ODWH G 2OGHU FRPPXQLWLHV LQ =QULFK DUHDV GHYHORS D PL[WXUH RI Plectonema DQG SURWRQHPD RI VSHFLHV RI WKH PRVV Dicranella DQG WKLV LQ WXUQ JLYHV ULVH WR LQWHUPLQJOHG Plectonema DQG OHDI\ Dicranella 3ODWH I DQG VRPHWLPHV DOVR OLFKHQV HVSHFLDOO\ Collema VSS 7KH \RXQJHU VWDJHV RI VXFK FRPPXQLWLHV KDYH PDQ\ VLPLODULWLHV ZLWK WKH FU\SWRELRWLF FUXVWV RI VHPLDULG UHJLRQV VHH DERYH DOWKRXJK WKH\ FDQ GHYHORS XQGHU FRQGLWLRQV RI KLJK UDLQIDOO 2OG VKHDWKV RI Plectonema DQG GHFD\LQJ SDUWV RI WKH PRVV DSSHDU WR SOD\ WKH PDLQ UROH LQ GHYHORSLQJ D WUXH VRLO RQ PDQ\ VSRLO KHDSV WKRXJK VSHFLHV RI Collema DQG Peltigera ZLWK F\DQREDFWHULDO V\PELRQWV VRPHWLPHV RFFXU DW WKHVH VLWHV DQG FRQWULEXWH IL[HG QLWURJHQ WR WKH HFRV\VWHP DXWKRU V XQSXEOLVKHG GDWD 7KHUH DUH PDQ\ UHFRUGV RI F\DQREDFWHULD WROHUDWLQJ FRQWDPLQDWLRQ E\ RLO RU SHUKDSV HYHQ EHLQJ IDYRXUHG E\ LWVSUHVHQFHWKRXJK WKHVHGHDOPRVWO\ ZLWK VKDOORZ ZDWHUV UDWKHU WKDQ VRLO 6HYHUDOVWXGLHVKDYHQRWHG WKH RFFXUUHQFH RI WKH QLWURJHQIL[HU Nostoc commune LQ RU RQ VRLOVQDWXUDOO\FRQWDPLQDWHGE\ RLO WDUVDQGV RI $OEHUWD 0 +LFNPDQ DQG WKH DXWKRU XQSXEOLVKHG
GDWD RUSROOXWHG E\ SHWUROHXPSURGXFWV HJ $WODV HW DO 7KH H[WHQVLYH OLVW RI F\DQREDFWHULD JURZLQJ RQ FRDO WDU UHSRUWHG E\ %DUKDWH DQG 7DUDU GRHV QRW LQFOXGH DQ\ KHWHURF\VWRXV IRUPV EXW QR VWXGLHV ZHUH GRQH WR HVWDEOLVK ZKHWKHU DQ\ RIWKH VSHFLHV OLVWHG ZHUH QRQKHWHURF\VWRXV QLWURJHQIL[HUV
6SRUWV 7XUI 6RLO F\DQREDFWHULD LQ JROI FRXUVHV DQG RWKHU DPHQLW\ JUDVVODQGV PD\ UHDFK GHQVLWLHV VXIILFLHQW WR KDYH IDYRXUDEOH HIIHFWV RQ VRLO SURSHUWLHV EXW VXUIDFH JURZWKV FDQ DOVR EH H[WHQVLYH HQRXJK WR FDXVH PDQDJHPHQW SUREOHPV %DOGZLQ DQG :KLWWRQ 7ZR GLIIHUHQW W\SHV RI VLWXDWLRQ KDYH EHHQ UHFRJQL]HG 2QHLV ZKHUH WKHGHYHORSPHQW RI Nostoc commune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scillatoria VSS Nostoc VS DQG Desulfovibrio VSS LQ VDQG FROXPQV VKRZHG WKDW EODFNHQLQJ RFFXUUHG RQO\ ZKHQ F\DQREDFWHULD DQG Desulfovibrio ZHUH ERWK SUHVHQW %ODFNHQLQJ ZDV PRVW LQWHQVH LQ UHVSRQVH WR DGGHG VXOSKXU FKHODWHG LURQ RUDPL[WXUHRI WKH WZR
,9 5LFHILHOGV &\DQREDFWHULD ZHUH ILUVW UHSRUWHG WR EH DEXQGDQW LQ ULFHILHOGV E\ )ULWVFK D F DQG WKHLU LPSRUWDQFH LQ KHOSLQJ WR PDLQWDLQ ULFHILHOG IHUWLOLW\ GXH WR 1 IL[DWLRQ ZDV VXJJHVWHG E\ 'H &\DQREDFWHULD
%$:KLWWRQ DUH HVSHFLDOO\ HYLGHQW LQ ZHWODQG ULFHILHOGV ZKLFK VXSSO\ RI WKH ZRUOG UHTXLUHPHQWIRUULFH /DGKD DQG 5HGG\ 0DQ\ ULFHILHOG VRLOV QRW RQO\ FRQWDLQ D KLJK GHQVLW\ RI F\DQREDFWHULD EXW SRVVHVV YLVXDOO\ REYLRXV JURZWKV RI F\DQREDFWHULD DW RU IORDWLQJ DERYH WKH VXUIDFH DW OHDVW GXULQJ VRPH VHDVRQV 7\SLFDOO\ DERXW KDOI WKH F\DQREDFWHULDO JHQHUD SUHVHQW DUH KHWHURF\VWRXV Anabaena, Aulosira, Calothrix, Cylindrospermum, Fischerella, Gloeotrichia, Nostoc, Scytonema, Tolypothrix, Wollea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nabaena, Calothrix, Nostoc DOO UHVSRQGHG GLIIHUHQWO\ WR OHYHOV RI LUUDGLDWLRQ +LJKHU DEXQGDQFH RI Nostoc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cytonema mirabile ZHUH
&KDSWHU6RLOVDQG 5LFH)LHOGV VRPHWLPHV DEXQGDQW ERWK RQ WKH PRLVW VRLO DQG IORDWLQJ LQ WKH VDPHILHOG ZKHQ IORRGHG 0DQ\ RI WKH ULFHILHOG VSHFLHV IRUPDNLQHWHV VR LW ZRXOG EH XVHIXO WR NQRZ ZKDW IDFWRUV OHDG WR DNLQHWH IRUPDWLRQ LQ SDUWLFXODU VSHFLHV $ VWUDLQ RI Fischerella muscicola LVRODWHG IURP D ILHOG DW WKH &XWWDFN 5LFH 5HVHDUFK ,QVWLWXWH IRUPHG DNLQHWHV XQGHU 3 OLPLWDWLRQ 0LVKUD DV GR DERXW KDOI WKH VWUDLQV RI F\DQREDFWHULD LVRODWHG IURP GLYHUVH KDELWDWV :KLWWRQ +RZHYHU QRW DOO ULFHILHOG VSHFLHV IRUP DNLQHWHV LQ UHVSRQVH WR 3 OLPLWDWLRQ 5HGG\ $ VWXG\ :KLWWRQ RI VXUIDFH SKRVSKRPRQRHVWHUDVH DFWLYLW\ RI PDFURVFRSLFFRORQLHVLQSDGG\ILHOGV LQ WKH 3KLOLSSLQHV VKRZHG WKDW Aphanothece stagnina WHQGHG WR KDYH ORZ DFWLYLW\ ZKHUHDV Gloeotrichia DOZD\V VKRZHG KLJK DFWLYLW\ $PRQJ Nostoc, N. commune KDG KLJK DFWLYLW\ ZKHUHDV RWKHU VSHFLHV WHQGHG WR KDYH OHVV DFWLYLW\ $V SKRVSKRPRQRHVWHUDVH DFWLYLW\ LV HQKDQFHG LQ DOOWKHVH VSHFLHV XQGHU FRQGLWLRQV RI 3 OLPLWDWLRQ LW ZDV FRQFOXGHG WKDW Aphanothece W\SLFDOO\ JUHZ XQGHU 3ULFK FRQGLWLRQV ZKHUHDV Gloeotrichia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i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ulosira,
Aphanothece RU Gloeotrichia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olypothrix tenuis. 6WXGLHV RQ WKH XVH RI LQRFXOD KDYH EHHQ GLVFRQWLQXHG LQ -DSDQ EXW QXPHURXV VWXGLHV KDYH EHHQ PDGH HOVHZKHUH HVSHFLDOO\ LQ ,QGLD.DXVKLN DQG PRUHUHFHQWO\ LQ (J\SW HJ
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
%$:KLWWRQ IORRG FRQGLWLRQV E\ LPPRELOL]LQJ WKHP RQ SRO\XUHWKDQH IRDP .DQQDL\DQ HW DO 7KLV VWXG\LQFOXGHGD ILHOGH[SHULPHQWWKHUHVXOWVRI ZKLFK LQGLFDWHG WKDW IRDPLPPRELOL]HG Anabaena azollae H[FUHWHG FRQVLGHUDEOH DPRXQWV RI DPPRQLD LQWR WKH IORRG ZDWHU DQG WKDW WKLV OHG WR LQFUHDVHG ULFH JUDLQ DQG VWUDZ \LHOGV $OWKRXJK WKH VWUDLQ ZDV LVRODWHG IURP Azolla, HYLGHQFH ZDV QRW LQFOXGHG WR FRQILUP WKDW LW ZDV LQ IDFW WKH HQGRV\PELRQW Anabaena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zolla ZLWK LWV V\PELRWLF QLWURJHQIL[LQJ Anabaena &KDSWHU KDVEHHQ LQYHVWLJDWHG LQ PDQ\ VWXGLHV /XPSNLQ DQG 3OXFNQHWW :KLWWRQ DQG 5RJHU .DQQDL\DQ :KLWWRQ 7KH Azolla LV XVHG LQ D YDULHW\ RI ZD\V 2QHDSSURDFKLV
&KDSWHU6RLOVDQG 5LFH)LHOGV WRDGGDODUJHLQRFXOXPZLWKLUULJDWLRQZDWHUWRILHOGV DWDQHDUO\VWDJHLQWKHULFHFURSOHDYLQJWKHAzolla WR JURZIRUDIHZZHHNVIRU DIHZZHHNV WKHQSDUWLDOO\ GUDLQLQJ WKH ILHOGV DQG SORXJKLQJ WKHAzolla LQWR WKH VRLO $QRWKHU DSSURDFK LV WR JURZAzolla RQ LWV RZQ DQG WKHQFRPSRVWWKHSODQWV DQGSORXJK WKH GHFD\LQJ PDWHULDO LQWR WKH VRLO (LWKHU ZD\ PRVW RI WKH QLWURJHQIL[HGE\WKH V\PELRQWHYHQWXDOO\UHDFKHVWKH VRLO RU WKH IORRG ZDWHU DQG WKXV VRPH EHFRPHV DYDLODEOH IRU WKH ULFH .DQQDL\DQ 5RJHU D
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a PDNHV LW HDV\ WR GLVWLQJXLVK WKH VPDOOHU F\DQREDFWHULD IURP EDFWHULD &RPELQDWLRQV RIH[FLWDWLRQDQGVXSSUHVVLRQ ILOWHUV PD\ EH FKRVHQ WR GLVWLQJXLVK WKH DFFHVVRU\ SLJPHQWV SK\FRF\DQLQ DQG SK\FRHU\WKULQ :\QQ :LOOLDPV ZKLOHWKHIOXRURFKURPH$XUDPLQH2 FDQ EH XVHG WR GLIIHUHQWLDWH OLYLQJ IURP GHDG FHOOV +DZHVDQG'DYH\ UHSRUWHGWKDWYLDEOHFHOOVRI Phormidium IOXRUHVFHG\HOORZZKLOHQRQYLDEOHFHOOV UHWDLQHGRQO\WKHUHG IOXRUHVFHQFH &XOWXUHGPDWHULDO VWDLQHG PXFK EHWWHU WKDQ ILHOG PDWHULDO VXJJHVWLQJ WKDW WKLV PHWKRG UHTXLUHV IXUWKHU GHYHORSPHQW EHIRUH EHLQJSXWWRURXWLQHXVH 3ODWLQJ WHFKQLTXHV IRU FRXQWLQJ F\DQREDFWHULD GHSHQG RQ WKH VXLWDELOLW\ RI WKH FKRVHQ PHGLXP IRU JURZWK RIF\DQREDFWHULD 5H\QDXG DQG 5RJHU DQG WKH UHOLDELOLW\ RI WKH GLOXWLRQ PHWKRG XVHG $ JHQHUDO JXLGH RQ FKRLFH RI PHWKRGV IRU TXDQWLI\LQJ VRLO F\DQREDFWHULD LV JLYHQ E\ :KLWWRQ ,W VHHPV SUREDEOH WKDW PRVW VRLO F\DQREDFWHULD JURZ UHODWLYHO\ HDVLO\ LI SODWHG RQ VXLWDEOH QXWULHQW DJDU 7KH DEVHQFH RI F\DQREDFWHULD UHSRUWHG LQ WKH OLWHUDWXUH IRU VRLOV RI VHYHUDO UHJLRQV HJ SDUWV RI
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
%$:KLWWRQ
VRLOVUDQJHIURP J GU\ZHLJKWRI VRLO ZLWK D PHDQ RI [ $ IXUWKHU SUREOHP LQ LQWHUSUHWLQJ WKH GDWD FRQFHUQLQJ &)8V RQ DQ DJDU VXUIDFHUHVXOWV IURPWKH IDFW WKDW PDQ\ F\DQREDFWHULD UDSLGO\ XQGHUJR FHOO RU ILODPHQW GLYLVLRQ RQ WUDQVIHU WR QXWULHQWULFK PHGLD :KLWWRQ 7KH UHOHDVH RI PRWLOHKRUPRJRQLDLV IUHTXHQW ,QVRPHFDVHV HJ Calothrix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alothrix LQ EDWFK FXOWXUH
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
&KDSWHU 6RLOV DQG 5LFH)LHOGV PDQ\ VRLOV )LJ 1), TXDQWLWDWLYH GDWD RIWHQ QHHGV D KLJK QXPEHU RI UHSOLFDWHV VR ELRPDVV QHHGV WR EH TXDQWLILHG E\ D VLPSOH PHWKRG &KORURSK\OO a LV WKH XVXDOYDULDEOHFKRVHQIRUWKLV SXUSRVHVHHDERYH
& $VVD\V IRU )HUWLOLW\ DQG 7R[LFLW\ $ PHWKRGRORJ\ IRU XVLQJ DOJDH WR DVVHVV WKH QXWULHQW VWDWXV RI VRLOV ZDV GHYHORSHG E\ 7FKDQ 7FKDQ HW DO ORQJ EHIRUH D VLPLODU DSSURDFK ZDV XVHG IRU ZDWHU 7KLV PDGH XVH RI D PL[HG LQRFXOXP WKRXJK DSSDUHQWO\ PRVWO\ JUHHQ DOJDH +RZHYHU Nostoc commune ZDV VHOHFWHG E\ 3HGHUVRQ DQG 6KXEHUW IRU DVVD\LQJ WKH WR[LFLW\ RI VXEVWDQFHV OLNHO\ WR FRQWDPLQDWHVRLO 7KLVLQYROYHVJURZLQJWKH RUJDQLVP RQ PHPEUDQHILOWHUV VXSHULPSRVHG RQ WKH VXUIDFH RI QXWULHQW DJDU FRQWDLQLQJ YDULRXV FRQFHQWUDWLRQV RI WKH VXEVWDQFH XQGHU WHVW WKLV WHFKQLTXH PDLQWDLQV WKH DLUVROLG LQWHUIDFHLPSRUWDQW IRU PDQ\ VRLO VXUIDFHF\DQREDFWHULD 3UHVXPDEO\ WKH PHWKRG FRXOG EH PRGLILHG VXFK WKDW WKH PHPEUDQH ZLWK WKH Nostoc FRORQ\ LV SODFHG GLUHFWO\ RQ WKH VXUIDFH RI D VRLO VDPSOH
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a. 7KLV LV SDUWO\EHFDXVHRI WKHSRVVLEOHRU NQRZQ LPSRUWDQFH RI WKH FRQWULEXWLRQE\ F\DQREDFWHULDO QLWURJHQ IL[DWLRQ LQ PDQ\ HFRV\VWHPV EXW DOVR EHFDXVH RI WKH UHODWLYH
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p 0&= 3DOPD 50 DQG &RORPER . ([RSRO\VDFFKDULGH RI Nostoc muscorum F\DQREDFWHULD LQ WKH DJJUHJDWLRQ RI VRLO SDUWLFOHV - $SSO 3K\FRO GH +DOSHULQ '5 GH &DQR 06 GH 0XOp 0&= DQG GH &DLUH *= 'LD]RWURSKLF F\DQREDFWHULD IURP $UJHQWLQH SDGG\ ILHOGV 3K\WRQ 'HVLNDFKDU\ 79 &\DQRSK\WD ,QGLDQ &RXQFLO RI $JULFXOWXUDO5HVHDUFK 1HZ 'HOKL 'HVHUWRYi % 6RPH LQWHUHVWLQJ DOJDH IURP VRLO $UFK +\GURELRO 6XSSO GH :LQGHU % 3OXLV - GH 5HXV / DQG 0XU /5 &KDUDFWHUL]DWLRQ RI D F\DQREDFWHULDO DOJDO GXQH FUXVW LQ WKH FRDVWDO GXQHV LQ WKH 1HWKHUODQGV ,Q &RKHQ < DQG 5RVHQEHUJ ( HGV 0LFURELDO 0DWV 3K\VLRORJLFDO (FRORJ\ RI %HQWKLF
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ostoc F\DQREDFWHULD LQRFXODWLRQ RQ WKH VWUXFWXUH DQG VWDELOLW\ RI FOD\ VRLOV %LRO )HUWLO 6RLOV )HUQiQGH]9DOLHQWH( 4XHVDGD $ 3URVSHUL & 1LHYD 0 /HJDQpV ) DQG 8FKD $ 6KRUW DQG ORQJWHUP HIIHFWV RI DPPRQLXP RQ SKRWRGHSHQGHQW QLWURJHQ IL[DWLRQ LQ ZHWODQG ULFH ILHOGV RI 6SDLQ %LRO )HUWLO 6RLOV )RUHVW +6 7KH VRLO DOJDO FRPPXQLW\ ,, 6RYLHW VRLO VWXGLHV -3K\FRO )ULWVFK )( D 7KH VXEDHULDO DQG IUHVKZDWHU DOJDO IORUD RI WKH WURSLFV $QQDOV RI %RWDQ\ )ULWVFK )( E 7KH U{OH RI DOJDO JURZWK LQ WKH FRORQL]DWLRQ RI QHZ JURXQG DQG LQ WKH GHWHUPLQDWLRQRI VFHQHU\ *HRJ - )ULWVFK )( F $ JHQHUDO FRQVLGHUDWLRQ RI VXEDHULDO DQG IUHVKZDWHU DOJDO IORUD RI &H\ORQ $ FRQWULEXWLRQ WR WKH VWXG\ RI WURSLFDO DOJDO HFRORJ\ 3DUW , 6XEDHULDO DOJDH DQG DOJDH RI WKH LQODQG IUHVKZDWHUV 3URF 5R\ 6RF /RQGRQ 6HULHV % )ULWVFK )( 7KH WHUUHVWULDO DOJD - (FRO *DOORQ -5 +DVKHP 0$ DQG &KDSOLQ $( 1LWURJHQ IL[DWLRQ E\ Oscillatoria VSS XQGHU DXWRWURSKLF DQG SKRWRKHWHURWURSKLF FRQGLWLRQV - *HQ 0LFURELRO *DQWDU 0 .HUE\ 1: 5RZHOO 3 DQG 2EUHKW = D &RORQL]DWLRQ RI ZKHDW Triticum vulgare / E\ 1IL[LQJ F\DQREDFWHULD , $ VXUYH\ RI VRLO F\DQREDFWHULDO LVRODWHV IRUPLQJ DVVRFLDWLRQV ZLWK URRWV 1HZ 3K\WRO *DQWDU 0 .HUE\ 1: DQG 5RZHOO 3 E &RORQL]DWLRQ RI ZKHDW Triticum vulgare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ryza
251 sativa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a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alothrix ' 0LFURELRV ,VODP 05 DQG :KLWWRQ %$ E 3KRVSKRUXV FRQWHQW DQG SKRVSKDWDVH DFWLYLW\ RI WKH GHHSZDWHU ULFHILHOG F\DQREDFWHULXP EOXHJUHHQ DOJD Calothrix ' 0LFURELRV -HIIULHV '/ .ORSDWHN -0 /LQN 62 DQG %ROWRQ + -U $FHW\OHQH UHGXFWLRQ E\ FU\SWRJDPLF FUXVWV IURP D EODFNEUXVK FRPPXQLW\ DV UHODWHG WR UHVDWXUDWLRQ DQG GHK\GUDWLRQ 6RLO %LR %LRFKHP -KD 01 -KD 81 $KPDG 1 DQG 0DOOLN 0. &\DQREDFWHULDO IORUD RI ULFH ILHOG VRLOV RI 3XVD DQG LWV DGMRLQLQJ DUHDV 3K\NRV
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nabaena variabilis. $SSO 6RLO (FRO .DPXUX ) $OEUHFKW 6/$OOHQ /+ DQG 6KDQPXJDP .7 'U\ PDWWHU DQG QLWURJHQ DFFXPXODWLRQ LQ ULFH LQRFXODWHG ZLWK D QLWURJHQDVHGHUHSUHVVHG PXWDQW RI Anabaena variabilis. $JURQRP\ - .DQQDL\DQ 6 $]ROOD %LRIHUWLOL]HU 7HFKQRORJ\ IRU 5LFH &URS 7DPLO 1DGX $JULFXOWXUDO 8QLYHUVLW\ &RPEDWRUH ,QGLD .DQQDL\DQ S 1LWURJHQ FRQWULEXWLRQ E\ Azolla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zolla DV ELRIHUWLOL]HU IRU ULFH ,Q 6XEUDPDQLDQ * .DXVKLN %' DQG 9HQNDWDUDPDQ *6 HGV &\DQREDFWHULDO %LRWHFKQRORJ\ 6FLHQFH 3XEOLVKHUV ,QF (QILHOG 86$ .XODVRRUL\D 6$ 5RJHU 3$ %DUUDTXLR :/ DQG :DWDQDEH , %LRORJLFDO QLWURJHQ IL[DWLRQ E\ HSLSK\WLF PLFRUJDQLVPV LQ ULFH
%$ :KLWWRQ ILHOGV ,55, 5HVHDUFK 3DSHUV ,QWHUQDWLRQDO 5LFH 5HVHDUFK ,QVWLWXWH 32 %R[ 0DQLOD 3KLOLSSLQHV /DGKD -. DQG 5HGG\ 30 ([WHQVLRQ RI QLWURJHQ IL[DWLRQ WR ULFH QHFHVVLW\ DQG SRVVLELOLWLHV *HR -RXUQDO /DQJH 2/ %HOQDS - DQG 5HLFKHQEHUJHU + 3KRWRV\QWKHVLV RI WKH F\DQREDFWHULDO VRLOFUXVW OLFKHQ Collema tenax IURP DULG ODQGV LQ VRXWKHUQ 8WDK 86$ UROH RI ZDHU FRQWHQW RQ OLJKW DQG WHPSHUDWXUH UHVSRQVHV RI &2 H[FKDQJH )XQFWLRQDO (FRORJ\ /DQJH : 6SHFXODWLRQV RQ D SRVVLEOH HVVHQWLDO IXQFWLRQ RI WKH JHODWLQRXV VKHDWK RI EOXHJUHHQ DOJDH &DQ - 0LFURELRO /HJDQp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ostoc commune IURP D KHUEDULXP VSHFLPHQ HLJKW\VHYHQ \HDUV ROG %XOO 7RUUH\ &OXE /XPSNLQ 7$ DQG 3OXFNQHWW '/ Azolla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ostoc muscorum. %LRWHFKQRO /HWW 0RRUH $: 2FFXUUHQFH RI QRQV\PELRWLF QLWURJHQIL[LQJ PLFURRUJDQLVPV LQ 1LJHULDQ VRLOV 3ODQW DQG 6RLO
&KDSWHU 6RLOV DQG 5LFH)LHOGV 0UR]LQVND 7 $HURSK\WLF DOJDH IURP 1RUWK .RUHD $OJRORJLFDO 6WXGLHV 1DWHVDQ 5 DQG 6KDQPXJDVXQGDUDP 6 ([WUDFHOOXODU SKRVSKDWH VROXELOL]DWLRQ E\ WKH F\DQREDFWHULXP Anabaena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´ 6RYLHW 6RLO 6FL 3DUNHU %& 6FKDQHQ 1 DQG 5HQQHU 5 9LDEOH VRLO DOJDH IURP WKHKHUEDULXPRI WKH 0LVVRXUL%RWDQLFDO*DUGHQ $QQ 0LVVRXUL %RW *DUG 3DUNV -0 DQG 5LFH (/ (IIHFWV RI FHUWDLQ SODQWV RI ROGILHOG VXFFHVVLRQ RQ WKH JURZWK RI EOXHJUHHQ DOJDH %XOO 7RUU %RW &OXE 3HDUVRQ +: +RZVOH\ 5 .MHOGVRQ &. DQG :DOVE\ $( $HURELF QLWURJHQDVHDFWLYLW\ E\ D[HQLF FXOWXUHVRI WKH EOXH JUHHQ DOJD Microcoleus chthonoplastes. )(06 0LFURELRO /HWW 3HGHUVRQ &/ DQG 6KXEHUW / 8VH RI PHPEUDQH ILOWHUV IRU VRLO DOJDO ELRDVVD\V - $SSO 3K\FRO 3HGXUDQG 3 DQG 5H\QDXG 3$ 'R F\DQREDFWHULD HQKDQFH JHUPLQDWLRQ DQG JURZWK RI ULFH" 3ODQW DQG 6RLO 3HWHUVRQ +* %RXWLQ & )UHHPDUN .( DQG 0DUWLQ 3$ 7R[LFLW\ RI KH[D]LQRQH DQG GLTXDW WR JUHHQ DOJDH GLDWRPV F\DQREDFWHULD DQG GXFNZHHG $TXDWLF 7R[LFRORJ\ 3RWWV 0 DQG %RZPDQ 0$ 6HQVLWLYLW\ RI 1RVWRF commune 87(; &\DQREDFWHULD WR ZDWHU VWUHVV $UFK 0LFURELRO 3UDVDG %1 DQG 6ULYDVWDYD 31 6\VWHPDWLF DQG HFRORJLFDO VWXGLHV RQ DOJDH RI ODNDOLQH 8VDU VRLOV 3KXNRV 3URVSHUL & %ROXGD / /XQD & DQG )HUQDQGH]9DOLHQWH ( (QYLURQPHQWDO IDFWRUV DIIHFWLQJ in vitro QLWURJHQDVH DFWLYLW\ RI F\DQREDFWHULDLVRODWHGIURPULFHILHOGV- $SSO 3K\FRO 4XHVDGD $ /HJDQLV ) DQG )HUQiQGH]9DOLHQWH ( (QYLURQPHQWDO IDFWRUV FRQWUROOLQJ 1 IL[DWLRQ LQ 0HGLWHUUDQHDQ ULFH ILHOGV 0LFURE (FRO 4XHVDGD $ 1LHYD 0 /HJDQLV ( 8FKD $ 3URVSHUL & DQG )HUQiQGH]9DOLHQWH ( $FFOLPDWLRQ RI F\DQREDFWHULDO FRPPXQLWLHV LQ ULFHILHOGVDQGUHVSRQVHRI QLWURJHQDVHDFWLYLW\ WR OLJKW UHJLPH 0LFURE (FRO 5DQD %& *RSDO 7 DQG .XPDU +' 6WXGLHV RQ WKH ELRORJLFDO HIIHFWV RI LQGXVWULDO ZDVWHV RQ WKH JURZWK RI DOJDH (QYLURQ +HDOWK 5DR '/1 DQG %XPV 5* 7KH HIIHFW RI VXUIDFH JURZWK RI EOXHJUHHQ DOJDH DQG EU\RSK\WHV RQ VRPH PLFURELRORJLFDO ELRFKHPLFDO DQG SK\VLFDO VRLO SURSHUWLHV %LRO )HUWLO 6RLOV 5DWK % DQG $GKLNDU\ 63 7R[LFLW\ RI IXUDGDQ WR VHYHUDO 1 IL[LQJ F\DQREDFWHULDIURP WKH ULFH ILHOGV RI FRDVWDO 2ULVVD ,QGLD 7URSLFDO $JULFXOWXUH
5HGG\ 30 &RQWURO RI VSRUXODWLRQ LQ VRPH EOXHJUHHQ DOJDH $UFK +\GURELRO 6XSSO 5HGG\ 30 DQG 5RJHU 3$ '\QDPLFV RI DOJDO SRSXODWLRQV DQG DFHW\OHQHUHGXFLQJDFWLYLW\ LQ ILYH ULFH VRLOV LQRFXODWHG ZLWK EOXHJUHHQ DOJDH %LRO )HUWLO 6RLOV 5H\QDXG 3$ DQG )UDQFKH & ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI QRQKHWHURF\VWRXV WURSLFDO F\DQREDFWHULD JURZLQJ RQ QLWURJHQIUHHPHGLXP 0/5&(1 - 5H\QDXG 3$ DQG 5RJHU 3$ 0LOLHX[ VpOHFWLIV SRXU OD QXPLUDWLRQ GHV DOJXHV HXFDU\RWHV SURFDU\RWHV HW IL[DWULFHV G¶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µXQ F\FOH GH FXOWXUH GDQV XQH UL]LqUH 6DKHOLHQQH 5HY (FRO %LRO 6RO 5RJHU 3$ DQG 5H\QDXG 3$ /D ELRPDVVH DOJDOH GDQV OHV UL]LqUHV GH 6pQpJDO LPSRUWDQFH UHODWLYH GHV F\DQRSK\Fp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|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zolla DQG EOXHJUHHQ DOJDO ELRIHUWLOL]DWLRQ RI ULFH - $JULF 6FL 6LQJK /- 7LZDUL '1 DQG 6LQJK +1 (YLGHQFH IRU JHQHWLF FRQWURO RI KHUELFLGH UHVLVWDQFH LQ D ULFHILHOG LVRODWH RI Gloeocapsa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
%$ :KLWWRQ 6PLWK *' /\QFK 50 -DFREVRQ * DQG %DPHV &- &\DQREDFWHULDO QLWURJHQ IL[DWLRQ LQ DULG VRLOV RI &HQWUDO $XVWUDOLD )(06 0LFURE (FRO 6ROKHLP % (QGDO $ DQG 9LJVWDG + 1LWURJHQ IL[DWLRQ LQ $UFWLF YHJHWDWLRQ DQG VRLOV IURP 6YDOEDUG 1RUZD\ 3RODU %LRORJ\ 6SLOOHU + /DWWRUH & +DVVDQ 0( DQG 6KDQPXJDQ .7 ,VRODWLRQDQGFKDUDFWHUL]DWLRQRI QLWURJHQDVHGHSUHVVHGPXWDQW VWUDLQV RI F\DQREDFWHULXP Anabaena variabilis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p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
:\QQ:LOOLDPV '' 7HOHYLVLRQ LPDJH DQDO\VLV RI PLFURELDOFRPPXQLWLHVLQ$QWDUFWLFIHOOILHOGV 3RODUIRUVFKXQJ :\QQ:LOOLDPV'' 0LFURELDOFRORQL]DWLRQSURFHVVHVLQ $QWDUFWLF IHOOILHOG VRLOV DQH[SHULPHQWDO RYHUYLHZ 3URF 1DWLRQDO ,QVWLWXWH RI 3RODU 5HVHDUFK 6\PSRVLXP RQ 3RODU %LRORJ\
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loeocapsa ZKLOH VLWHV ZLWK PDUNHG WHPSRUDO GLIIHUHQFHV LQ DPELHQW SKRVSKDWH DUH RIWHQ GRPLQDWHG E\ KDLUIRUPLQJ JHQHUD HJ Rivularia &DOFLXP FDUERQDWH GHSRVLWLRQ B A Whitton and M Potts (eds). The Ecology of Cyanobacteria. © Kluwer Academic Publishers Printed in the Netherlands
$ 3HQWHFRVWDQG%$ :KLWWRQ
VRPHWLPHVOHDGVWRDSHUPDQHQWLQFUHDVHLQWKHDPRXQWRIOLPHVWRQHSUHVHQW DVZLWK WUDYHUWLQHLQIUHVKZDWHUV RU VWURPDWROLWHVLQERWKIUHVKZDWHUDQGPRUHHVSHFLDOO\PDULQHHQYLURQPHQWV (QGROLWKLFVSHFLHVRFFXULQDYDULHW\RIVLWXDWLRQVEXWSDUWLFXODUO\ ZKHUHWKHOLPHVWRQHLV VRIWDQGSRURXV )RU H[DPSOHVHYHUDOJHQHUDHJHyella JURZDFWLYHO\LQWRWKHOLPHVWRQHDWVLWHV ZKHUHUHPRYDORIVXUIDFHPDWHULDO WHQGV WR RFFXU 6XFK HQGROLWKLF FRPPXQLWLHV DUH EHVW GHYHORSHG RQ YHUWLFDO VXUIDFHV LQ WKH LQWHUWLGDO ]RQH RI WURSLFDODQGVXEWURSLFDOVHDVZKHUHWKHPRUHVXSHUILFLDOSDUWVRIWKHFRPPXQLWLHVDUHFRQWLQXDOO\EHLQJUHPRYHG E\JUD]LQJDQLPDOV
, ,QWURGXFWLRQ ,WKDV ORQJEHHQNQRZQ WKDWOLPHVWRQH VXUIDFHV RIWHQ KDYH YLVXDOO\ FRQVSLFXRXV JURZWKV RI F\DQREDFWHULD DQG WKH VXUIDFHV DV D ZKROH PD\ EH ULFK LQ VSHFLHV -DDJ *ROXELF 7KLV FKDSWHU SURYLGHV DQ RYHUYLHZ RI WKH FRPPXQLWLHV SUHVHQW DQG WKHLU FRPSRQHQW F\DQREDFWHULD *ROXELF¶V UHYLHZ LQ DQ HDUOLHU YROXPH VWLOO SURYLGHV DQ H[FHOOHQW LQWURGXFWLRQ WR WKH VXEMHFW DQG DVSHFWV FRYHUHG LQ GHWDLOWKHUHDUHPHQWLRQHGRQO\EULHIO\KHUH 6RPH RI WKH GLYHUVLW\ RI F\DQREDFWHULD DVVRFLDWHG ZLWKOLPHVWRQHVXUIDFHVFDQSUREDEO\EHH[SODLQHGE\ WKH IHDWXUHV RIWKH OLPHVWRQHV WKHPVHOYHV 7KHVH DUH D GLYHUVH JURXS RI VHGLPHQWV OLQNHG E\ WKHLU KLJK FDOFLXP FDUERQDWH FRQWHQW ! ZW 7KH\ UDQJH IURP PHWDPRUSKRVHG OLPHVWRQHV ZLWK YHU\ ORZ PDJQHVLXP FRQWHQW WR KLJKPDJQHVLDQ FDOFLWHV DQG GRORPLWHV ZKLFK PD\ UHDFK PROH 0J&2 7KH\ YDU\ ZLGHO\ LQ RULJLQ DQG LQ WKHLU SRURVLW\ GHQVLW\ DQG HURGDELOLW\ 7XFNHU DQG :ULJKW 7KH\DOVRYDU\ZLGHO\LQFKHPLFDOFRQVWLWXHQWV ZKLFK PLJKW EH H[SHFWHG WR LQIOXHQFH F\DQREDFWHULDO JURZWK VXFK DV SKRVSKDWH 'DYLV DQG 5DQGV DQGRUJDQLFFRQWHQW 7KH F\DQREDFWHULDO FRPPXQLWLHV RI OLPHVWRQH VXUIDFHV LQFOXGH WKRVH RI GHVHUW URFNV FOLIIV RI GU\ DQG KXPLG UHJLRQV YDULRXV W\SHV RI HPHUJHQW VWUXFWXUHIRUPHGE\HURVLRQWKHLQXQGDWHGOLPHVWRQHV RIODNHV ULYHUV DQG VWUHDPV DQG PDULQH H[DPSOHV LQ WKH LQWHUWLGDO ]RQH +LJKO\ FDOFDUHRXV IUHVKZDWHUV VRPHWLPHV EXW QRW DOZD\V KDYH FRQVSLFXRXV F\DQREDFWHULDO FRPPXQLWLHV &\DQREDFWHULD FDQ DOVR IRUPLPSRUWDQWFRPPXQLWLHV RQOLPHVWRQHDUWLIDFWV RU FRPPHUFLDO SURGXFWV ZKHUH OLPHVWRQH KDV EHHQ LQFRUSRUDWHG $ ZLGH UDQJH RIHFRV\VWHPV ZLOO WKXV EH FRQVLGHUHG LQ UHODWLRQ WR ZDWHU UHJLPH DQG FOLPDWLF WLGDO DQG KXPDQ IDFWRUV 7KH RUJDQLVPV ZKLFK LQFOXGH WKH F\DQROLFKHQV PD\ EH HSLOLWKLF RU HQGROLWKLF 6RPH F\DQREDFWHULD DUH QRWHG IRU WKHLU DELOLW\ WR SUHFLSLWDWH WUDS DQG ELQG SDUWLFOHV RI FDOFLXP FDUERQDWH WR IRUP DFFXPXODWLRQV RI OD\HUHG WUDYHUWLQH RUVWURPDWROLWH ,QDGGLWLRQ VRPH DUHDEOH WR HWFK OLPHVWRQH DQG JURZ ZLWKLQ WKH VXEWUDWH *ROXELF HW DO 7KLV DELOLW\ RI D VSHFLHV WR
GHSRVLWRUWRHURGHOLPHVWRQHSURYLGHVLWZLWKDSRWHQW PHDQVRIPRGLI\LQJLWVORFDOHQYLURQPHQW
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hormidium DQG Schizothrix VSS VKRZHG OLWWOH YLVXDOO\ REYLRXV FKDQJH VR WKDW DQ RUGHU RIPDJQLWXGH FKDQJH LQ + KDGQRPHDVXUDEOHHIIHFW $VWXG\'DYLVDQG5DQGV RQWKH JURZWKRI Hapalosiphon intricatus LQ D )ORULGD FDYH LQ UHODWLRQ WR WKH PLQHUDO FRPSRVLWLRQ RI WKH XQGHUO\LQJ OLPHVWRQH IRXQG WKDW WKH SKRVSKDWH FRQFHQWUDWLRQ LQ WKHOLPHVWRQHZDV J J J3 J ZKLOH 12 UDQJHGIURP JJ JN J DQG 1+ IURP J J J 1 J ,Q VROXWLRQV LQ HTXLOLEULXP ZLWK OLPHVWRQH VDPSOHV WKH SKRVSKDWHFRQFHQWUDWLRQUDQJHGIURP J/ J 3 / 7KH DXWKRUV VXJJHVWHG WKDW WKH FRPELQHG QLWURJHQ FRXOG KDYH EHHQ GHULYHG SDUWO\ IURP VHHSDJH DEVRUSWLRQ RI DPPRQLD IURP WKH UHVLGHQW EDW SRSXODWLRQ DQG QLWURJHQ IL[DWLRQ E\ WKH Hapalosiphon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
,,, (QYLURQPHQWV DQG WKHLU &KDUDFWHULVWLF &RPPXQLWLHV $ 7HUUHVWULDO 2YHUYLHZ 7HUUHVWULDO OLPHVWRQHV LQFOXGH GLYHUVH PRUSKRORJLHV VXFK DV FOLIIV IODWSDYHPHQW DQG HPHUJHQWSLQQDFOHV 6XFKVXUIDFHVDUHPRVWO\ GRPLQDWHG E\F\DQREDFWHULD DQG OLFKHQV LQ YDU\LQJ SURSRUWLRQV 7KH SKRWRWURSK RI WKH OLFKHQV LV XVXDOO\ D JUHHQ DOJD EXW F\DQROLFKHQV DUH VRPHWLPHV LPSRUWDQW VHH EHORZ &\DQREDFWHULD DSSHDU WR EH WKH GRPLQDQWV RI VXEWURSLFDO DQG WURSLFDO OLPHVWRQHV ZKHUH Tolypothrix byssoidea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
$ 3HQWHFRVW DQG%$:KLWWRQ
Fig. 1. ([WHQVLYHDUHDV RIIODWOLPHVWRQHSODWLQ RQ$OGDEUD $WROO ODUJHO\EDUH RIKLJKHUSODQWYHJHWDWLRQ DUH FRYHUHGE\ DF\DQREDFWHULDO FRPPXQLW\GRPLQDWHGE\Tolypothrix byssoidea.
PRLVW HQRXJK WR FDUU\ RXW PHWDEROLF DFWLYLW\ ,Q PDQ\ VLWXDWLRQV VXFK DV WKH FRPPXQLWLHV RQ FOLIIV GHVFULEHG EHORZ LW LV SRVVLEOHWR FRPSDUH DUDQJH RI F\DQREDFWHULDO FRPPXQLWLHV DVVRFLDWHG ZLWK LQFUHDVLQJ SHULRGV RI PRLVW RU LQ VRPH FDVHV WHPSRUDULO\VXEPHUJHGFRQGLWLRQV 0DQ\ VWXGLHV KDYH UHSRUWHG WKDW F\DQREDFWHULD FDQ EHLPSRUWDQWLQWKHZHDWKHULQJRIWHUUHVWULDOOLPHVWRQH VXUIDFHV RIWHQ LQ DVVRFLDWLRQ ZLWK RWKHU RUJDQLVPV VXFK DV WKH FRPELQHG HIIHFWV RI F\DQREDFWHULD DQG OLFKHQV UHSRUWHG IRU KLOOVLGHV LQ WKH 1HJHY ,VUDHO 'DQLQ DQG *DUW\ :HDWKHULQJ GXH WR F\DQREDFWHULD LQYROYHV D YDULHW\ RI SURFHVVHV LQFOXGLQJ WKH JURZWK RI HQGROLWKV &KDSWHU 6RPHWLPHV WKH JURZWK RI HQGROLWKV PD\ EH FRQVLGHUHG RSSRUWXQLVWLF PDNLQJ XVH RI FUDFNV DQG KROHV DOUHDG\ SUHVHQW VXFK DV ZLWK WKH HQGROLWKLF FRPPXQLW\ GRPLQDWHG E\ Gloeocapsa LQ WKH GRORPLWLF OLPHVWRQH VWXGLHG E\ )HUULV DQG /RZVRQ VHH 6HFWLRQ ,, DERYH +RZHYHU ERULQJ E\ F\DQREDFWHULDPD\ DOVR SOD\ DUROH 9LOHV ,W LV IDU IURP FOHDU ZK\ VRPH OLPHVWRQH VXUIDFHV DUH PRUHVXEMHFWWRERULQJWKDQRWKHUVEXWLWVHHPVOLNHO\ WKDW DYDLODELOLW\ RI QXWULHQWV HVSHFLDOO\ SKRVSKDWH PD\ EH DQ LPSRUWDQW IDFWRU 3UHVXPDEO\ LW LV RQO\
HQHUJHWLFDOO\ SRVVLEOH IRU FKDUDFWHULVWLF ERULQJ VSHFLHV VXFK DV Plectonema terebrans, WR FDUU\ RXW WKH PHWDEROLF DFWLYLW\ UHTXLUHG IRU HWFKLQJ OLPHVWRQH XQGHUHVSHFLDOO\IDYRXUDEOHQXWULHQWFRQGLWLRQV 'HSRVLWLRQ PD\ DOVR RFFXU VRPHWLPHV LQ FORVH SUR[LPLW\ WR HURVLRQDO SURFHVVHV )RU LQVWDQFH 9LOHV V VWXG\ RI F\DQREDFWHULDO ERULQJ RQ WHUUHVWULDO VXUIDFHV RI $OGDEUD IRXQG WKDW WKHUH ZDV DOVR SUHFLSLWDWLRQ RI PLFULWH DURXQG WKH ILODPHQWV VRPHWLPHV IRUPLQJ DFUXVW VHYHUDO PP WKLFN ,QWKH FDVH RIIyengariella endolithicu LQ D VSULQJIHG ODNH LQ 1( 0H[LFR WKH RUJDQLVP ZDV RIWHQ ERULQJ LQWR WKH URFN XQGHUQHDWK D ]RQH RI DFWLYH WUDYHUWLQH GHSRVLWLRQ 6HHOHU DQG *ROXELF 'HSRVLWLRQ DQG HURVLRQDO SURFHVVHV DUH WUHDWHG PRUH IXOO\ LQ 6HFWLRQV 9 DQG 9, UHVSHFWLYHO\ 7ZR W\SHV RI WHUUHVWULDO HQYLURQPHQW DUH GHVFULEHG LQ PRUH GHWDLO KHUH
7LQWHQVWULFKH &RORXUIXO YHUWLFDO VWUHDNV RI F\DQREDFWHULD PD\ EH IRXQGRQVWHHSFOLIIVZRUOGZLGH3ODWHVDE DQG DUH DVVRFLDWHG ZLWK WHPSRUDU\ ZDWHU UXQQHOV ZKLFK PD\EHFRQVLGHUHGDVH[WUHPHH[DPSOHVRIHSKHPHUDO
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cytonemo gloeocapsetum WR D Tolypothricetum byssoideae DQG ILQDOO\ D Dichothricetum compactaea. &KDUDFWHULVWLF VSHFLHV RI WKH ILUVW PRVW [HULF DVVRFLDWLRQ LQFOXGH Scytonema myochrous, Gloeocapsa compacta, G. kuetzingiana DQG G. sanguinea, ZKLOH WKH ZHWWHVW VLWHV KDG Dichothrix compacta, Phormidium autumnale DQG Xenococcus kerneri. 8QGHU ORZ LOOXPLQDWLRQ DVXLWHODUJHO\GRPLQDWHGE\Schizothrix VSSRFFXUV 7KH DXWKRUV KDYH REVHUYHG HOHPHQWV RI WKHVHDVVRFLDWLRQVLQWKH8QLWHG .LQJGRPDQG VLPLODU FRPPXQLWLHV RFFXU HOVHZKHUH HJ )MHUGLQJVWDG 7KH VWXG\ RI WLQWHQVWULFKH DGGV WR WKH LQIRUPDWLRQ RQ F\DQREDFWHULDO ZDWHU UHODWLRQV IURP RWKHU W\SHV RI HQYLURQPHQW &KDSWHUV )RU H[DPSOH Gloeocapsa calcarea, DIWHU UHPDLQLQJ LQ D GHVLFFDWRU IRU PRQWKV EHJDQ WR IL[ FDUERQ VRRQ DIWHU UHZHWWLQJ 3HQWHFRVW E ZKLOH WKH Schizothrix VSHFLHV RI SHUPDQHQWO\ ZHW WLQWHQVWULFKH DSSHDU PXFK OHVV WROHUDQW 7KRVH VSHFLHV ZLWK WKH WKLFNHVW DQG RIWHQ SLJPHQWHG VKHDWKV RFFXU DW WKH GULHVW VLWHV ZKLOH WKLQVKHDWKHG VSHFLHV RFFXU LQ VWUHDPV 7KH UROH RI WKH \HOORZEURZQ VKHDWK SLJPHQW VF\WRQHPLQ *DUFLD3LFKHO DQG &DVWHQKRO] LQ SURWHFWLQJ DJDLQVW XOWUDYLROHW OLJKW DQG WKH SRVVLEO\ VLPLODU UROH RIWKH S+VHQVLWLYH JORHRFDSVLQ DUHGHVFULEHGLQ&KDSWHU6HFWLRQ,,$
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phanocapsa grevillei, Chroococcus turgidus YDU spelaeus DQG Gloeocapsa kuetzingiana. 6LPLODU DVVRFLDWLRQV KDYH EHHQ QRWHG LQ WKH 8. DXWKRUV XQSXEOLVKHGGDWD DQG&KLQD 7LDQDQG+H DQG SUREDEO\ RFFXU ZRUOGZLGH +RZHYHU D VWXG\ RI VSHFLHVGLYHUVLW\ LQ D YHU\ ROG FDYH LQ ,VUDHO VKRZHG WKDW WKH SURSRUWLRQ RI FRFFRLG WR ILODPHQWRXV IRUPV GHFUHDVHG DV LUUDGLDQFH JRW OHVV 9LQRJUDGRYD HW DO 7ZHQW\ JHQHUDDQG VSHFLHV ZHUH IRXQG LQ WKH FDYH DQG JHQHULF DQG VSHFLHV GLYHUVLW\ ERWK GHFUHDVHGZLWKGHFUHDVLQJLUUDGLDQFH &DYH Gloeocapsa VSS KDYH EHHQ IRXQG ZKHUH WKH OLJKWLQWHQVLW\LVDVORZDV OX[FDSPROSKRWRQ P V 3$5 &R[ HW DO &HOO GLDPHWHUV DYHUDJH P DQG DW WKLV ORZ LUUDGLDQFH ZRXOG H[SHULHQFHDIOX[RI SKRWRQ V 3$5 ZKLFK ZLWK DQ HVWLPDWHG SKRWRV\WKHWLF HIILFLHQF\ RI ZRXOG UHVXOWLQDFHOOGLYLVLRQWLPHRIGD\V 7KHFHOOV RI FDYHLQKDELWLQJ F\DQREDFWHULD FRQWDLQ GHQVHO\ SDFNHG WK\ODNRLGV &R[ HW DO &RXWp &RXWp DQG %XU\ D FOHDU DGDSWDWLRQ WR ORZOLJKWOHYHOV 8QOLNHWKHORZOLJKWFRQGLWLRQVWREH IRXQGLQGHHS ZDWHU VSHFWUDOOLJKWTXDOLW\ LQFDYHVLV SUDFWLFDOO\ LQYDULDQW ZLWK LOOXPLQDWLRQ DQG FKURPDWLF DGDSWDWLRQKDVQRWEHHQREVHUYHG 0RVW RIWKHVH F\DQREDFWHULD FDQ DOVR EH IRXQG RQ VKDGHG OLPHVWRQH FOLIIV EXW D IHZ DSSHDU WR EH VWULFW FDYHUQLFROHV 7ZR VSHFLHV RI Geitleria DUH NQRZQ IURP FDYHV ZKHUH WKH\ PD\ EH UHFRJQLVHG DV D GHOLFDWH DVKHQ IHOW RQ FDYH ZDOOV G. calcarea LV ZLGHO\ GLVWULEXWHG LQ ZDUP FOLPDWHV )ULHGPDQQ ZKHUH LWPD\ EH IRXQG DW YHU\ ORZ LUUDGLDQFH /HFOHUF HW DO &RORQLHV RI Scytonema julianum IRUP D VLPLODU IHOW EXWWKLV VSHFLHV LV PRUH ZLGHO\ GLVWULEXWHG DQG LV QRW UHVWULFWHG WR FDYH HQWUDQFHV ,QERWKFDVHVWKHXQXVXDOSDOHYLROHWJUH\ FRORXULVFDXVHGE\FDOFLILFDWLRQ &DOFLWHLVGHSRVLWHG
$ 3HQWHFRVWDQG%$ :KLWWRQ
Fig. 2. 6FDQQLQJHOHFWURQPLFURJUDSKRI6F\WRQHPDMXOLDQXPIURP6SULQJ&DYHV0DOKDP(QJODQG 7KHVKHDWKV DUHFRYHUHGZLWK VSLN\ RFFDVLRQDOO\GHQGULWLFFU\VWDOVRIFDOFLWH $LUGULHGVDPSOH$X3GFRDWHG %DU P
RQ WKH ILODPHQWV DV GHQVH UDGLDWLQJ PLFURFU\VWDOV IRUPLQJDOD\HUXSWRPWKLFN)LJ 7KHPLQHUDOLVDWLRQRIGeitleria DSSHDUV WR EH PRUH UHJXODU WKDQ WKDW RI Scytonema )ULHGPDQQ &RXWp DQG %XU\ DQG WKLV KDV OHG WR WKH VXJJHVWLRQ WKDW LW LV ELRORJLFDOO\ FRQWUROOHG 7KHVH HQFUXVWDWLRQV PD\ KRZHYHU VLPSO\ EH WKH UHVXOW RI VWHDG\ HYDSRUDWLRQ RI FDUERQDWHULFK ZDWHU VHHSDJHV DURXQG WKH ILODPHQWV $ELRJHQLF FDOFLWH QRGXOHV DUH FRPPRQ DW FDYH HQWUDQFHV DQG QHHGOHILEUH FDOFLWHV PRUSKRORJLFDOO\ VLPLODU WR WKHVH F\DQREDFWHULDO FDOFLWHV DUH D FRPPRQ IHDWXUH RI FDYH VHHS ]RQHV 9HUUHFFKLDDQG9HUUHFFKLD 1HYHUWKHOHVVWKH UHJXODU DUUD\V RIPLFURFU\VWDOV RFFDVLRQDOO\ IRXQG RQ Geitleria PDNH D FRPSHOOLQJ DUJXPHQW WKDW VRPH IRUPRIELRORJLFDOO\LQGXFHGFRQWUROLVWDNLQJSODFH .QRZOHGJH RI FDYH F\DQREDFWHULD LV DW EHVW UXGLPHQWDU\ \HW FDYHV SURYLGH DQ H[FHOOHQW QDWXUDO ODERUDWRU\ IRU WKH LQYHVWLJDWLRQ RI FKDQJLQJ LOOXPLQDWLRQ XQGHU FRQGLWLRQV RI DOPRVW FRQVWDQW WHPSHUDWXUH 7KH\DZDLWIXUWKHUH[SORLWDWLRQ
% )UHVKZDWHU 7KH IUHVKZDWHU HQYLURQPHQW DSSHDUV WKH OHDVW VWUHVVIXO RIFDOFDUHRXV HQYLURQPHQWV EXW WKLV LV RQO\ WUXHZKHUHWKHUHLVSHUPDQHQWZDWHUFRYHU 7KHUHDUH D QXPEHU RI UHFRUGV RI ODNHV ZLWK DQ H[WHQVLYH ERWWRP FRYHU RI ILUP VWURPDWROLWLF VWUXFWXUHV ZKHUH F\DQREDFWHULD DSSHDU WR SOD\ DQ LPSRUWDQW UROH LQ WKHLU IRUPDWLRQ Rivularia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hormidium, Lyngbya DQG Pleurocapsa;
&KDSWHU/LPHVWRQHV Entophysalis, Lyngbya DQG Pleurocapsa; Pleurocapsa, Dichothrix gypsophila DQG Schizothrix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cytonema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yella VSS Solentia, Plectonema terebrans, Mastigocoleus testarum, Kyrtuthrix DQG Brachytrichia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
$ 3HQWHFRVWDQG %$:KLWWRQ
Table 1. )UHTXHQWO\UHFRUGHGF\DQREDFWHULDRI OLPHVWRQHVXUIDFHVLQIUHVKZDWHUVHVSHFLDOO\WKRVHZKHUHWUDYHUWLQHGHSRVLWLRQLV WDNLQJ SODFH (FRORJLFDOJHQHUDOL]DWLRQV DUHEDVHG ODUJHO\RQREVHUYDWLRQV LQ QRUWKHUQ DQGFHQWUDO (XURSH 7KHRUGHUV OLVWHG DUHWKRVH RI &DVWHQKRO] DQG :DWHUEXU\ 0 PHWHRJHQH 7 WKHUPRJHQH VFDOHLQGLFDWLQJIUHTXHQF\LQ SDUWLFXODUFDWHJRU\ ZLWK LQIUHTXHQWWR YHU\ ZLGHVSUHDG
6SHFLHV &KURRFRFFDOHV Aphanocapsa grevillei
07 1RWHV
5HIHUHQFHV
Gloeocapsa calcarea G. rupestris Synechococcus elongatus S. minervae
6PDOOGLVFUHWHFRORQLHVDPRQJ ILODPHQWRXVPLFUREHV1R FDOFLILFDWLRQZLWKLQVKHDWK :LGHVSUHDGDVDERYH 6XSHUILFLDOFUXVWVRQ VLWHVOLDEOHWRGHVLFFDWLRQ 2 6RPHWLPHVSURYLGLQJQXFOHL IRUDUDJRQLWHGHSRVLWLRQ 'LVWLQFWLYHVSHFLHVZLWKHORQJDWHFHOOVQRWNQRZQ WRFDOFLI\
3OHXURFDSVDOHV Hyella fontana
Pleurocapsa minor
,VRODWHG ILODPHQWV PD\ EHKDYHDVHQGROLWKVLQ VORZO\ GHSRVLWLQJWUDYHUWLQH 3UREDEO\VRPHWLPHVHQGROLWKLFFDOFLILFDWLRQRFFDVLRQDOO\ UHSRUWHG
*UQLQJHU 3HQWHFRVW 3HQWHFRVW E 3HQWHFRVWF &RSHODQG 3HQWHFRVW F *ROXELF .UXPEHLQ 3RWWV 2UGRQH] *DUFLD GHO &XUD
2VFLOODWRULDOHV Homoeothrix balearica
H. crustacea
H. janthina
Lyngbya martensiana
L. valderianum
L. vandenberghenii
Phormidium calcicola
/DUJHVSHFLHVIRUPLQJKDUG\HOORZLVKQRGXOHV+HDYLO\ FDOFLILHG :HOOGHILQHG\HOORZEURZQ FRORQLHV PP LQGLDPHWHU +HDYLO\FDOFLILHG3ODWHG :HDNO\ FDOFLILHGVSHFLHVFKDUDFWHULVHGE\ WKHP ILODPHQWV DQG GDUN JUHHQFRORXUen PDVVH 6HYHUDORWKHU FDOFLILHGVSHFLHVDUHNQRZQ 2IWHQDVVRFLDWHGZLWK Phormidium incrustatum DQG GLVWLQJXLVKHG E\ ZLGHUSP WULFKRPHV 8VXDOO\ DVVRFLDWHG ZLWK P. incrustatum, EXWQDUURZHU SP WULFKRPHV 6PDOO ULYHUV ZLWK KDUGZDWHUXQGHUORZ IORZXQGHUVXFK FRQGLWLRQVWKHFRPELQDWLRQRI WKHRUJDQLVPDQGDWKLQ FDUERQDWHGHSRVLWJLYHWKHVXUIDFHD FKDFWHULVWLF PLON\ EOXH DSSHDUDQFH 6LPLODUWRP. incrustaturn, EXWDSSDUHQWO\OHVVFRPPRQ
FRORQL]DWLRQ UHTXLUHV WKH IRUPDWLRQ RI VPDOO FHOOV IURP QRQILODPHQWRXV IRUPV RU KRUPRJRQLD IURP ILODPHQWRXV IRUPV $O7KXNDLU DQG *ROXELF VKRZHG WKDW WKH VPDOO FHOOV EDHRF\WHV RI Hyella immanis ZHUH PRWLOH DQG SKRWRWD[LF IHDWXUHV ZKLFK VHHP OLNHO\ WR EH FKDUDFWHULVWLFRI WKH UHSURGXFWLYH VWUXFWXUHV RI DOOHXHQGROLWKV $JLWDWLRQRI ODERUDWRU\ FXOWXUHV RI WKLV VSHFLHV HQKDQFHG WKH IUHTXHQF\ RI ERULQJV DQG WKHLQLWLDOERULQJUDWHEXW KDG QR HIIHFW RQWKHFRQWLQXLQJERULQJDFWLYLW\ $OWKRXJK UHPRYDO RI WKH VXSHUILFLDOSDUWV RI WKH WKDOOXV E\ JUD]LQJ DQG HURVLRQ LV D NH\ SUREOHPIRU PRVW HQGROLWKLF F\DQREDFWHULD VRPHWLPHV WKH RUJDQLVPV JURZ LQ VLWXDWLRQV ZKHUH FDOFLXP FDUERQDWHLV LQFUHDVLQJDW WKH VXUIDFHDV VKRZQ IRU Plectonema terebrans LQ FUXVWRVH UHG DOJDH *KLUDGHOOL %RWK OLYH DQG GHDG DOJDH ZHUH LQFOXGHG LQ *KLUDGHOOL V VWXG\ EXW /H &DPSLRQ
3HQWHFRVW :LQVERURXJKHW DO 3HQWHFRVW &DVDQRYD
6\PRHQV 0DOODLVH 3HQWHFRVW HW DO 6\PRHQV YDQ GHU :HUII +ROPHV :KLWWRQ .DQQ
$OVXPDUG HW DO VKRZHG WKDW WKH HQGROLWKLF FRPPXQLWLHV RI OLYH DQG GHDG FRUDO RQ 0RRUHD )UHQFK 3RO\QHVLD GLIIHUHG 7KH HQGROLWKV RI OLYH FRUDODUHVHOHFWHGIRUSRVLWLYHSKRWRWURSLVPDQG IDVW JURZWK WR FRPSHWH ZLWK WKH DFFUHWLRQ UDWHV RI WKH FRUDOVNHOHWRQDV ORQJ DV WKHFRUDOUHPDLQVDOLYHWKH HQGROLWKV DUH SURWHFWHG IURP JUD]LQJ 7KH PDLQ HQGROLWK RI WKHFRUDO Porites lobata RQ 0RRUHD ZDV WKH JUHHQ DOJD Ostreobium quekettii. 'HDG DQG GHQXGHGSDUWVRI WKHFRUDOVNHOHWRQZHUHFRORQL]HGDW WKH VXUIDFH DQG ERUHG LQZDUG E\ D VXFFHVVLRQ RI HXHQGROLWKV VWDUWLQJZLWK WKH JUHHQDOJD Phaeophila dendroides, IROORZHG E\ Plectonema terebrans DQG Mastigocoleus testarum DQG OHDGLQJ ZLWKLQ \U WR D VWDEOH FRPPXQLW\ GRPLQDWHG E\ Ostreobium quekettii. 7KHHQGROLWKVLQ GHDG FRUDOV ZHUH JUD]HG E\ PROOXVFV HFKLQRGHUPV DQG VFDULG ILVK 6XFK JUD]LQJ LV LPSRUWDQW IRU WKH PDLQWHQDQFH RI WKH
&KDSWHU /LPHVWRQHV
Table1 FRQWLQXHG
6SHFLHV
07 1RWHV
P. incrustatum
P. laminosum
Plectonema tomasianum Schizothrix affinis S. calcicola S. coriacea S. fasciculata
S.rubella V\Q S. lardacea Spirulina labyrinthiformis
5HIHUHQFHV
3UHIHUHQFHIRUVKDGHGVLWHVZKHUH LW IRUPVµIUHVKZDWHU VWURPDWROLWHV¶ $EXQGDQWRQWUDYHUWLQHWHUUDFHVZKHUH IRUPVVPRRWKJUHHQ VWUDWD ZKLFK PD\ EHFRPHFDOFLILHG 6XSHUILFLDOJURZWKVUDUHO\HQFUXVWHG )RUPVDEURZQLVKQRGXODUFUXVW $µFOXVWHU¶VSHFLHVIRURWKHUFDOFLILHGSchizothrix )RXQG DV OLJKWO\HQFUXVWHGEURZQLVKVWUDWD3ODWHH 3DOH EURZQZHOOFDOFLILHGQRGXODUFRORQLHVRIWHQLQ VLWHV SURQH WR RFFDVLRQDOGHVLFFDWLRQ7KHUHDUH VHYHUDORWKHU FDOFLILHGVSHFLHV )RUPV SLQNUHGWKLQFUXVWV RQ LQWHUPLWWHQWO\PRLVWOLPHVWRQH URFNV3ODWHF /RRVHIORFVLQ WKHUPDO HIIOXHQWV XVXDOO\ FRQWDLQLQJVXOSKLGH 8VXDOO\ XQFDOFLILHG
)ULWVFK 3HQWHFRVWE )UH\WHW 3OD]LDW 3HQWHFRVW HW DO *OD]HN &DXGZHOO 3HQWHFRVW $OOHQ 3HQWHFRVW *ROXEL& *ROXEL&
3HQWHFRVW D
Nostocales Calothrix parietina
Dichothrix gypsophila
Nostoc sphaeroides Rivularia biasolettiana R. haematites
Scytonema crustaceum
Scytonema myochrous
Stigonematales Fischerella laminosus
)UHTXHQW DW LQWHUIDFHEHWZHHQIUHVKZDWHUDTXDWLFDQG WHUUHVWULDO 6RPHWLPHVRQRQFRLGVZKHUH VKHDWK FDOFLILFDWLRQKDVEHHQUHSRUWHG 6LPLODUKDELWVWRDERYHWROHUDQWRI KLJK VDOLQLW\ 2WKHU VSHFLHV JURZ RQ WKHUPRJHQHWUDYHUWLQHVXVXDOO\ FDOFLILHG 6XSHUILFLDO XVXDOO\ XQFDOFLILHGJURZWKVLQVXPPHU :HDNO\ FDOFLILHGPHUJLQJLQWRVSHFLHVEHORZ 'DUN EURZQKHDYLO\FDOFLILHGFRORQLHVZLWK FRQFHQWULF]RQHV RI FDOFLWH 2QH RI VHYHUDOVSHFLHVIRUPLQJGDUNVXSHUILFLDOWXUIV RQ WUDYHUWLQHV XVXDOO\ HQFUXVWHG EHORZ )RUPV ODUJH DQG FKDUDFWHULVWLF FXVKLRQV LQPRLVWFDOFDUHRXV HQYLURQPHQWVWUDSSLQJSDUWLFOHV 0D\ IRUPDZHDNO\FDOFLILHGFUXVW :LGHO\ GLVWULEXWHG LQ KRWVSULQJV
HXHQGROLWKLF FRPPXQLW\ DV RWKHUZLVH WKH VXUIDFH EHFRPHVRYHUJURZQZLWKDQHSLOLWKLFDOJDOWXUI /H%ULV HW DO UHSRUWHG RQ IXUWKHU GHWDLOHG VWXG\ RQ WKH FRORQL]DWLRQ RI EORFNV RI Porites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streobium. 7KH
3HQWHFRVW )MHUGLQJVWDG 3DULKDU 3DQW *ROXELF 2EHQOQHVFKORVV 7KXQPDUN :KLWWRQ HW DO
3HQWHFRVW 7HUU\
DXWKRUV VXJJHVWHGWKDWWKHH[WHQWRIHSLOLWKLF FRYHULQJ RUWKHULFKQHVVRIWKHHQGROLWKLFIORUDFRXOGEHXVHGDV D JOREDO LQGLFDWRU RIWKH TXDOLW\ DQG KHDOWK RID UHHI WKH KLJKHU WKH UDWLR RI HQGROLWKV WR HSLOLWKV DQG WKH KLJKHUWKHGLYHUVLW\RIWKHHQGROLWKV WKHPRUHKHDOWK\ WKHUHHI
,9 7KH2UJDQLVPV $ )UHHOLYLQJ6SHFLHV $ UHODWLYHO\ ZLGH UDQJH RI JHQHUD DQG VSHFLHV KDV EHHQ UHSRUWHG IURP FDOFDUHRXV HQYLURQPHQWV )RU LQVWDQFH RYHU VSHFLHV ZHUH UHSRUWHG IURP WHUUHVWULDO VXUIDFHV RI$OGDEUD $WROO :KLWWRQ :KLWWRQ DQG 'RQDOGVRQ 3RWWV DQG :KLWWRQ ,Q YLHZ RI WKH IDFW WKDW WKHUH DUH H[WHQVLYH DUHDV RI OLPHVWRQH LQ GLYHUVH FOLPDWLF UHJLRQV IRU
ZKLFK WKHUH KDYH VR IDU EHHQ QR UHFRUGV RI WKHLU F\DQREDFWHULDO IORUD OHW DORQH D PRGHUQ DWWHPSW WR DVVHVV WKHLU ELRGLYHUVLW\ LW VHHPV OLNHO\ WKDW WKH ZRUOG OLVW RI VSHFLHV ZLOO SURYH WR EH PXFK KLJKHU )RU LQVWDQFH WKH PDVVLYH OLPHVWRQH SLQQDFOHV IRXQG LQYDULRXVSDUWVRIWKHZHWWURSLFV VXFKDV0XOX3DUN LQ,QGRQHVLD KDYH\HWWREH VWXGLHG ([DPSOHV IURP PRVWO\(XURSHDQ VLWHV ZLWKWUDYHUWLQHGHSRVLWLRQDUH OLVWHG LQ 7DEOH 7KHVH VSHFLHV PD\ EH EURDGO\ FODVVLILHG LQWR WZR JURXSV WKRVH ZKLFK FDOFLI\ DQG HYHQWXDOO\ EHFRPH LQFRUSRUDWHG LQWR WKH GHSRVLW DQG WKRVH ZKLFK KDYH D PRUH IOHHWLQJ H[LVWHQFH DUH QRUPDOO\XQPLQHUDOLVHGDQGUDUHO\LIHYHUIRVVLOL]H 7KH SRVVLELOLW\ WKDW FDOFLILFDWLRQ PD\ SURYLGH D FULWHULRQ RIWD[RQRPLF YDOXH &DVWHQKRO] KDV EHHQGLVFXVVHGE\3HQWHFRVWDQG 5LGLQJ 0HU] DQG 0HU] DQG =DQNO $ FRQVHUYDWLYH WD[RQRPLF YLHZSRLQW KDV EHHQ DGRSWHG LQ SUHSDULQJ 7DEOH ZLWKFDOFLI\LQJIRUPV VRPHWLPHVUHFRJQL]HG DV VHSDUDWH VSHFLHV +RZHYHU PROHFXODU VWXGLHV DUH UHTXLUHG EHIRUH LW FDQ EH HVWDEOLVKHG ZKHWKHU WKH GLIIHUHQFH EHWZHHQ D FDOFLI\LQJ DQG DQ RWKHUZLVH FORVHO\ VLPLODU QRQFDOFLI\LQJ IRUP UHDOO\ MXVWLILHV UHFRJQLWLRQDWWKHVSHFLHVOHYHO 0RVW RIWKH HSLOLWKLF IRUPV KDYH VKHDWKV DQG IRUP REYLRXV FRORQLHV 7KH KHWHURF\VWRXV JHQHUD DUH PRVWO\ WKRVH ZLWK WULFKRPHV HQGLQJ LQ KDLUV DQ LQGLFDWLRQ WKDWWKH\ H[LVW IRU PXFK RIWKH WLPH XQGHU FRQGLWLRQV RI3 OLPLWDWLRQ 6HFWLRQ9,, $ 6RPH RI WKH ODUJHU FRORQLHV FDQ EH YHU\ ORQJOLYHG )RU LQVWDQFH Rivularia haematites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hormidium incrustatum LV IUHTXHQWO\ DWWDFNHG 7KH SDWKRJHQ KDV QRW EHHQ LGHQWLILHG EXW LQIHFWHG WULFKRPHVEHFRPHEOHDFKHGDQGRYHUUXQE\EDFWHULD
$ 3HQWHFRVWDQG%$:KLWWRQ %&\DQROLFKHQV 7KHVH OLFKHQV DUH GLVWLQJXLVKHG E\ WKHLU F\DQREDFWHULDO SKRWRELRQW DQG DUH D IDLUO\ VXFFHVVIXO JURXS FRORQL]LQJ D YDULHW\ RI WHUUHVWULDO DQG IUHVKZDWHUVXEVWUDWD 7KH OLFKHQL]HG F\DQREDFWHULDRI OLPHVWRQHV KDYH EHHQ ODUJHO\ LJQRUHG E\ HFRORJLVWV \HW DPRQJ WKH OLPHVWRQH PDFUROLFKHQV RIWKH %ULWLVK ,VOHVGHVFULEHGLQ3XUYLVHWDO DOPRVWDUH F\DQROLFKHQV D VLJQLILFDQWO\ KLJKHU SURSRUWLRQ WKDQ WKDWIRXQG RQ PRVW RWKHU URFN W\SHV /DUJH WKDOOL RI Collema DQG Leptogium DUH FRPPRQ RQ KDUG OLPHVWRQHV ZKHUHWKHUHLVSOHQW\ RIPRLVWXUH VXFK DV RQVWHHS VORSHVIDFLQJDZD\IURPWKHVXQ ,QVXUIDFH GHSUHVVLRQV ZKHUH ZDWHU IRUPV HSKHPHUDO SRROV WKH F\DQROLFKHQV Lempholemma botryosum DQG Leptogium schraderi RIWHQ JURZ ZLWK IUHHOLYLQJ Nostoc 0F&DUWK\ ,Q PRUH [HULF HQYLURQPHQWV PDFUROLFKHQV WHQG WR EH UHSODFHG E\ PLFUROLFKHQV ZLWK JUHHQ DOJDO SKRWRELRQWV 2QH PLJKW H[SHFW VXFK VLWHV WR EH IDYRXUDEOH IRU F\DQREDFWHULD EXW WKHUH DUH VXUSULVLQJO\ IHZ HQFUXVWLQJ F\DQROLFKHQV RQ OLPHVWRQH H[FHSW IRU PHPEHUV RIWKH 3\UHQRSVLGDFHDH DQ XQFRPPRQ DQG SRRUO\ NQRZQ IDPLO\ RI PLFUROLFKHQV DQG PHPEHUV RI WKH JHQXV Pyrenocollema 3XUYLV HW DO 7KH FRPPRQ HQFUXVWLQJ VSHFLHV LQ QRUWKHUQ (XURSH Placynthium nigrum, LV QRW D PHPEHU RIWKLV IDPLO\ DQG LV LQ IDFW FKDUDFWHULVWLF RIGDPS UDWKHU WKDQ GU\ OLPHVWRQHV0F&DUWK\ 7KH SKRWRELRQW RI DOO RI WKHVH OLFKHQV DSDUW IURP Placynthium SRVVLEO\ Dichothrix DQG Pyrenocollema Gloeocapsa LV Nostoc. 7KH Nostoc DSSDUHQWO\ DOZD\V FRQWDLQV REYLRXV KHWHURF\VWV DQG WKXV SUHVXPDEO\ IL[HV QLWURJHQ 7KH F\DQREDFWHULD VHHPWR JDLQ VWUXFWXUDO LQWHJULW\ IURPWKH P\FRELRQW DOORZLQJ WKHP WR FRORQL]H RWKHUZLVH XQIDYRXUDEOH VXUIDFHV )RU H[DPSOH Collema cristatum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
&KDSWHU/LPHVWRQHV
Fig. 3. 7KH IROLRVH F\DQROLFKHQ &ROOHPD FULVWDWXP JURZLQJ RQ KDUG OLPHVWRQH DW DQ XSODQG VLWH LQ 1RUWKHUQ (QJODQG ZKHUH WKH PHDQ DQQXDOWHPSHUDWXUHLV& 7KHOLFKHQH[SDQGVUDGLDOO\DVDVHULHVRIGDUNEDQGV 7KHRXWHUPRVWEDQGOHIWDUURZ KDGQRWSURJUHVVHGGXULQJ WKHSUHYLRXV\UEXWWKHIHUWLOHSDWFKZLWKDSRWKHFLDULJKWDUURZ DQGWKHLQQHUEDQGFHQWUHDUURZ ERWKH[SDQGHGE\ PP\U
& %LRPDVV
,9'HSRVLWLRQ
7KHUH DUH VXUSUL]LQJO\ IHZ TXDQWLWDWLYH GDWD RQ F\DQREDFWHULDO VWDQGLQJ FURS IURP OLPHVWRQH HQYLURQPHQWV EXW D IHZ YDOXHV H[SUHVVHG DV ELRYROXPH SHU XQLW DUHD DUH JLYHQ LQ 7DEOH WR HQFRXUDJH RWKHUV WR UHFRUG VXFK GDWD :LWKLQ WKLV OLPLWHGGDWDVHWWKHKLJKHVWYDOXHRFFXUUHGLQDPDWRI Phormidium ambiguum, ZKHUH WKH F\DQREDFWHULD JUHZ LQ D WKLFN XQFDOFLILHG OD\HU H[SRVHG WR GLUHFW VXQOLJKW ,Q FRQWUDVW WKH HQGROLWKLF DQG WUDYHUWLQH DVVRFLDWHGFRPPXQLWLHV KDG DELRPDVV PRUH WKDQWHQ WLPHV OHVV SUHVXPDEO\ EHFDXVH DFFHVV WR OLJKW DQG SHUKDSV QXWULHQWV ZDV UHVWULFWHG $ VLPLODU ORZ ELRPDVV LV DSSDUHQW LQ WKH XQFDOFLILHG WLQWHQVWULFKH FRPPXQLW\ ZKHUH WKH DOJDH ZHUH LQIOXHQFHG PDUNHGO\E\GHVLFFDWLRQ
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
$ 3HQWHFRVWDQG%$:KLWWRQ
Table 2. ([DPSOHVRIELRPDVVPHDVXUHPHQWVRIF\DQREDFWHULDOFRPPXQLWLHVDVFHOOXODUELRYROXPHSHUXQLWDUHD $ZDWHUEORRPLV LQFOXGHGIRUFRPSDULVRQ
6XEVWUDWXP
/RFDWLRQ
'RPLQDQW 6SHFLHV
0HWHRJHQH WUDYHUWLQH
:DWHUIDOO %HFN (QJODQG
/LPHVWRQH WLQWHQVWULFKH
0DOKDP &RYH (QJODQG
Schizothrix calcicola agg Calothrixparietina Gloeocapsa alpina Gloeocapsa kuetzingiana
(QGROLWKV RQ VXEPHUJHG OLPHVWRQH FREEOHV 0DW LQ ZDUP HIIOXHQW :DWHUEORRP
0DOKDP &RYH (QJODQG
Phormidium favosum Schizothrix perforans 7DII :HOOV :DOHV Phonnidium ambiguum / /XFHUQH 6ZLW]HUODQG Oscillatoria rubescens
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in vitro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
%LRYROXPH
5HIHUHQFH
3HQWHFRVW
3HQWHFRVW
$3XQSXEOGDWD
$3 XQSXEO GDWD =LPPHUPDQQ
PPFP
7KHUH PD\ EH VSHFLILF SURSHUWLHV RI WKH H[WUDFHOOXODU VKHDWK ZKLFK HQFRXUDJH ELR PLQHUDOL]DWLRQ 3HQWHFRVW DQG %DXOG DQG WKH FRORQ\ DUFKLWHFWXUH RIPhormidium incrustatum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
Ca(HCO3)2 = CaCO3 + H2O + CO2 6HYHUDO ZHOO GHILQHG W\SHV RI WUDYHUWLQH RFFXU QRWDEO\ WKH thermogenes, ZKLFK DUH EDVLFDOO\ K\GURWKHUPDO GHSRVLWV DVVRFLDWHG ZLWK KRW VSULQJV DQG WKH meteogenes ZKLFK DUH NDUVW ZDWHU GHSRVLWV
&KDSWHU/LPHVWRQHV
ULSDULDQYHJHWDWLRQ 7ULFKRPHVFRQWLQXHWRJURZXSZDUGVIROORZHG E\UHOHDVHRIKRUPRJRQLD 0LFULWHLVGHSRVLWHG DV DUHVXOWRIERWK &2 HYDVLRQDQGSKRWRV\QWKHVLVIRUPLQJDORRVHVSRQJ\OD\HU Autumn 7ULFKRPHJURZWKVORZV DQGDQRWKHUGHQVHODWHURIFDOFLWH EHJLQVWRIRUPDQGWKLVFRQWLQXHVWKURXJKWKHZLQWHU
IRUPHG IURP FRROHU ZDWHUV 0DQ\ F\DQREDFWHULD ILQGWUDYHUWLQHDVXLWDEOHHQYLURQPHQWIRUJURZWK DQG GHYHORSERWKXSRQDQGZLWKLQWKHGHSRVLWV 7DEOH DOPRVW DOO DFWLYH WUDYHUWLQHV H[SRVHG WR OLJKW DUH FRORQL]HG 7KLV FORVH DVVRFLDWLRQ RI F\DQREDFWHULD ZLWK WUDYHUWLQH KDV EHHQUHFRJQL]HG IRU PRUH WKDQ D FHQWXU\3LD
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ig. 4. 'HYHORSPHQW RI D ODPLQDWH Phormidium incrustaturn WUDYHUWLQH RYHU WKH FRXUVH RI RQH \HDU LQ DQ XSODQG VWUHDP LQ (QJODQG &DOFLWHFU\VWDOVDUHVKRZQLQEODFN Winter 7KH WULFKRPHV DUH RULHQWHG PRUH RU OHVV SHUSHQGLFXODU WR WKH GHSRVLW VXUIDFH DQG JURZ OLWWOH RU QRW DWDOO ZKLOH WHPSHUDWXUH DQGLUUDGLDQFHLVORZ &DOFLXPFDUERQDWH FRQWLQXHV WR SUHFLSLWDWH DURXQG WKHWULFKRPHV DV DUHVXOW RI &2 HYDVLRQ IRUPLQJ DGHQVH PLFURFU\VWDOOLQHOD\HU Spring 7KH WULFKRPHV EHJLQ WR JURZ XSZDUGV DQG DOVR SRVVLEO\ GRZQZDUGV DQG VPDOO FU\VWDOV RIFDOFLWH PLFULWH DOVR IRUP $W WKLVWLPHFKLURQRPLGODUYDHRIWHQEXLOGWXQQHOVLQWKHVXUIDFHOD\HU DQGIHHGRQWKHIUHVKF\DQREDFWHULDOFURS Summer 7KLV LV XVXDOO\ WKH SHULRG RIPD[LPXP JURZWK ZLWK WKH KLJKHUWHPSHUDWXUHVEXWJURZWKPD\EHUHGXFHGGXHWRVKDGLQJE\
& 0HWHRJHQH7UDYHUWLQHV 7KH FDOFLXP ELFDUERQDWH LQ PHWHRJHQH WUDYHUWLQHV IRUPV DV D UHVXOW RI VRLOJHQHUDWHG FDUERQ GLR[LGH UHDFWLQJZLWK VXEVXUIDFHOLPHVWRQHV 7KH FRPPRQHVW F\DQREDFWHULXP RI WKHVH WUDYHUWLQHV DSSHDUV WR EH Phormidium incrustatum, ZKLFK LV ZLGHO\ GLVWULEXWHG LQ (XURSH DQG 1RUWK $PHULFD 3HQWHFRVW E +RZHYHU DOWKRXJK WKLV DQG RWKHU Phormidium VSS DUH ZLGHVSUHDG DQG VRPHWLPHV DEXQGDQW WKHUH LV DOVR RIWHQ D GLYHUVH IORUD RIRWKHU F\DQREDFWHULD )RU LQVWDQFH RI WKH QRQGLDWRP
WD[DIRXQGLQWZRVPDOOIDVWIORZLQJULYHUVLQ$XVWULD WKH FDOFDUHRXV ,VDU DQG SDUWLDOO\ FDOFDUHRXV *VFKQLW]EDFK ZHUHF\DQREDFWHULDDQG RIWKHVH EHORQJHGWRChamaesiphon). :DWHU UHODWLRQV DUH NQRZQ WR LQIOXHQFH WKH GLVWULEXWLRQ RI F\DQREDFWHULD LQ WUDYHUWLQHGHSRVLWLQJ VWUHDPV3HQWHFRVW E ZLWKFRFFRLGIRUPV (Aphanocapsa, Gloeocapsa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vice versa? 7KH PHWHRJHQHWUDYHUWLQHV RI(XURSH $3 XQSXEOLVKHG GDWD DFFUHWH DW PP \U ZKLFK ZRXOG DSSHDU WR EH VXIILFLHQWO\ VORZ WR DOORZ WKH F\DQREDFWHULD WR NHHS SDFH ,Q ODERUDWRU\ FXOWXUHV WULFKRPHV RIP. incrustatum FDQJURZ PP LQ D IHZ GD\V VR JURZWK VKRXOG QRW EH OLPLWHG E\ FDUERQDWH GHSRVLWLRQ +RZHYHU JURZWK UDWHV RI FDOFLILHGFRORQLHV PHDVXUHG LQWKHILHOG DUHVLPLODUWR WKRVHRIWKHWUDYHUWLQHVWKHPVHOYHV ,Q YLHZ RI LWV LPSRUWDQFH WKH JURZWK RI Phormidium incrustatum ZLOO EH GHVFULEHG LQ VRPH GHWDLO 7KH VSHFLHV IRUPV GDUN JUHHQ VWUDWD RIWHQ FRPSOHWHO\ FRYHULQJ WKH GHSRVLW VXUIDFH LQ VKDG\ VLWXDWLRQV 7KH WULFKRPHV ZKLFK DUH P LQ GLDPHWHU XVXDOO\ RFFXU LQ IDVFLFOHV DQG RIWHQ IRUP VOLJKWO\ UDGLDWLQJ EXVKOLNH VWUXFWXUHV XS WR PP LQ GLDPHWHU )UH\WHW DQG 3OD]LDW 7KH IDVFLFOHV DUH XVXDOO\ FURVVFXW ZLWK DOWHUQDWLQJ OLJKW DQG GDUNODPLQDWLRQV FDXVHG E\GLIIHUHQFHV LQ FDOFLWH FU\VWDO VL]H )LJ DQG Phormidium ELRPDVV 'XULQJ WKH VSULQJ DQG VXPPHU ZKHQ Phormidium JURZWK LV UDSLG VPDOO FU\VWDOV RI PLFULWH FDOFLWH FU\VWDOV P GLDPHWHU DUH GHSRVLWHG DPRQJ WKH WULFKRPHV SURYLGLQJ D SRURXV VSRQJHOLNH VWUXFWXUH 'XULQJ ZLQWHU ZKHQ JURZWK LV PLQLPDO D PRUH GHQVH PDFURFU\VWDOOLQH OD\HU LV IRUPHG SURYLGLQJ D WKLQQHU OLJKWHU FRORXUHG ODPLQD ZKHQ YLHZHG LQ UHIOHFWHG OLJKW ,Q (XURSH DQG 1RUWK $PHULFD WKH FRXSOHWV DYHUDJH PP LQWKLFNQHVV DQG KDYHEHHQ XVHGWRPHDVXUHWUDYHUWLQHGHSRVLWLRQUDWHV 3HQWHFRVW DQG9LOHV LQSUHVV 'XULQJ WKH ZDUPHUPRQWKV WKH WULFKRPHV RI Phormidium ULVH DERYH WKH WUDYHUWLQH VXUIDFH *UQLQJHU DQG DUH SUHVXPDEO\ ORVWWR
$ 3HQWHFRVWDQG %$:KLWWRQ
Fig 5. &DOFLILHG FRORQLHV RI 'LFKRWKUL[ FDOFDUHD IURP D VKDOORZ OLPHVWRQHSRRO7KRUDJLOO1RUWK
Fig. 6. 6HFWLRQ WKURXJK D FRORQ\ RI Rivularia haematites :DWHUIDOO%HFN1RUWK
JUD]HUV DEUDVLRQ DQG LI QXWULHQW FRQGLWLRQV DUH IDYRXUDEOH E\ KRUPRJRQLXP SURGXFWLRQ 7KLV VSHFLHV LV IUHTXHQWO\ DVVRFLDWHG ZLWK D VSHFLDOLVW FKLURQRPLG IDXQD 7KLHQHPDQQ ZKLFK DORQJ ZLWKQHPDWRGHVJUD]HVWKHWULFKRPHV 6HYHUDO PRUSKRORJLFDOO\ VLPLODU Phormidium VSHFLHV DUH IUHTXHQWO\ IRXQG DVVRFLDWHG ZLWK P. incrustatum: P. martensiana DQG P. valderianum. 7KH\ DUH GLVWLQJXLVKHG ODUJHO\ E\ WULFKRPH GLDPHWHU ZKLFK GLIIHUV VWDWLVWLFDOO\ .DQQ DWWHPSWHG D UHYLVLRQ RI WKH P. incrustatum FRPSOH[ DQG
&KDSWHU/LPHVWRQHV FRQFOXGHG WKDW WKHUH ZHUH RQO\ WZR VSHFLHV P. incrustatum DQG P. calcicola, WKHODWWHUFKDUDFWHULVHG E\ WKH SUHVHQFH RI D FDO\SWUD EXW WKH WD[RQRPLF VWDWXV RIWKLV FKDUDFWHUKDV \HW WR EH YHULILHG 2WKHU IRUPV RI Phormidium DUH NQRZQ IURP WKHUPRJHQH WUDYHUWLQHV ZKHUH WKH\ FRPPRQO\ WKULYH DW ZDWHU WHPSHUDWXUHV RI & P. laminosum LV D ZLGHO\ GLVWULEXWHGVSHFLHV IRUPLQJVPRRWKFRULDFHRXV VWUDWD ZKHUH WKH WUDYHUWLQH LV QRW GHSRVLWHG WRR UDSLGO\ 3HQWHFRVWHWDO &\DQREDFWHULDZLWKVKHDWKVFRQWDLQLQJWZRRUPRUH WULFKRPHV FDQ EH DEXQGDQW RQ WUDYHUWLQHV HJ *ROXELF 6]XOF DQG 6P\N HVSHFLDOO\ WKRVH RFFXUULQJ DW ODNH HGJHV DQG H[SRVHG DUHDV SURQHWR GHVLFFDWLRQ.DQQ +HUH Schizothrix VSHFLHVRIWHQSUHGRPLQDWH ZKLFKEXLOG\HOORZEURZQ FDOFLILHG VKHHWV S. calcicola DQG S. fasciculata RU QRGXOHVS. fasciculata IR semiglobosa, S. pulvinata WKH ODWWHU RIWHQ DVVRFLDWHG ZLWK QRGXOHV RI Homoeothrix crustacea 3HQWHFRVW ,Q WKH QRGXODU IRUPV S. fasciculata, 6 pulvinata WKH WULFKRPHV IRUP UDGLDWLQJ EXVKOLNH VWUXFWXUH D IHZ PLOOLPHWUHV LQ GLDPHWHU 7KHVH EHORQJ WR WKH VXEJHQXV,QDFWLVDQGDUHRIWHQKHDYLO\HQFUXVWHGZLWK FDOFLWH 7KH\DUHSDUWLFXODUO\ FRPPRQ RQWUDYHUWLQHV H[SRVHG WR GULSSLQJ RU IORZLQJ ZDWHU RU WKRVH IRUPLQJDWWKHHGJHVRIODNHV ZKHUHWKH\PD\SHUVLVW
IRU PDQ\ \HDUV 6SHFLHV IRUPLQJ OHDWKHU\ FUXVWV S. coriacea, S. lateritia DUH RIWHQ IRXQG SHHOLQJ DZD\ IURPURFNVXUIDFHV DQG DUH XVXDOO\ FRQILQHG WR DUHDV RI LQWHUPLWWHQW IORZ 7KHVH EHORQJ WR WKH VXEJHQXV +\SKHRWKUL[ DQG UDUHO\ SHUVLVW +RZHYHU FXUUHQW FRQFHSWV RI WKLV JHQXV DUH GDWHG DQG DZDLW FODULILFDWLRQ &RQFUHWLRQDU\ IRUPV DUH DOVR HQFRXQWHUHG LQ RWKHU VSHFLHV RI Homoeothrix, EXW PRUH HVSHFLDOO\ LQ Dichothrix )LJ DQG Rivularia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
Fig. 7. 7UDYHUWLQH GHSRVLWLRQ DVVRFLDWHG ZLWK F\DQREDFWHULDO FRPPXQLW\ Dichothrix, Scytonema myochrous LQ WULFNOHV DGMDFHQW WR $O 7DQXPDKZDWHUIDOOQHDU$EKD6DXGL$UDELD:KLWWRQHWDO
WKH F\DQREDFWHULD DUH RIWHQ DVVRFLDWHG ZLWK EU\RSK\WHV IRUPLQJ VLPLODU VWUXFWXUHV :DQJ HW DO 7KHUH DUH WZR SRVVLEOH UHDVRQV IRU JUHDWHU SUHFLSLWDWLRQ XQGHU KLJKHU LUUDGLDQFH ELRJHQLF SUHFLSLWDWLRQ DVVRFLDWHG ZLWKSKRWRV\QWKHWLF XSWDNHRI &2 RU JUHDWHU HYDSRUDWLYH ORVV DQG KHQFH PRUH GHSRVLWLRQ 6KHDWK SLJPHQWV ZRXOG HQKDQFH UDGLDWLRQDEVRUSWLRQDQGKHQFHHYDSRUDWLRQ
' 2QFRLGV 2QFRLGV FRQVLVWLQJ RI URXQGHG DQG FRQFHQWULFDOO\ ODPLQDWHG WUDYHUWLQH SHEEOHV DUH ZLGHVSUHDG LQ FDOFDUHRXV VWUHDPV DQG FRQWDLQ D ZLGH UDQJH RI HQFUXVWLQJ F\DQREDFWHULD 0RVW RQFRLGV SRVVHVV D FUXVW RI ILODPHQWRXV VSHFLHV EHORQJLQJ WR WKH JHQHUD Calothrix, Dichothrix, Homoeothrix, Phormidium, Rivularia RU Schizothrix 6FKDIHU DQG 6WDI 3HQWHFRVW 5RWW %HFDXVH WKH GHSRVLWV DUH FRQWLQXRXVO\ PRYHG E\ ZDWHU FXUUHQWV WKH\ SURJUHVV VORZO\ GRZQVWUHDP HQODUJLQJ DV WKH JR 7KH\ SURYLGH DQ XQXVXDO HFRORJLFDO QLFKH ZKHUH SHULRGV RI SDUWLDO EXULDO RVFLOODWH ZLWK SHULRGV RI FDUERQDWH GHSRVLWLRQ $W RQH VLWH LQ QRUWKHUQ (QJODQG 3HQWHFRVW JURZWK VWXGLHV KDYH VKRZQ WKDW RQFRLGWUDYHUWLQHDFFXPXODWLRQLV VHDVRQDO DQG FRUUHODWHG ZLWK ZDWHU WHPSHUDWXUH 7KH WUDYHUWLQH ZDV FRORQL]HG WKURXJKRXW WKH \HDU E\ Calothrix parietina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omoeothrix crustacea DQG Schizothrix fasciculata DQG WKH UHPDLQLQJ LQQHUSDUWRI WKHRQFRLGV Rivularia 7KLV PD\ UHIOHFW HQYLURQPHQWDO FKDQJHV GXULQJ WKH SHULRG RI RQFRLG GHYHORSPHQW VXFKDV DSRVVLEOH LQFUHDVHLQ DTXHRXV FRPELQHG QLWURJHQ
$ 3HQWHFRVWDQG%$:KLWWRQ 9, &RORQL]DWLRQ 6XFFHVVLRQ DQG :HDWKHULQJ LQ7HUUHVWULDO DQG )UHVKZDWHU (QYLURQPHQWV &\DQREDFWHULD DUH DPRQJ WKH ILUVW SKRWRWURSKV WR FRORQL]H FOHDQ OLPHVWRQH VXUIDFHV )UHVKO\ H[SRVHG WUDYHUWLQHV DQG VXEVWUDWD VXFK DV ,FHODQG VSDU FDOFLWH VRRQ DFTXLUH D IHOW RI QDUURZ ILODPHQWRXV F\DQREDFWHULD HVSHFLDOO\ Phormidium DQG Schizothrix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loeocapsa WRRN VHYHUDO\HDUV WRFRORQL]HFDOFDUHRXV ZDOOV LQ&DQDGD $W 0DOKDP &RYH (QJODQG DQ P KLJK VRXWK IDFLQJ OLPHVWRQH FOLII FRFFRLG F\DQREDFWHULD IRUP GDUN WLQWHQVWULFKH UDUHO\ H[SRVHG WR ZDWHU 2QH VWUHDNZDVFOHDQHG RILWVF\DQREDFWHULD Chlorogloea microcystoides, Gloeocapsa VSS DQG DIWHUWHQ \HDUV RI REVHUYDWLRQ WKH ELRPDVV ZDV EDUHO\ RI DQ DGMDFHQW XQFOHDUHG FRQWURO DUHD ,Q WKH FDVH RI D WLQWHQVWULFKH UHFHLYLQJ UHJXODU ZHWWLQJ DSSUR[ GD\V SHU \HDU WKH ELRPDVV RI D Schizothrix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chizothrix WLQWHQVWULFKH QRWHG DERYH
&KDSWHU/LPHVWRQHV
Fig. 8. &RORQL]DWLRQ DQG PLFULWL]DWLRQ RI D GHQVH FU\VWDOOLQH OLPHVWRQH DV REVHUYHG LQ D VPDOO KLOO VWUHDP :DWHUIDOO %HFN (QJODQG 7KHZKROHSURFHVVWDNHVDERXWILYH\HDUVLQWKLVXSODQG VWUHDP ZKRVHPHDQZDWHUWHPSHUDWXUHLV RQO\DERXW& &DOFLWH FU\VWDOVDUHVKRZQLQEODFN $ 2Q VXEPHUVLRQ WKH OLPHVWRQH DFTXLUHV D WKLQ DXIZXFKV FRQWDLQLQJPXFLODJLQRXVF\DQREDFWHULDDQGGLDWRPV % Schizothrix VSHFLHV EHJLQ WR HWFK WKH FDOFLWH FU\VWDOV XVXDOO\ IROORZLQJOLQHVRIZHDNQHVVVXFKDVFU\VWDOERXQGDULHV & )XUWKHUFRORQL]DWLRQOHDGVWRGHHSHUSHQHWUDWLRQ ' ,QWHQVHFRORQL]DWLRQUHYHDOV WKHF\DQREDFWHULDIRUPLQJDFULVV FURVV SDWWHUQJXLGHGE\ FU\VWDOSODQHV ZHDNHQLQJWKHPLQHUDO DQG HYHQWXDOO\EUHDNLQJLWXSLQWRPLFULWH
ZKHUH DOOEXWRQHVSHFLHV Stigonema ocellatum KDG UHFRORQL]HG DIWHU D \HDU 6LQFH WLQWHQVWULFKH FRPPXQLWLHV UDUHO\ SRVVHVV PXFK VWUXFWXUH DQG ]RQDWLRQ UDSLG VWDELOLVDWLRQ LV QRW XQH[SHFWHG +RZHYHU 7XULDQ UHSRUWHG DQ H[DPSOH RI VXFFHVVLRQ EHJLQQLQJ ZLWK Gloeocapsa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
PDUNHG H[SDQVLRQ RQ ZHWWLQJ DQG DUH DOVR OLNHO\ WR DVVLVW LQ SDUWLFOH GHWDFKPHQW RQ OLPHVWRQH ,Q DGGLWLRQ WR PHFKDQLFDO DFWLRQ FKHPLFDO HWFKLQJ LV DOVR OLNHO\ WR EH VLJQLILFDQW 'DQLQ HW DO IRXQGWKDWOLPHVWRQHDQGGRORPLWHVXUIDFHVLQSDUWVRI ,VUDHO ZHUH FRYHUHG ZLWK SLWV FRQWDLQLQJ F\DQREDFWHULD 7KH SLWV ZHUH PP GHHS DQG FRORQL]HGE\Gloeocapsa VSHFLHVDQGOLFKHQV ,WZDV VXJJHVWHG WKDW WKH SLWV VKLHOGHG WKH DOJDH IURP VRODU UDGLDWLRQ DQG LW LV OLNHO\ WKDW WKH\ ZHUH IRUPHG E\ GLVVROXWLRQ 9LOHV DOVR REVHUYHG D OLPHVWRQH
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
9,, ,QIOXHQFHRI7HPSRUDO&KDQJHVLQ 1XWULHQWV $ 6HDVRQDO 7KH GRPLQDQW F\DQREDFWHULD RI DTXDWLF WUDYHUWLQH GHSRVLWLQJ VLWHV RIWHQ VKRZ PDUNHG VHDVRQDOFKDQJHV LQ JURZWK DQG FDOFLWH GHSRVLWLRQ 7KLV LV HVSHFLDOO\ REYLRXV LQ VSHFLHV IRUPLQJ PXOWLFHOOXODU KDLUV 6WXGLHV RQ VPDOO VWUHDPV GRPLQDWHG E\ Rivularia LQ QRUWKHUQ(QJODQG 3ODWHD DQG LVRODWHV IURPWKHVH VWUHDPV SURYLGH LQVLJKW LQWR ZKDW PD\ EH PRUH JHQHUDO SKHQRPHQD LQ OLPHVWRQH VWUHDPV 7KH PRVW GHWDLOHG VWXGLHV KDYH EHHQ PDGH LQ 8SSHU 7HHVGDOH ZKHUH VWUHDP ZDWHU LV GHULYHG SDUWO\ IURP OLPHVWRQH VSULQJVDQGSDUWO\IURPSHDWGUDLQDJH $ VHDVRQDO VWXG\ RIIRXU VWUHDPV /LYLQJVWRQH DQG :KLWWRQ VKRZHGWKDWSKRVSKDWHFRQFHQWUDWLRQV
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ivularia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ivularia FI R. biasolettiana FRORQLHV LQ 8SSHU 7HHVGDOH FDOFLI\ RQO\ ZHDNO\ DQG UHODWLYHO\ OLWWOH SHUPDQHQW GHSRVLW LV IRUPHG +RZHYHU LW VHHPV OLNHO\ WKDW VHDVRQDO FKDQJHV LQ DTXHRXV 3 LQIOXHQFH JURZWK IHDWXUHV LQ PRUH W\SLFDO WUDYHUWLQH GHSRVLWLQJ VLWHV VXFK DV WKH ODPLQDWLRQV LQ RQFRLGV PHQWLRQHGDERYHDQGWKHFKDUDFWHULVWLFODPLQDWLRQVRI R. haemutites. 0HDVXUHPHQWV RI F\DQREDFWHULDO JURZWK UDWHV DQG QXWULHQW EXGJHWV LQ :DWHUIDOO %HFN (QJODQG DOVR VXJJHVWHG WKDW IRU PRVW RI WKH \HDU QLWUDWH1 LV DGHTXDWH EXW SKRVSKDWH PD\ EHFRPH OLPLWLQJ GXULQJ VXPPHU 3HQWHFRVW 6SLUR DQG 3HQWHFRVW 1RFRORQ\IRUPLQJD[HQLFVWUDLQRI D VWUHDPRivularia KDV \HWEHHQ EURXJKW LQWR FXOWXUH EXW D QXPEHU RI ODERUDWRU\ VWXGLHV KDYH EHHQ PDGH RQ DQ LVRODWH RI QRQFDOFLI\LQJ Calothrix parietina IURPRQHRIWKH8SSHU7HHVGDOHVWUHDPV ZKLFK VKRZ WKDW LWV QLWURJHQDVH DQG SKRVSKDWDVH DFWLYLWLHV
&KDSWHU 9 /LPHVWRQHV UHVSRQG VLPLODUO\WR FKDQJHV LQ3 VWDWXV DV Rivularia /LYLQJVWRQH DQG :KLWWRQ /LYLQJVWRQH HW DO :KLWWRQ *UDLQJHUHWDO &KDQJHV LQ ILHOG SRSXODWLRQV RI & parietina LQ WKHVH VWUHDPV SUREDEO\ UHIOHFW WKH VPDOOHU SKRVSKDWH SHDNV RFFXUULQJ DW YDULRXV WLPHV RI\HDU DQG WKH RUJDQLVP WKHUHIRUH XQGHUJRHV F\FOHV RIPRUSKRORJLFDO FKDQJHV PRUH IUHTXHQWO\ WKDQ RQFH D \HDU 2QFRLGV ZLWK KLJKO\ FDOFLILHG Rivularia FRORQLHV DW VHYHUDO VLWHV LQ QRUWKHUQ(QJODQG HJ 3ODWH F DVVD\HG LQ VXPPHU IRU VXUIDFH SKRVSKDWDVH DOVR VKRZHG KLJK DFWLYLW\ VXJJHVWLQJ PDUNHG 3 OLPLWDWLRQ -0
% /RQJWHUP(IIHFWV (XWURSKLFDWLRQ RI WUDYHUWLQHGHSRVLWLQJ V\VWHPV LV NQRZQWREHGHOHWHULRXV (PLJ QRWHGGDPDJH LQ DQ 2NODKRPD FDWFKPHQW FDXVHG E\ FDWWOH JUD]LQJ DQG LQ (QJODQG (GZDUGV DQG +H\ZRRG IRXQG WKDWFDUERQDWHHQFUXVWDWLRQV GLVDSSHDUHG EHORZ D VHZDJH RXWIDOO WKRXJK WKH HXNDU\RWLF DOJDO IORUD UHPDLQHGXQFKDQJHG 6HYHUDO VWUHDPVLWHV DUHNQRZQ ZKHUH Rivularia FRORQLHV ZHUHRQFHSUHVHQW EXWQRZ DEVHQW %$: XQSXEOLVKHG GDWD 3KRVSKDWH LQKLELWLRQ RIFDOFLWH QXFOHDWLRQ LV JHQHUDOO\ UHJDUGHG DV WKH FDXVH 5DLVWULFN &DVDQRYD DQG /DIRQW WKHWUDYHUWLQHFUXVWEHLQJORVWWRJHWKHUZLWKLWV DVVRFLDWHG FRPPXQLW\ +RZHYHU HQKDQFHG SKRVSKDWH ZRXOG DOVR EH H[SHFWHG WR HOLPLQDWH RUJDQLVPV DGDSWHG WR KLJKO\ YDULDEOH OHYHOV RI DPELHQW3VXFKDVWKHKDLUIRUPLQJJHQHUDCalothrix, Dichothrix DQG Rivularia :KLWWRQ 7KH GLVDSSHDUDQFH RIRQFRLGV LQ /LWWOH &RQHVWRJD &UHHN 86$ ZKLFK IRUPHG DURXQG FRORQLHV RI P. incrustatum ZDV DWWULEXWHG WR SROOXWLRQ E\ DFLGLF LQGXVWULDO HIIOXHQWV *ROXEL& *ROXELF DQG )LVFKHU 7KH H[WHQWRISROOXWLRQ OLNHO\WR OHDGWR FKDQJHV LQ WUDYHUWLQHGHSRVLWLRQ VHHPVWRYDU\EHWZHHQ VLWHV ,Q VRXWKHUQ (QJODQG IRU LQVWDQFH PRVW WUDYHUWLQH GHSRVLWLQJ VWUHDPV DUH VXEMHFW WR QXWULHQW HQULFKPHQW
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ivularia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hormidium crossbyanum, SHUVLVWHG LQ D QRQ FDOFLILHG VWDWH +RZHYHU LW PLJKW HTXDOO\ ZHOO EH DUJXHG WKDW WKH DUULYDO RI QHZ FDOFLILHG VSHFLHV UHIOHFWHG WKH IDFW WKDW WKHUH KDG EHHQ D VKLIW LQ WKH HQYLURQPHQW
9,,, &RQFOXGLQJ &RPPHQWV 7KHUH LV D FRQVLGHUDEOH OLWHUDWXUH RQ GHSRVLWLRQ DQG HURVLRQ RI OLPHVWRQH DW WHUUHVWULDO IUHVKZDWHU DQG PDULQHVLWHV ,Q WKH FDVH RIIUHVKZDWHUV LW LV VWLOO IDU IURPFOHDU ZK\ GHSRVLWLRQ RFFXUV DWRQH VLWH DQG QRW DQRWKHU ZKLFK DWOHDVWVXSHUILFLDOO\ PD\ VHHPTXLWH VLPLODU 0RUH LQIRUPDWLRQLV QHHGHG RQ WKHH[WHQWWR ZKLFK RWKHUIDFWRUV LQIOXHQFHWKHLQKLELWRU\ HIIHFWV RI SKRVSKDWH RQ GHSRVLWLRQ 7KH VDPH DSSOLHV WR LQGLYLGXDO VSHFLHV VXFK DVPleurocapsa minor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
$FNQRZOHGJHPHQWV 7KDQNVWR'U+HDWKHU$9LOHVIRUXVHIXOVXJJHVWLRQV
5HIHUHQFHV $OOHQ 7)0 0XOWLYDULDWH DSSURDFKHV WR WKH HFRORJ\ RI DOJDH RQ WHUUHVWULDO URFN VXUIDFHV LQ 1RUWK :DOHV - (FRO $O7KXNDLU $$ DQG *ROXELF 6 1HZ HQGROLWKLF F\DQREDFWHULD IURP WKH $UDELDQ *XOI Hyella immanis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air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pHVHQFURXWDQWVGX 9DU)UDQFH 9HUK,QW 9HUHLQ/LPQRO &DVWHQKRO]5: 6XEVHFWLRQ ,,, 2UGHU2VFLOODWRULDOHV ,Q %HUJH\ V0DQXDORI6\VWHPDWLF%DFWHULRORJ\9RO :LOOLDPVDQG:LONLQV%DOWLPRUH0DU\ODQG &DVWHQKRO] 5: DQG :DWHUEXU\ -% 7D[D RI WKH &\DQREDFWHULD ,Q %HUJH\ V 0DQXDO RI6\VWHPDWLF %DFWHULRORJ\
$ 3HQWHFRVWDQG %$ :KLWWRQ 9ROXPH SS :LOOLDPV DQG :LONLQV %DOWLPRUH 0DU\ODQG &DXGZHOO & (WXGH H[SHULPHQWDOH GH OD IRUPDWLRQ GH PLFULWH HW GH VSDULWH GDQV OHV VWURPDWROLWHV G HDX j Rivularia. %XOO6RF*HRO )U &RSHODQG--
&KDSWHU /LPHVWRQHV )MHUGLQJVWDG( 7KHDOJDOIORUDRIVRPH 7LQWHQVWULFKH LQWKH
$OSHV0DULWLPHV )UDQFH 6FKZHL]=+\GURO )UH\WHW 3 DQG 3OD]LHW $ /HV IRUPDWLRQV VWURPDWROLWLTXHV WXIV FDOFDLUHV UHFHQWHV GH OD UHJLRQ GH 7RXUQXV 6D{QH HW /RLUH *HRELRV )UH\WHW 3 DQG 3OD]LHW $ 0RGHP IUHVKZDWHU PLFURELDO FDUERQDWHV WKH 3KRQQLGLXP VWURPDWROLWHV WXIDWUDYHUWLQH RI 6RXWKHDVWHUQ%XUJXQG\3DULV%DVLQ )UDQFH )DFLHV )ULHGPDQQ ( , 7KH JHQXV *HLWOHULD GLVWULEXWLRQ RI * calcarea DQG*IORULGDQDQVS 3ODQW6\VW(YRO 131: 169-178. )ULWVFK )( 7KH OLPHHQFUXVWHG 3KRQQLGLXPFRPPXQLW\ RI %ULWLVKVWUHDPV 9HUK,QW9HUHLQ/LPQRO )ULWVFK )( 3KRUPLGLXP LQFUXVWDWXP 1DHJ *RP DQ LPSRUWDQWPHPEHURIWKHOLPHHQFUXVWHG FRPPXQLWLHV RIIORZLQW ZDWHU %LRORJLVFK-DDUERHN *DUFLD3LFKHO ) DQG &DVWHQKRO] 5: &KDUDFWHULVDWLRQ DQG ELRORJLFDO LPSOLFDWLRQV RIVF\WRQHPLQ a cyanobacterium sheath pigment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lOOH $EK.DUVWX+EKOHQNXQGH( +HDWK &5 /HDGEHDWHU %&6 &DOORZ 0( (IIHFWV RI LQKLELWRUV RQ FDOFLXP FDUERQDWH GHSRVLWLRQ PHGLDWHG E\ IUHVKZDWHUDOJDH -$SSO3K\FRO +HUPDQ -6 DQG /RUDK 00 &2 RXWJDVVLQJ DQG FDOFLWH SUHFLSLWDWLRQ LQ )DOOLQJ 6SULQJ &UHHN 9LUJLQLD 86$ &KHP *HRO +ROPHV 17+DQG:KLWWRQ%$ 3K\WREHQWKRV RIWKH 5LYHU 7HHVDQGLWVWULEXWDULHV )UHVKZDW%LRO -DDJ 2 8QWHUVXFKXQJHQXEHUGLH9HJHWDWLRQXQG %LRORJLH GHU $OJHQ GHV QlFNWHQ *HVWHLQV LQ GHQ $OSHQ LP -XUD XQG LP VFKZHL]HULVFKHQ 0LWWHODQG %HLWU .U\SWRJDPHQ )ORUD 6FKZHL] .DQQ ( 'LH HXOLWRUDO $OJHQ]RQH LP 7UDXQVHH $UFK +\GURELRO .DQQ( %HPHUNXQJHQ]XU6\VWHPDWLNXQGgNRORJLHHLQLJHU PLW .DON LQNUXVWLHUWHU 3KRQQLGLXP $UWHQ 6FKZHL] = +\GURO
.HPSH 6 .D]PLHUF]DN- /DQGPDQQ* .RQXN7 5HLPHU$ DQG /LSS$ /DUJHVWNQRZQPLFURELDOLWHV GLVFRYHUHG LQ/DNH 9DQ7XUNH\ 1DWXUH .UXPEHLQ :( DQG 3RWWV 0 *LUYDQHOODOLNH VWUXFWXUHV IRUPHG E\ 3OHFWRQHPD JORHRSKLOXP %RU]L &\DQRSK\WD IURP WKH%RUUHJRGHVHUWLQ6 &DOLIRUQLD *HRPLFURELRO - /H%ULV 6 /H &DPSLRQ$OVXPDUG 7 DQG 5RPDQR -& &KDUDFWHULVWLFV RI HSLOLWKLF DQG HQGROLWKLF DOJDO WXUI H[SRVHG WR GLIIHUHQW OHYHOV RIELRHURVLRQ LQ )UHQFK 3RO\QHVLDQ FRUDO UHHIV 2FHDQRORJLFD$FWD /H &DPSLRQ$OVXPDUG 7 Hyella WD[D HQGROLWKLF F\DQRSK\WHV IURP WURSLFDO HQYLURQPHQWV /L]DUG ,VODQG *UHDW %DUULHU5HHI $UFK+\GURELRO /H &DPSLRQ$OVXPDUG 7 *ROXELF 6 DQG +XWFKLQJV 3 0LFURELDOHQGROLWKVLQVNHOHWRQV RIOLYHDQGGHDG FRUDOV 3RULWD lobata 0RRUHD )UHQFK 3RO\QHVLD 0DU (FRO 3URJU 6HU /HFOHUF - & &RXWp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alothrix parietina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¶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jQGH] HGV 3KRVSKDWDVHVLQWKH(QYLURQPHQW .OXZHU'RUGUHFKW 2EHQOXQHVFKORVV - %LRORJLH XQG gNRORJLH YRQ GUHL UH]HQWHQ 6VVZDVVHU5LYXODULDHQ &\DQREDNWHULHQ hEHUWUDJEDUNHLWDUWVSH]LILVFKHU9HUNDONODQJVWUXFWXUHQ DXIIRVVLOH )RUPHQ *|WWLQJHU$UE*HRO3DODHRQW
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lyengariella endolithica VS QRYD D FDUERQDWH ERULQJ 6WLJRQHPDWDOHDQ F\DQREDFWHULXP IURP D ZDUP VSULQJIHG ODNH QDWXUH WR FXOWXUH $UFK +\GURELRO 6PLWK 5- DQG :LONLQV $ $ FRUUHODWLRQ EHWZHHQ LQWUDFHOOXODU FDOFLXP DQG LQFLGHQW UDGLDWLRQ LQ 1RVWRF 1HZ3K\WRO 6RPHUV*)DQG%URZQ0 7KHDIILQLW\RIWULFKRPHVRIEOXH JUHHQDOJDHIRUFDOFLXPLRQV (VWXDULHV 6SLUR%DQG3HQWHFRVW$ 2QHGD\LQWKHOLIHRIDVWUHDP D GLXUQDOLQRUJDQLFFDUERQPDVVEDODQFHIRUDWUDYHUWLQHGHSRVLWLQJ VWUHDP:DWHUIDOO%HFN
&KDSWHU /LPHVWRQHV 7LDQ < DQG +H ) $ SUHOLPLQDU\ VWXG\ RQ WKH PLFURFRPPXQLW\ HFRORJ\ RI NDUVW &\DQRSK\WD LQ PDW IURP )HQGRQJ&DYH 3DQ[LDQ *XL]KRX &DVDORJLFD6LQLFD 7XFNHU 0( DQG :ULJKW 93 &DUERQDWH 6HGLPHQWRORJ\ %ODFNZHOO/RQGRQ 7XULDQ * &\DQRSK\FpH ³G¶RUpH´ Chrysocapsa SLRQQLpUH GHODFRORQL]DWLRQEXEHWRQ 6DXVVXUHD 8VGRZVNL(+RHIV-DQG0HQVFKHO* 5HODWLRQVKLSEHWZHHQ & DQG O2 IUDFWLRQDWLRQ DQG FKDQJHV LQ PDMRU HOHPHQW FRPSRVLWLRQ LQ D UHFHQW FDOFLWHGHSRVLWLQJ VWUHDP$ PRGHO RI FKHPLFDO YDULDWLRQV ZLWK LQRUJDQLF &D&2 SUHFLSLWDWLRQ (DUWK 3ODQHW6FL/HWW YDQ GHU :DO 3 GH -RQJ / :HVWEURHN 3 GH %UXLMQ :& DQG 0XOGHU6WDSHO$$ 3RO\VDFFKDULGHORFDOL]DWLRQ FRFFROLWK IRUPDWLRQ DQG JROJL G\QDPLFV LQ WKH FRFFROLWKRSKRULG Hymenomonas carterae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three ZHVWHUQ,QGLDQ2FHDQLVODQGV$VWRYH)DUTXKDU DQG6W3LHUUH $WROO5HVHDUFK%XOOHWLQ :KLWWRQ%$ .KRMD70 DQG$ULI,$ :DWHUFKHPLVWU\DQG DOJDOYHJHWDWLRQRIVWUHDPVLQWKH$VLU0RXQWDLQV 6DXGL$UDELD +\GURELRORJLD :KLWWRQ %$
=LPPHUPDQQ 8 2NRORJLVFKH XQG SK\VLRORJLVFKH 8QWXUVXFKXQJHQ DQ GHU SODQNWLVFKHQ %ODXDOJD Oscillatoria rubescens '& XQWHU EHVRQGHUHU %HUFNVLFKWLJXQJ YRQ /LFKW XQG 7HPSHUDWXU 6FKZHL]= +\GURO
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
B . A . Whitton and M. Potts (eds). The Ecology of Cyanobacteria. . © Kluwer Academic Publishers . Printed in the Netherlands.
$ 2UHQ
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icrocoleus chthonoplustes, DFRVPRSROLWDQ PDWEXLOGLQJ F\DQREDFWHULXP IRXQG IURP VHDZDWHU VDOLQLW\ WR VDOLQLWLHV H[FHHGLQJ R DQG WKH XQLFHOOXODU Aphanothece halophytica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scillatoria limnetica, LVRODWHG IURP6RODU/DNH ZDV XVHGDVDODERUDWRU\PRGHOIRUWKHVWXG\RIDQR[\JHQLFSKRWRV\QWKHVLVDQG DQDO\VLVRIWKH SKHQRPHQRQ DWWKH PROHFXODUOHYHOFRPPHQFHG 7R EH DEOH WR ZLWKVWDQG WKH KLJK RVPRWLF SUHVVXUH FDXVHG E\ WKH VDOWFRQFHQWUDWLRQV LQWKHLU VXUURXQGLQJ PHGLXP F\DQREDFWHULDOLYLQJ DWKLJK VDOLQLWLHV SRVVHVV PHFKDQLVPV WRPDLQWDLQRVPRWLFHTXLOLEULXPDQGFHOOWXUJRU ,RQV1D .& FDQWHPSRUDULO\HQWHUWKHFHOOVWRFRXQWHUDFW UDSLG LQFUHDVHV LQ PHGLXP VDOLQLW\ KRZHYHU IRU WKH ORQJ WHUP RUJDQLF VROXWHV DUH DFFXPXODWHG WR SURYLGH RVPRWLF EDODQFH DV H[SHFWHG LQ RUJDQLVPV ZKRVH HQ]\PDWLF PDFKLQHU\ LV LQKLELWHG E\ VDOW 'LIIHUHQW RUJDQLF RVPRO\WHV ZHUH IRXQG LQ F\DQREDFWHULD WKH GLVDFFKDULGHV VXFURVH DQG WUHKDORVH HVSHFLDOO\ LQ WKH OHVV VDOW WROHUDQWW\SHV JOXFRV\OJO\FHUROLQPRGHUDWHO\KDORWROHUDQWVSHFLHV DQGJO\FLQHEHWDLQHLQ$ halophytica DQG D IHZ RWKHU IRUPV WKDW WROHUDWH YHU\ KLJK VDOW FRQFHQWUDWLRQV 2XU XQGHUVWDQGLQJ RI WKH RVPRUHJXODWRU\ PHFKDQLVPV LQF\DQREDFWHULDLQFUHDVHG JUHDWO\ LQUHFHQW\HDUV HVSHFLDOO\ ZLWKUHJDUG WR WKH PROHFXODUDQDO\VLV RI JOXFRV\OJO\FHURO PHWDEROLVP LQ Synechocystis 3&& $FFXPXODWLRQ RI RUJDQLF RVPRWLF VROXWHV KDV LPSRUWDQWLPSOLFDWLRQV IRUWKHFDUERQF\FOH LQ K\SHUVDOLQHHQYLURQPHQWV ZKHUHKDORSKLOLF F\DQREDFWHULDSURYLGH WKHJUHDWHVWFRQWULEXWLRQWRWKHSULPDU\SURGXFWLYLW\
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icrocoleus chthonoplastes, WKH PDLQ PDW EXLOGLQJ F\DQREDFWHULXP LQ WKH PDULQH OLWWRUDO DQG LQWHUWLGDO IODWV FDQ EH IRXQG LQ K\SHUVDOLQH HQYLURQPHQWV ZRUOGZLGH XS WR VDOLQLWLHV RI R DQG KLJKHU 'XELQLQ HW DO D -DYRU ,Q DOO FDVHV D VLQJOH FRVPRSROLWDQIRUPZLWKWKHDELOLW\ WRDGDSWWR D ZLGH UDQJH RI VDOLQLWLHV FKDUDFWHUL]HV WKHVH HQYLURQPHQWV*DUFLD3LFKHOHWDO 7KHVWUDLQ Synechocystis 3&& DQ LVRODWH ZLGHO\ XVHG LQ WKH VWXG\ RI WKH SK\VLRORJ\ RI VDOW DGDSWDWLRQ LQ F\DQREDFWHULD ZDV LVRODWHG IURP D IUHVKZDWHU
&KDSWHU 6DOWVDQG%ULQHV
Table 1. 7HUPV XVHG LQ WKLV FKDSWHU WR GHVFULEH VDOLQLW\ DQG VDOW FRPSRVLWLRQ RIEULQHV DQG WKH UHVSRQVH RIPLFURRUJDQLVPV WR KLJK VDOW FRQFHQWUDWLRQV
6DOLQLW\ 7KDODVVRKDOLQH $WKDODVVRKDOLQH +DORSKLOLF +DORWROHUDQW 6WHQRKDOLQH (XU\KDOLQH 2OLJRKDOLQH 0HVRKDOLQH 3RO\KDOLQH
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
7HUPVGHULYHGIURP*ROXELF
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
,, +\SHUVDOLQH (QYLURQPHQWVDQGWKHLU &\DQREDFWHULDO &RPPXQLWLHV $WWHPSWV WR UHYLHZ WKH RFFXUUHQFH RI F\DQREDFWHULD LQ K\SHUVDOLQH HQYLURQPHQWV DUH KDPSHUHG E\ WKH SRRU VWDWH RI F\DQREDFWHULDO WD[RQRP\ 7KH VSHFLHV FRQFHSW LV QRW ZHOO GHILQHG ZLWKLQ WKH JURXS
(VVHQWLDOO\ DOO FODVVLILFDWLRQ LV EDVHG RQ FHOO RU ILODPHQW PRUSKRORJ\ DQG GLIIHUHQW DXWKRUV RIWHQ XVH GLIIHUHQW GHVLJQDWLRQV IRU ZKDW PD\ EH WKH VDPH PRUSKRORJLFDO W\SH 2Q WKH RWKHU KDQG ZKDW PD\ EHUHIHUUHG WR E\ GLIIHUHQW DXWKRUV DV WKH VDPH JHQXV DQG VSHFLHV PD\ HQFRPSDVV D EURDG UDQJH RI GLIIHUHQW RUJDQLVPV HJ Aphunothece halophytica ,Q WKLV FKDSWHU QR DWWHPSW ZDV PDGH WR DGRSW D XQLIRUP QRPHQFODWXUH V\VWHP DQG PRVW VSHFLHV GHVLJQDWLRQV DUHWKRVH ZKLFK DSSHDUHG LQ WKH RULJLQDO UHSRUWV DV FLWHG LQ WKH OLVW RI UHIHUHQFHV $ QRQ H[KDXVWLYH OLVW RI JHQHUD DQG VSHFLHV WKDW RFFXU DW KLJK VDOW FRQFHQWUDWLRQV DV GHILQHG DERYH LQFOXGHV DV UHYLHZHG LQ %RURZLW]ND *ROXELF 2UHQDQG3RWWV 8QLFHOOXODU W\SHV Aphunothece RU Aphunocupsu URXQG RU RYRLG FHOOV VXUURXQGHG E\ PXFLODJH Synechococcus LGHP ZLWKRXW PXFLODJH QRWH WKDW WKH SUHVHQFH DQG WKLFNQHVV RID VOLPH FDSVXOH FDQ EH KLJKO\ GHSHQGHQW RQ WKH HQYLURQPHQW PDNLQJ LW GLIILFXOW DW EHVW WR GLVFHUQ VRPH VWUDLQV RI Aphunothece IURP Synechococcus; Synechocystis; Gloeothece URGVKDSHG FHOOV GLYLGLQJ LQ D VLQJOH SODQH DQG Gloeocapsa XVXDOO\ VSKHULFDO FHOOV ZKLFK GLYLGH LQ WZR RU WKUHH SODQHV *ROXELF %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ FODVVLILHVURXQG WRFRFFRLG FHOOV ORQJHUWKDQ P DV Cyanothece, VPDOOHU FHOOV DV Synechococcus, DQG FHOOV ZLWK D ZHOOGHILQHG VKHDWK DV Gloeothece :DWHUEXU\ DQG 5LSSND Aphunothece halophytica DOVR GHVLJQDWHG Aphunocupsa halophytica LV DWUXHKDORSKLOH %URFN DQG LV IRXQG ZRUOGZLGH DW VDOLQLWLHV RI DERYH R WR YDOXHV DSSURDFKLQJ KDOLWH VDWXUDWLRQ 7KH ORQJ VSLQGOHVKDSHG Dactylococcopsis salina LV RIWHQ
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phanothece halophytica DQG Schizothrix arenaria (KUOLFK DQG 'RU *HUGHVHWDO
&KDSWHU 6DOWVDQG%ULQHV 7KH YDULHG PLFURELDO FRPPXQLWLHV RI 6RODU /DNH DUHGLVFXVVHGLQ6HFWLRQ,9$
% +\SHUVDOLQH/DNHV &\DQREDFWHULD DUH IRXQG LQ PDQ\ K\SHUVDOLQH ODNHV ERWK ZLWK D WKDODVVRKDOLQH DQG DWKDODVVRKDOLQH VDOW FRPSRVLWLRQ $ QXPEHU RI FKDUDFWHULVWLF H[DPSOHV LQFOXGH
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unaliella VSS F\DQREDFWHULD DUH D FKDUDFWHULVWLF FRPSRQHQW RI WKH ODNH V ELRWD Aphanothece halophytica ZDV IRXQG DW WKH KLJKHVW VDOLQLWLHV ,Q D FODVVLF VWXG\ %URFN UHFRJQL]HG WKH H[LVWHQFH RI KDORSKLOLF F\DQREDFWHULD EDVHG RQ WKH RFFXUUHQFH RIAphanothece LQ WKH *UHDW 6DOW/DNH DQG RQ WKH SURSHUWLHV RIWKH LVRODWHG VWUDLQ LQ FXOWXUH ,Q DGGLWLRQ VSHFLHV RI Phormidium RU Oscillatoria, DV ZHOO DV Microcoleus lyngbyaceus, Spirulina major, DQG Nodularia spumigena ZHUH IRXQGLQWKHVKDOORZVHGLPHQWVRIWKHODNH)HOL[DQG 5XVKIRUWK 3RVW 5XVKIRUWK DQG )HOL[
7KH'HDG6HD 7KH'HDG6HDLVDQDWKDODVVRKDOLQHLQODQGODNHRQWKH ERUGHU EHWZHHQ ,VUDHO DQG -RUGDQ ,WV H[WUHPHO\ VDOLQH ZDWHUV DURXQG J/ WRWDO GLVVROYHG VDOWV KDYHDQXQXVXDOLRQLFFRPSRVLWLRQ 3UHVHQWO\WKHODNH FRQWDLQV PRUH WKDQ 0 0J DQG 0 &D LQ DGGLWLRQ WR DERXW 0 1D 7KH GRPLQDQW DQLRQ LV & IROORZHGE\%U DERXWRIWKHDQLRQIUDFWLRQ 7KH PLFURRUJDQLVPV EHVW DGDSWHG WR WKLV KRVWLOH HQYLURQPHQW DUH VSHFLHV RI KDORSKLOLF $UFKDHD DQG
RQH SULPDU\ SURGXFHU WKH XQLFHOOXODU JUHHQ DOJD Dunaliella pawa. 7KH ELRORJ\ RI WKH 'HDG 6HD LQFOXGLQJ LQIRUPDWLRQ RQ WKH G\QDPLFV RI WKH FRPPXQLWLHVDVLQIOXHQFHGE\FKDQJHVLQWKHSK\VLFDO VWUXFWXUHRIWKH ZDWHUFROXPQ ZDVUHYLHZHGE\2UHQ &\DQREDFWHULD GR QRW IRUP D TXDQWLWDWLYHO\ LPSRUWDQW SDUW RI WKH 'HDG 6HD HFRV\VWHP 7KLV LV H[SHFWHGWRVRPHH[WHQWIURPWKHORZS+RIWKH'HDG 6HD EULQHV a F\DQREDFWHULD DUH QRW QRUPDOO\ DEXQGDQWDWORZS+YDOXHV%URFN +RZHYHU VWUDLQV RIAphanothece, Microcystis, Phormidium DQG Nostoc, DOO GHYHORSLQJ DW VDOWFRQFHQWUDWLRQV RIXS WR J / ZHUH REWDLQHG LQ HQULFKPHQW FXOWXUHV 9ROFDQL 2QH REOLJDWHO\ KDORSKLOLF Aphanocapsa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phanothece halophytica) VKRZHG PDVVLYH GHYHORSPHQW LQ DUWLILFLDO VRODU SRQGV HVWDEOLVKHG DW WKH VKRUH RI WKH 'HDG 6HD 'RU DQG +RUQRII D 'RUDQG3D] VHHDOVR 6HFWLRQ ,,( 5HFHQW DWWHPSWV WR HQULFK IRU SKRWRV\QWKHWLF PLFURRUJDQLVPV XVLQJ 'HDG 6HDVHGLPHQWV IURP P GHSWK DV LQRFXOXP DQG HPSOR\LQJ PHGLD FRQWDLQLQJ WR SDUWV RI 'HDG 6HD ZDWHU JDYHULVH WR WKH GHYHORSPHQW RI XQLFHOOXODU SynechococcusW\SHFHOOV2UHQXQSXEOLVKHGUHVXOWV
6RODU/DNH 6LQDL 6RODU /DNH LV D VPDOO K\SHUVDOLQH KHOLRWKHUPDO ODNH RQWKH VKRUHRIWKH 6LQDL FRDVW ,Q VXPPHU WKH ODNH PD[LPXP GHSWK P LV PL[HG ZLWK D XQLIRUP VDOLQLW\ RI aR 6HHSDJH RI VHDZDWHU IURP WKH *XOI RI $TDED R VDOLQLW\ DQG RFFDVLRQDO UDLQ IORRGV FDXVH WKH IRUPDWLRQ RID OHVV VDOLQH OD\HU DV ORZDVR LQDXWXPQ ZLQWHUDQGWKHODNHUHPDLQV VWUDWLILHG IURP 2FWREHU XQWLO 0D\ 'XULQJ VWUDWLILFDWLRQ D FKHPRFOLQH VHSDUDWHV DQ DQDHURELF K\SROLPQLRQ FRQWDLQLQJ XSWR SSP+6 IURP DQ R[LGL]HGHSLOLPQLRQ 7KHGLIIHUHQFHLQGHQVLW\ RIWKH XSSHUOHVVVDOLQH PRIWKHZDWHUFROXPQDQGWKH GHHS K\SHUVDOLQH EULQHV SUHYHQWV PL[LQJ DQG KHOLRWKHUPDOKHDWLQJFDXVHVWKH K\SROLPQLRQ WRUHDFK
PD[LPDOWHPSHUDWXUHVRIXSWR& LQZLQWHU $ GHWDLOHG DFFRXQW RI WKH OLPQRORJ\ RI WKH ODNH ZDV JLYHQE\&RKHQHWDODD $ ULFK F\DQREDFWHULDO IORUD ZDV IRXQG ERWK LQ WKH ZDWHUFROXPQDQGLQWKHEHQWKLFPLFURELDOPDWV 7KH OLVW RI VSHFLHV IRXQG LQ 6RODU /DNH LQFOXGHG Aphanocapsa concharum, Aphanothece stagnina, A. rnicroscopica, A. halophytica, A. litoralis, A. pallida, Chroococcidiopsis VS Dactylococcopsis salina, D. acicularis, Entophysalis VS Gloeothece VS Johannesbaptista pellucida, Lyngbya aestuarii, L. confewoides, L. digueti, Microcoleus chthonoplastes, Oscillatoria redekei, O. limnetica, O. salina, Phormidium hypersalinum, P. hypolirnneticum, Pleurocapsa VS Pseudanabaena catenata, Spirulina labyrinthiformis, DQG Synechococcus VS &DPSEHOO DQG *ROXELF &RKHQ HW DO D E -¡UJHQVHQHWDO 3RWWV 7KHPL[HGZDWHU FROXPQ GXULQJ WKH VXPPHU PRQWKV FRQWDLQHG Aphanothece FHOOV DW D ORZ GHQVLW\ Aphanothece ZDVDOVRIRXQGLQWKHXSSHUZDWHUOD\HUGXULQJZLQWHU VWUDWLILFDWLRQ $W WKH ERUGHU EHWZHHQ WKH DHURELF HSLOLPQLRQ DQG WKH KRW DQDHURELF K\SROLPQLRQ D SRSXODWLRQ RI Dactylococcopsis salina ZDV IRXQG 3RWWV YDQ5LMQDQG&RKHQ :DOVE\HWDO D. salina DOVR GHVLJQDWHG Myxobactron salinurn E\ 'DYLV DQG *LRUGDQR LV JDV YDFXRODWH ZLWK \HOORZ WR RUDQJH FHOOV DQG JURZV LQ FXOWXUH EHWZHHQ VDOW RSWLPXP DQG XS WR & 7KH KRW VXOILGH FRQWDLQLQJ ZDWHU OD\HU DW WKH ERWWRP RI WKH ODNH FRQWDLQV D GHQVH SRSXODWLRQ RI Oscillatoria limnetica, ZKLFK JURZV DQDHURELFDOO\ XVLQJ VXOILGH DV HOHFWURQ GRQRU LQ DQ DQR[\JHQLF W\SH RISKRWRV\QWKHVLV VHH EHORZ 7KH SURSHU VSHFLHV GHVLJQDWLRQ RIWKLV DERXW P ZLGH ILODPHQWRXV F\DQREDFWHULXP LV XQFHUWDLQ 7KH RULJLQDOGHVFULSWLRQ RIO. limnetica UHODWHV WR DIUHVK ZDWHU VSHFLHV ZLWK D PXFK VPDOOHU ILODPHQW ZLGWK WKDQWKH6RODU/DNHLVRODWH *ROXELF FODVVLILHG WKHRUJDQLVPDVDPhormidium VSHFLHV 'LIIHUHQW W\SHV RIEHQWKLF F\DQREDFWHULDO PDWV FDQ EH IRXQG DW6RODU/DNH 7KH VXUIDFH PDWV GLIIHUHQW PDW PRUSKRORJLHV EHLQJ GHVFULEHG DV VKDOORZ IODW PDW GHHS IODW PDW DQG EOLVWHU PDW DOO DW VDOLQLWLHVDURXQGR DQGWHPSHUDWXUHVEHWZHHQ & ZHUHGRPLQDWHGE\Microcoleus, Phormidiurn, Aphanothece, Aphanocapsa, DQG Synechococcus. 1LWURJHQ IL[DWLRQ DV PHDVXUHG E\ WKH DFHW\OHQH UHGXFWLRQDVVD\WHFKQLTXH ZDVVKRZQWRRFFXULQWKH Microcoleus PDWV RI6RODU /DNH 3RWWV 7KH EURZQUHG VXPPHU PDW ZDV GRPLQDWHG E\ Aphanothece halophytica DQG Aphanothece littoralis &RKHQ D -¡UJHQVHQ HW DO .UXPEHLQ HW
$2UHQ DO 5HYVEHFK HW DO 7KH SHULSKHUDO FUXVW DURXQG WKH SRRO FRQWDLQHG Pseudanabaena, DQG WKH J\SVXP FUXVW FORVH WR ZDWHU ZDV LQKDELWHG E\ Entophysalis 3RWWV Lyngbya aestuarii DSSHDUHG DV GLVFRQWLQXRXV SDWFK\ ILOPV RYHU WKH VXUIDFH RI 6RODU /DNH PDWV 3RWWV $Q DSSUR[LPDWHO\ PP WKLFN JHODWLQRXV SRO\VDFFKDULGHULFK PDW ZDV IRXQG LQ ZLQWHU LQ WKH WKHUPRFOLQH DUHD VDOLQLW\ DERXW R WHPSHUDWXUH DURXQG& 7KLVPDWODFNHGMicrocoleus, DQGZDV ULFK LQ Phormidium, Aphanothece halophytica, Aphanocapsa DQG Pleurocapsa, ZKLFK LPSDUWHG D EULJKW RUDQJH FRORU WR WKH XSSHU PP RI WKH PDW %HORZWKLVGHSWKWKHFRORUFKDQJHGWROLJKWJUHHQGXH WR WKH SUHVHQFH RI Oscillatoria VS DFFRPSDQLHG E\ A. halophytica.
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yanothece, Oscillatoria, DQG Spirulina VSHFLHV %XUNH $ YDULHW\ RI LQODQG ODNHV DUH IRXQG RQ WKH $XVWUDOLDQFRQWLQHQWZLWKVDOLQLWLHVIURP J/ WR DOPRVW J / 'LIIHUHQWW\SHV RIILODPHQWRXV DQG XQLFHOOXODU F\DQREDFWHULD ZHUH IRXQG LQ WKHVH HQYLURQPHQWV%DXOG +RW/DNH:DVKLQJWRQ D VWUDWLILHG ODNH ZLWK J / WRWDO GLVVROYHG VDOWV LQ WKH VXUIDFH OD\HU LQFUHDVLQJWRJ/ QHDUWKHERWWRP ZDVUHSRUWHG WR EH LQKDELWHG E\ Anacystis thermalis, Gomphosphaeria aponina, Oscillatoria chlorina, DQG Plectonema nostocorurn $QGHUVRQ /DNH %RQQH\ LQ WKH 'U\ 9DOOH\V RI$QWDUFWLFD KDV EHQWKLFPDWVGRPLQDWHGE\Lyngbya martensiana DQG Phormidium frigidum, RFFXUULQJ DWDVDOLQLW\ RIDERXW R :KDUWRQHWDO /DNH 9DQGD LQ :ULJKW 9DOOH\ $QWDUFWLFD ZDV UHSRUWHG WR FRQWDLQ FRFFRLG (Synechococcus VS DQG ILODPHQWRXV F\DQREDFWHULD (Phorrnidium W\SH DERXW FHOOV DQG ILODPHQWV /UHVSHFWLYHO\ DW P GHSWKVDOLQLW\DERXWR*ROGPDQHWDO
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
& +\SHUVDOLQH6XOIXU6SULQJV 7KH VSULQJVRI+DPHL0D]RURQ WKH ZHVWHUQ VKRUH RI WKH'HDG 6HDDUHFKDUDFWHUL]HG E\DKLJKVDOWFRQWHQW DURXQG J/ WRWDOGLVVROYHGVDOWV DKLJKVXOILGH FRQWHQW DERXW P0 DQG D S+ RI 7KH WHPSHUDWXUH RIWKH ZDWHU DW WKH VRXUFH LV & 7KH SRROV DQGUXQRIIFKDQQHOV IRUPHGE\WKH VSULQJ ZHUH FRYHUHG E\ D OD\HU RI ILODPHQWRXV F\DQREDFWHULD 2VFLOOXWRULDVS3ODWH D DFFRPSDQLHGE\SDWFKHV RISXUSOH VXOIXUEDFWHULD7KLRFDSVD VS 2UHQ 3ODWHE 7KH PDVV RFFXUUHQFH RIF\DQREDFWHULDLQ WKLVHQYLURQPHQWLV XQXVXDO LQYLHZRIWKHORZS+ RI WKH ZDWHU DV F\DQREDFWHULD DUH UDUHO\ IRXQG DW S+ YDOXHV EHORZ 5.5 WR %URFN %URFN 3KRWRDVVLPLODWLRQ RI &2 E\ WKH F\DQREDFWHULDO FRPPXQLW\ ZDV QRW LQKLELWHG E\ '&08 DQG WKH SUHVHQFH RI VXOIXU JUDQXOHV DVVRFLDWHG ZLWK WKH F\DQREDFWHULDO ILODPHQWV VXJJHVWHG WKDW SKRWRV\QWKHVLV ZDV RI WKH DQR[\JHQLF W\SH ZLWK VXOILGHDVHOHFWURQGRQRU2UHQ 6HFWLRQ,9%
' 6DOWHUQV 6DOWHUQV DUH PDQPDGH VKDOORZ SRQGV XVHG IRU WKH SURGXFWLRQ RI KDOLWH 1D&O IURP VHDZDWHU 0RVW VDOWHUQV\VWHPVDUHORFDWHGLQWURSLFDODQGVXEWURSLFDO DUHDVDQGDUHEXLOWDV PXOWLSRQGV\VWHPV 6HDZDWHU LVSXPSHGWRWKHILUVWVHWRISRQGV $IWHUHYDSRUDWLRQ KDV FDXVHG D VXIILFLHQW ULVH LQ VDOLQLW\ WKH ZDWHU LV WUDQVIHUUHGWRWKHQH[WVHWRISRQGV DQGVRRQXQWLO D
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
$2UHQ WKH GRPLQDQW SODQNWRQLF VSHFLHV LQ WKH K\SHUVDOLQH SRQGVDW
&KDSWHU 6DOWVDQG%ULQHV 7KLV DEVRUSWLRQ ZDV GXH WR D KLJK FRQWHQW RI \HW XQLGHQWLILHG P\FRVSRULQHOLNH DPLQR DFLGV 0$$ V 2UHQ 6SHFWUD RI WKH XSSHU F\DQREDFWHULDO OD\HU LQ WKH J\SVXP FUXVW RI WKH 6DOLQVGH*LUDXG VDOWHUQVKRZHGDVKDUSULVHLQDEVRUEDQFHIURPWR QP WKH ORZHVW ZDYHOHQJWK PHDVXUHG &DXPHWWH HWDO 7KHVHGDWDVXJJHVWWKDW895DEVRUELQJ FRPSRXQGV PD\ EH SUHVHQW WKHUH LQ KLJK FRQFHQWUDWLRQV 7KH SRVVLEOH IXQFWLRQ RI WKHVH FRPSRXQGV LQ K\SHUVDOLQH HQYLURQPHQWV LV GLVFXVVHG LQ6HFWLRQ9% Aphanothece (Coccochloris) PDWV DVVRFLDWHG ZLWK J\SVXP FUXVWV ZHUH DOVR UHSRUWHG WR RFFXU LQ $XVWUDOLDQVDOWHUQV&ROHPDQDQG:KLWH -RQHV HWDO 6DPP\ DQG EURZQUHGOD\HUVRI Aphanothece LQ VOLPH DQG ULFK LQ OLJKWSURWHFWLQJ FDURWHQRLGV ZHUH DOVR IRXQG LQ VDOWHUQV LQ (J\SW 7DKHUHWDO %UD]LO 'H0HGHLURV 5RFKD DQG &DPDUUD DQG ,QGLD 5DKDPDQ HW DO 6HVKDGULDQG%XFK $V R VDOLQLW\ LV H[FHHGHG WKH QXPEHU RI F\DQREDFWHULD GHFUHDVHV SRSXODWLRQV RI Dunaliella LQFUHDVH LQ DEXQGDQFH DQG ILQDOO\ WKH KDORSKLOLF DUFKDHDSUHGRPLQDWHLQWKHVDWXUDWHGSRQGV
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phanothece halophytica ZDV SUHVHQW DW D ZLGH UDQJH RI WHPSHUDWXUHV DQG VDOLQLWLHV & DQG J/ VDOW Phormidium hypolimneticum GHYHORSHG RSWLPDOO\ DW J/ VDOW DQG & 2WKHUVSHFLHVIRXQG ZHUH Chroococcidiopsis kashaii, Chlorogloea fritschii, Phormidium hypersalinum, DQG Spirulina subsalsa. 7KH VXPPHU EORRP RI Aphanothece halophytica DSSHDUHG KLJKO\ WROHUDQW RI
KLJK OLJKW LQWHQVLWLHV ,Q WKH GHHSHU ]RQH ZLWK WHPSHUDWXUHV LQ H[FHVV RI&Aphanothece IRUPHG D VRIW JHODWLQRXV PDW 'RU DQG 3D] 2WKHU SKRWRV\QWKHWLF PLFURRUJDQLVPV VXFK DV GLDWRPV ZHUHDOVRSUHVHQW'RUDQG(KUOLFK 'LIIHUHQW VWUDLQV RIAphanothece halophytica ZHUH LVRODWHG IURPWKH VRODUSRQGV $OO SURYHG HXU\KDOLQH DQG HXU\WKHUPDO EXW WKH\ GLIIHUHG LQ WKHLU RSWLPXP HQYLURQPHQWDO UDQJHV DQG LQ WKH H[WHQW RI SRO\PRUSKLVP GLVSOD\HG &HOO VL]H JHQHUDOO\ LQFUHDVHGZLWKLQFUHDVLQJVDOLQLW\ 7ZR LVRODWHVZHUH VWXGLHG ,Q ERWK WKH VDOLQLW\ DQG WHPSHUDWXUH UHTXLUHPHQWV ZHUH VKRZQ WR EH FRXSOHG DQG DQ LQFUHDVH LQ WHPSHUDWXUH ZDV FRPSHQVDWHG IRU E\ DQ LQFUHDVH LQ RSWLPXP VDOLQLW\ 'RU DQG +RUQRII DE
,,, 3K\VLRORJLFDO 3URSHUWLHVRIWKH0DMRU +DORSKLOLF &\DQREDFWHULD 7KRXJK D ZLGH UDQJH RIF\DQREDFWHULD EHORQJLQJ WR GLIIHUHQW WD[RQRPLF JURXSV ZHUH UHSRUWHG WR OLYH DW KLJK VDOLQLWLHV WZR PDMRU W\SHV GRPLQDWHG K\SHUVDOLQH HQYLURQPHQWV WKH ILODPHQWRXV Microcoleus chthonoplastes, D FRVPRSROLWDQ PDW EXLOGLQJ F\DQREDFWHULXP IRXQG IURP VHDZDWHU VDOLQLW\ WR VDOLQLWLHV H[FHHGLQJ J/ DQG WKH XQLFHOOXODU Aphanothece halophytica DOVR GHVLJQDWHG DV Aphanocapsa, Synechococcus, Cyanothece RU Coccochloris). 7KHVH VSHFLHV ZHUH WKH VXEMHFW RI PDQ\ VWXGLHV DQG PRUH LQIRUPDWLRQ LV NQRZQ DERXW WKHPWKDQDERXWRWKHUKDORSKLOLFF\DQREDFWHULD
$ 0LFURFROHXV FKWKRQRSODVWHV Microcoleus chthonoplastes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icrocoleus KDV DOVR EHHQ SHUIRUPHG .DUVWHQ DQG *DUFLD3LFKHO 3K\ORJHQHWLF DQDO\VLVRQWKHEDVLVRI 6 U51$VHTXHQFHVVKRZHG Microcoleus WR EH FORVHO\ UHODWHG WR WKH Oscillatoria
JURXS *DUFLD3LFKHO HW DO :LOPRWWH 'XH WR LWV PRGH RI JURZWK WKLFN EXQGOHV ZLWKLQ D FRPPRQ SRO\VDFFKDULGH VKHDWK Microcoleus LV RQH RI WKH PDLQ FRPSRQHQWV RI F\DQREDFWHULDO PDWV ZRUOGZLGH DQG RWKHU PLFURRUJDQLVPV ILQG WKHLU KDELWDWZLWKLQWKHFRKHUHQWPDWVEXLOWE\WKLVIRUP 7KHRSWLPDOWHPSHUDWXUHIRUJURZWKRIMicrocoleus ZDVDURXQG&DQGRSWLPDOUDWHVRISKRWRV\QWKHVLV ZHUHDFKLHYHGDWS+YDOXHVDURXQG EHVWEHWZHHQ DQG 'XELQLQ HWDO D 3KRWRV\QWKHVLV DV PHDVXUHG E\ R[\JHQ HYROXWLRQ DQGRU &2 SKRWRLQFRUSRUDWLRQ ZDVPRVWUDSLGDW R VDOLQLW\ EXWHYHQDWVDOLQLWLHVDVKLJKDVR SKRWRV\QWKHWLF DFWLYLW\ ZDV GHWHFWHG +RZHYHU WKH PRVW UDSLG LQFUHDVH LQ ELRPDVV H[SUHVVHG LQ WHUPV RIV\QWKHVLV RISURWHLQ RU FKORURSK\OO a ZDV IRXQG DW R DQG WKH PRVW UDSLG LQFUHDVH LQ FDUERK\GUDWH FRQWHQW DW R VDOLQLW\ 'XELQLQ HWDO D $Q LPSRUWDQW SDUW RI WKH SKRWRV\QWKHWLFDOO\IL[HG FDUERQ ZDV SUREDEO\ XVHG IRU WKH V\QWKHVLV RI JOXFRV\OJO\FHURO WKH RUJDQLF RVPRWLF VROXWH DFFXPXODWHG E\ Microcoleus, DQG XVHG WR DFKLHYH RVPRWLF EDODQFH ZLWK WKH KLJK VDOW FRQFHQWUDWLRQV RIWKH VXUURXQGLQJ PHGLXP 6HFWLRQ 9% 5DWHV RISKRWRV\QWKHVLV LQ Microcoleus PDWV PD\EHYHU\ KLJK DQGDUHOLPLWHG E\ WKH UDWH RI VXSSO\ RI &2 5RWKVFKLOG DQG 0DQFLQHOOL5RWKVFKLOG Microcoleus ILODPHQWV RIWHQ DUH IRXQG DW RU QHDU WKH LQWHUIDFH EHWZHHQ WKH DHURELF XSSHU OD\HU RIWKH VHGLPHQW DQG WKH DQDHURELF GHHSHU OD\HUV ZKHUH GLVVLPLODWRU\ VXOIDWH UHGXFWLRQ RFFXUV )LODPHQWV RIWHQ DUHH[SRVHG WR VXOILGH HVSHFLDOO\ DW WKH HQG RI WKH QLJKW DQG LQ HDUO\ PRUQLQJ ZKHQ VXOILGH SURGXFHG LQ WKH VHGLPHQW GLIIXVHV XSZDUG DQG VXOILGHXWLOL]DWLRQE\SKRWRV\QWKHWLF VXOIXUEDFWHULDLV OLPLWHG EHFDXVH RI ODFN RI DYDLODEOH OLJKW 7KRXJK QRW DV YHUVDWLOH DV Oscillatoria limnetica IURP 6RODU /DNH ZKLFKLVDEOHWRJURZZLWKKLJKFRQFHQWUDWLRQV RIVXOILGH DV HOHFWURQ GRQRU 2UHQ DQG 3DGDQ 2UHQ HW DO Microcoleus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icrocoleus PDWV-¡UJHQVHQHWDO Microcoleus, OLNH RWKHU ILODPHQWRXV
$2UHQ F\DQREDFWHULD FRQWDLQV SRO\XQVDWXUDWHG IDWW\ DFLGV & & DQG & WKH ELRV\QWKHVLV RIZKLFK LV R[\JHQGHSHQGHQW $V D FRQVHTXHQFH QR UHDO DQDHURELF JURZWK ZLOO RFFXU +RZHYHU LQ WKH SUHVHQFH RIR[\JHQ VXOILGH FDQ GULYH JURZWK ZLWKRXW WKH SDUWLFLSDWLRQ RI SKRWRV\VWHP ,, GH :LW HW DO $V DQ DGDSWDWLRQ WR SURORQJHG SUHVHQFH XQGHU DQDHURELF FRQGLWLRQV Microcoleus chthonoplastes DOVR SRVVHVVHV D ZHOO GHYHORSHG IHUPHQWDWLYH PHWDEROLVP WKDW SHUPLWV IHUPHQWDWLRQ RIHQGRJHQRXV FDUERK\GUDWH UHVHUYHV ZLWK DFHWDWH DV WKH PDLQ SURGXFW Microcoleus LV DOVR DEOH WR XVH HOHPHQWDO VXOIXU DV DQ HOHFWURQ DFFHSWRU IRU DQDHURELF UHVSLUDWLRQ DQG WKLV OHDGV WR WKH IRUPDWLRQ RI+6 LQ WKHGDUN'XELQLQDQG*HUDVLPHQNR
% $SKDQRWKHFHKDORSK\WLFD Aphanothece halophytica LV IRXQG ZRUOGZLGH LQ K\SHUVDOLQH HQYLURQPHQWV ,W ZDV ILUVW GHVFULEHG E\ +RIDQG )UpP\ LQ DQG ZDV H[WHQVLYHO\VWXGLHG VLQFH %URFN UHFRJQL]HG LWV LPSRUWDQFH DV D PRGHO RUJDQLVP IRU WKH VWXG\ RI OLIH DW KLJK VDOW FRQFHQWUDWLRQV 7KH QRPHQFODWXUH RIAphanothece halophytica DQG UHODWHG RUJDQLVPV LV H[WUHPHO\ FRQIXVLQJ DQG LW LV ZHOO SRVVLEOH WKDW PRUSKRORJLFDO W\SHV GHWHFWHG LQ QDWXUH DQG LVRODWHG VWUDLQV GHVFULEHG XQGHU GLIIHUHQW QDPHVGRLQIDFWEHORQJWRDVLQJOHIRUPDVGLVFXVVHG LQ%RURZLW]ND 7KH H[LVWLQJ FRQIXVLRQ LQ WKH QRPHQFODWXUH RIWKHVH RUJDQLVPV JUHDWO\ FRPSOLFDWHV WKH V\QWKHVLV RI GDWD RQ SK\VLRORJ\ DQG HFRORJ\ RI WKHGLIIHUHQWLVRODWHV 7KXV 'URXHWDQG'DLO\ LQFRUSRUDWHG A. halophytica LQ WKH QHZ WD[RQ Coccochloris elabens, DQGWKLVQDPHZDVXVHGIRUWKH GHVFULSWLRQ RI DQ LVRODWH IURP D VDOW SRQG LQ 3XHUWR 5LFR .DR HW DO %URFN XVHG WKH QDPHV Aphanothece halophytica DQG Aphanocapsa halophytica LQ DQ LQWHUFKDQJHDEOH ZD\ IRU KLV VWUDLQ IURP*UHDW6DOW/DNH8WDK 6SHOOLQJYDULDQWVVXFKDV Aphenotheca DQG Aphenocaspa VLF FDQ DOVR EH IRXQG LQ WKH OLWHUDWXUH )ULHGPDQQ HW DO 6WDQLHU HW DO SODFHG WKH RUJDQLVP LQ W\SRORJLFDO JURXS $ RIWKH Chroococcales VHHDOVR 7LQGDOO HW DO DQG LQ WKH FODVVLILFDWLRQ RI 5LSSNDHWDO WKHVSHFLHV EHORQJV WRWKH JHQXV Synechococcus. 7KXV WKH Synechococcus VWUDLQ GHVFULEHG IURP $EX *DEDUD /DNH LQ :DGL 1DWUXQ (J\SW ,PKRII HW DO DQG IURP VDOWHUQV LQ 4XHHQVODQG $XVWUDOLD &ROHPDQ DQG :KLWH PD\ EHORQJ WR WKLV JURXS 7KH JHQXV Cyanothece, FUHDWHG E\ .RPiUHN LQFOXGHV SDUW RI WKH
&KDSWHU 6DOWVDQG%ULQHV IRUPHU JHQXV Synechococcus. 7KH VWUDLQ LVRODWHG E\ 'H 3KLOLSSLV HW DO IURP D VDOW SDQ LQ 6RPDOLODQG DQG GHVFULEHG DV Cyanothece VS ZDV IRXQG WR EH YHU\ VLPLODU WR DQ LVRODWH IURP KHOLRWKHUPDOVDOWZRUNV LQ*UHHFHQDPHG Cyanothece halobia 5RXVVRPRXVWDNDNLDQG$QDJQRVWLGLV ,Q YLHZ RI WKH FRQIXVLQJ VWDWH RI WKH QRPHQFODWXUH HVSHFLDOO\ LQWKHJURXS RIXQLFHOOXODUF\DQREDFWHULD , JODGO\DJUHHZLWK WKH VWDWHPHQWE\%URFN WKDW LW VKRXOGEHHPSKDVL]HGWKDWWKHVSHFLHV GHVLJQDWLRQ KDV OLWWOH VFLHQWLILF PHDQLQJ $Q LQGHSWK SK\ORJHQWLF DQDO\VLV RI WKH XQLFHOOXODU H[WUHPHO\ KDORWROHUDQW F\DQREDFWHULD ZDV SXEOLVKHG UHFHQWO\ *DUFLD3LFKHOHWDO 7R P\ NQRZOHGJH IHZ PRGHUQ WD[RQRPLF SK\ORJHQHWLF VWXGLHV LI DW DOO KDYH EHHQ SHUIRUPHG ZLWK Aphanothece. 7KH PRO *& FRQWHQW RI LWV '1$ ZDV UHSRUWHG WR EH PRO :DWHUEXU\ DQG 5LSSND +RZHYHU LW LV VWLOO XQFOHDU ZKHWKHU Aphanothece halophytica )UpP\ LV D VLQJOH H[WUHPHO\ HXU\KDOLQH VSHFLHV DGDSWDEOH WR YDULHW\ RI VDOLQLWLHV RU D JURXS RI SK\ORJHQHWLFDOO\ GLVWLQFW IRUPV 6LJQLILFDQW GLIIHUHQFHV LQ VKDSH DQG VL]H ZHUH QRWHG IRU WKLV IRUP DQG FHOO PRUSKRORJ\ YDULDWLRQV FRPSOLFDWH WKH WD[RQRPLF SLFWXUH 7KH PRUSKRORJLFDO YDULDELOLW\ ZDV PHQWLRQHG E\ +RIDQG )UpP\ LQ DQG
SKRWRV\QWKHVLV *DUOLFN HW DO DQG WKH SURSHUWLHVRI58%,6&2$VDPLHWDO Aphanothece halophytica UHTXLUHV DPLQLPXP VDOW FRQFHQWUDWLRQ RI DURXQG J / DQG JURZV XS WR VDOLQLWLHV DV KLJK DV R 0RVW VWXGLHV LQGLFDWHG RSWLPXP FRQFHQWUDWLRQV IRU JURZWK LQ WKH UDQJH RI R %RURZLW]ND %URFN .DRHWDO 7LQGDOOHWDO
WUDQVFULSWLQFUHDVHGXSRQVXEMHFWLRQWRKHDWVWUHVVDQG DOVR XSRQ WUDQVIHU WR D K\SHURVPRWLF HQYLURQPHQW VXJJHVWLQJ D UROH RI 'QD. LQ WKH UHFRYHU\ RI A. halophytica IROORZLQJK\SHURVPRWLFVWUHVV/HHHWDO 2WKHU FHOOXODU FRPSRQHQWV RI A. halophytica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
$2UHQ RYHUWKH ZKROH UDQJH RIVDOW FRQFHQWUDWLRQV EXW ZDV JUHDWO\ LQFUHDVHG ZKHQ WKH FXOWXUHV UHDFKHG WKH VWDWLRQDU\ SKDVH RI JURZWK $ VLPLODU UHVXOW ZDV REWDLQHG ZLWK DQ LVRODWH GHVLJQDWHG Synechococcus IURP D J\SVXP FUXVW IURP D VDOWHUQ LQ :HVWHUQ $XVWUDOLD 1XWULHQW OLPLWDWLRQ ZDV SUREDEO\ UHVSRQVLEOH IRU WKH DFWLYDWLRQ RI H[WUDFHOOXODU SRO\VDFFKDULGHSURGXFWLRQ5RX[ $QDO\VLVRI WKH VOLPH VKRZHG WKH SUHVHQFH RI JOXFRVH IXFRVH PDQQRVH DQG JDODFWRVH LQ WKH UDWLR ZLWK PLQRU FRQVWLWXHQWV 8URQLF DFLGV RU QLQK\GULQSRVLWLYH FRPSRXQGV ZHUH QRW GHWHFWHG 7KH DPRXQWV RI VOLPH DFFXPXODWHG LQ Aphanothece FXOWXUHV ZHUH DOVR VKRZQ WR LQFUHDVH ZLWK LQFUHDVLQJ JURZWK WHPSHUDWXUH 'RU DQG +RUQRII D 7KH SUHVHQFH RI FRSLRXV DPRXQWV RI SRO\VDFFKDULGHV VXUURXQGLQJWKHFHOOV PDNHV LVRODWLRQRIAphanothece halophytica IURPQDWXUDOVDPSOHVGLIILFXOW 7KXVWKH D[HQLF FXOWXUH RI D VWUDLQ IURP D &DOLIRUQLDQ VRODU HYDSRUDWLRQ SRQG ZDV DFKLHYHG WKURXJK D SURWRFRO LQYROYLQJ WUHDWPHQW ZLWK GLIIHUHQWDQWLELRWLFV GHQVLW\ JUDGLHQW FHQWULIXJDWLRQ DQG XOWUDYLROHW UDGLDWLRQ
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scillatoria limnetica EHFDPH DODERUDWRU\PRGHO IRU WKH VWXG\ RI WKH ELRFKHPLVWU\ RI DQR[\JHQLF SKRWRV\QWKHVLVLQF\DQREDFWHULD
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scillatoria limnetica 6HFWLRQ ,,% IRUPV GHQVH EORRPV DWWKH ORZHUHQGRIWKHK\SROLPQLRQQHDUWKHERWWRP ZKHUH VXOILGH FRQFHQWUDWLRQV DUH WKH KLJKHVW %LRPDVV DQG SULPDU\ SURGXFWLYLW\ PD\ EH H[WUHPHO\ KLJK LQ WKLV OD\HU&RKHQ HWDO E 7KH VWUDLQ ZDV LVRODWHG DQG ZDV VWXGLHG H[WHQVLYHO\ GXULQJ WKH SDVW WZHQW\ \HDUV Oscillatoria limnetica FDQ VKLIWEHWZHHQ WZR W\SHV RI SKRWRV\QWKHVLV R[\JHQLF SKRWRV\QWKHVLV FKDUDFWHULVWLF RI DOO F\DQREDFWHULD LQ ZKLFK WZR SKRWRV\VWHPV FRRSHUDWH WR VSOLWZDWHU DQG DOORZ &2 SKRWRDVVLPLODWLRQ DQG DQR[\JHQLF SKRWRV\QWKHVLV LQZKLFKVXOILGHVHUYHVDVHOHFWURQGRQRULQDSURFHVV ZKLFK GRHV QRW UHTXLUH SDUWLFLSDWLRQ RI SKRWRV\VWHP ,, $FFRUGLQJO\ '&08 DQ LQKLELWRU RI SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUWDW WKH DFFHSWRU VLGH
RISKRWRV\VWHP ,, GRHV QRW LQKLELW VXOILGHGHSHQGHQW &2 SKRWRDVVLPLODWLRQ &RKHQ HW DO E 3KRWRV\VWHP,,GRHVQRWSDUWLFLSDWHLQ WKHSURFHVV DV VKRZQ IURP D FRPSDULVRQ RI SKRWRV\QWKHWLF DFWLRQ VSHFWUDLQWKH SUHVHQFH DQG DEVHQFH RI'&08 2UHQ HWDO 7KH SKHQRPHQRQLVGXHWRWKHIDFWWKDW ORZ FRQFHQWUDWLRQV RIVXOILGH LQKLELW HOHFWURQ IORZ DW WKH GRQRU VLGH RISKRWRV\VWHP ,, 2UHQ HW DO ,W PD\ EH QRWHG WKDW LQ RWKHU V\VWHPV H[DPLQHG WKH SDUWLFLSDWLRQ RISKRWRV\VWHP ,, WR WKH SKRWRV\QWKHWLF SURFHVVHV LQ WKH SUHVHQFH RI VXOILGH ZDV VLJQLILFDQW &RKHQ D 7KH RSWLPDO VXOILGH FRQFHQWUDWLRQ IRUWKHSURFHVV ZDV DURXQG P0 DQG HOHPHQWDO VXOIXU ZDV VKRZQ WR EH WKH VROH SURGXFW RI VXOILGH R[LGDWLRQ &RKHQ HW DO F 7KH VKLIW IURP R[\JHQLFWRDQR[\JHQLFSKRWRV\QWKHVLVUHTXLUHG DODJ SHULRGRIDIHZKRXUV DQG LQYROYHG V\QWKHVLV RIQHZ SURWHLQV2UHQDQG3DGDQ RQHRIZKLFKZDVD VXOILGHTXLQRQH UHGXFWDVH WKH NH\ HQ]\PH UHTXLUHG WR IHHG HOHFWURQV IURP VXOILGH LQWR WKH SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW FKDLQ EHWZHHQ WKH DFFHSWRU VLWH RI SKRWRV\VWHP ,, DQG WKH GRQRU VLWH RISKRWRV\VWHP , 7KLV HQ]\PH ZDV SDUWLDOO\ FKDUDFWHUL]HG $ULHOL HW DO Oscillatoria limnetica FDQ IL[ &2 LQ WKH OLJKW LQ WKH SUHVHQFH RIKLJK VXOILGH FRQFHQWUDWLRQV DQG FDQ DOVR JURZ DQDHURELFDOO\ LQ WKH SUHVHQFH RI VXOILGH 2UHQDQG3DGDQ *URZWK LQ WKH SUHVHQFH RI PROHFXODU R[\JHQ UHTXLUHV VSHFLDO DGDSWDWLRQV VXFK DV WKH DEVHQFH RI SRO\XQVDWXUDWHG IDWW\ DFLGV DEXQGDQW LQ WKH PHPEUDQH OLSLGV RI PRVW ILODPHQWRXV F\DQREDFWHULD DQG WKH SUHVHQFH RI DQ R[\JHQLQGHSHQGHQW SDWKZD\ WR V\QWKHVL]H PRQRXQVDWXUDWHG IDWW\ DFLGV Oscillatoria limnetica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scillatoria limnetica LV WKH SUHVHQFH RI GLIIHUHQW PRGHV RI HQHUJ\ JHQHUDWLRQ XQGHU DQDHURELF FRQGLWLRQV LQ WKH GDUN 2QH VXFK PHFKDQLVP LQYROYHV DQDHURELF UHVSLUDWLRQ RIHQGRJHQRXV VWRUDJH PDWHULDOV XVLQJ HOHPHQWDO VXOIXU IRUPHG GXULQJ WKH GD\ E\ SKRWRV\QWKHWLF R[LGDWLRQ RI VXOILGH DV HOHFWURQ DFFHSWRU ZLWK WKH IRUPDWLRQ RI &2 DQG
$2UHQ
VXOILGH2UHQDQG6KLOR ,QGLFDWLRQVH[LVWWKDW WKLV W\SH RI PHWDEROLVP PD\ EH RSHUDWLYH LQ 6RODU /DNH ZKHUH VXOILGH IRUPDWLRQ DW WKH H[SHQVH RI HOHPHQWDO VXOIXU ZDV REVHUYHG EHORZ WKH FKHPRFOLQH GXULQJ WKH QLJKW -¡UJHQVHQ HW DO $QRWKHU PRGH RIDQDHURELF JURZWKWKDWUHIOHFWV WKH PHWDEROLF IOH[LELOLW\ RIOscillatoria limnetica LV IHUPHQWDWLRQ RI HQGRJHQRXV FDUERQ UHVHUYHV ZLWK WKH IRUPDWLRQ RI ODFWDWH DV WKH PDLQ SURGXFW 2UHQ DQG 6KLOR :KHWKHU ODFWDWH LV WKH VROH SURGXFW DV VXJJHVWHG UHPDLQV WR EH DVFHUWDLQHG DFHWDWH DQG HWKDQRO ZHUH IRXQG DV DGGLWLRQDO SURGXFWV GXULQJ DQDHURELF IHUPHQWDWLRQ RI HQGRJHQRXV FDUERQ UHVHUYHV LQ D PDULQH Oscillatoria VS +H\HU HW DO 0RH]HODDUHWDO 6FUHHQLQJ RIWKH 6RODU /DNH F\DQREDFWHULDO LVRODWH Aphunothece halophytica VKRZHGWKDWWKLV VWUDLQ DOVR KDGWKH FDSDFLW\ WR XVH VXOILGH DV HOHFWURQ GRQRU LQ DQR[\JHQLF SKRWRV\QWKHVLV *DUOLFN HW DO +RZHYHU WKLV IRUP ZDV PXFK OHVV VXOILGH WROHUDQW WKDQ Oscillutoria limnetica, DQG WKH RSWLPDO VXOILGH FRQFHQWUDWLRQIRUWKHSURFHVVZDVP0RQO\ $OVR LQ WKH Aphanothece VWUDLQ HOHPHQWDO VXOIXU ZDV LGHQWLILHG DV WKHSURGXFWRIVXOILGH R[LGDWLRQ *DUOLFN HW DO Microcoleus LQ WKH 6RODU /DNH PLFURELDO PDW LV DQRWKHU YHUVDWLOH IRUPWKDW PD\ XVH R[\JHQLF DQG DQR[\JHQLF SKRWRV\QWKHVLV VLPXOWDQHRXVO\ LQ WKH SUHVHQFH RI ORZ VXOILGH FRQFHQWUDWLRQV ,Q WKLV FDVH SKRWRV\VWHP ,, ZDV SDUWLDOO\ LQKLELWHG DQG DQR[\JHQLF SKRWRV\QWKHVLV ZDV SDUWLDOO\ LQGXFHG &RKHQ E &RKHQ HW DO -¡UJHQVHQHWDO
% +DPHL0D]RU :KLOH LQ 6RODU /DNH WKH KLJK VXOILGH FRQFHQWUDWLRQV DUHGXHWRELRORJLFDODFWLYLW\ WKHVXOILGHFRQWHQW P0 RI WKH +DPHL 0D]HU VSULQJV 6HFWLRQ ,,& LV JHRWKHUPDO $V D UHVXOW RIWKH ORZ 3+ RIWKH ZDWHU aS+ PRVW RI WKH VXOILGH SUHVXPDEO\ H[LVWV DV WKH XQGLVVRFLDWHG IRUP +6 ZKLFK LV PXFK PRUH WR[LF WR OLIH WKDQ WKH LRQL]HG IRUPV +6 DQG 6 SUHVHQW DW KLJKHU 3+ YDOXHV 7KH Oscillatoria PDW 3ODWHV D E WKDW FRYHUHG WKH SRROV DQG RXWIORZ FKDQQHOV RI WKH VSULQJ GLVSOD\HG KLJK UDWHV RI &2 SKRWRDVVLPLODWLRQLQWKHSUHVHQFHRIVXOILGH DQGOLWWOH RU QR LQKLELWLRQ ZDV REVHUYHG E\ '&08 2UHQ /LWWOHDGGLWLRQDOZRUNZDVGRQHRQWKHVWUDLQ DQGQRFKDUDFWHUL]DWLRQRIIDWW\DFLGFRPSRVLWLRQZDV SHUIRUPHG WR DVVHVV ZKHWKHU PDHURELF JURZWK RFFXUUHGWKHUH
& 2WKHU(QYLURQPHQWV $QR[\JHQLF SKRWRV\QWKHVLV ZLWK VXOILGH DV HOHFWURQ GRQRU ZDV LPSOLFDWHG DV DQ HFRORJLFDOO\ LPSRUWDQW SURFHVV LQ D QXPEHU RI DGGLWLRQDO K\SHUVDOLQH HQYLURQPHQWV 7KH Synechococcus VWUDLQ LVRODWHG IURP WKH K\SHUVDOLQHDONDOLQHODNHVRI:DGL1DWUXQ ZDV VKRZQ WR EH DEOH WR SHUIRUP DQR[\JHQLF SKRWRV\QWKHVLV LQ WKHSUHVHQFHRIVXOILGH,PKRIIHWDO ,Q WKH J\SVXP FUXVWV RI WKH 6DOLQVGH*LUDXG VDOWHUQVLQWKHVRXWKRI)UDQFH VXOILGHIURPWKHORZHU OD\HUV ZDV IRXQG WR UHDFK WKH Phormidium OD\HU DW WKH HQG RI WKH QLJKW &DXPHWWH &DXPHWWH HW DO 7KH SRVVLELOLW\ WKDW LQ WKH HDUO\ PRUQLQJ KRXUV WKH F\DQREDFWHULDO FRPPXQLW\ R[LGL]HV VXOILGH SKRWRV\QWKHWLFDOO\LVZHOOZRUWKDQLQYHVWLJDWLRQ ,Q WKH ODJRRQV RI /DNH 6LYDVK HDVW &ULPHD Microcoleus PDWV DOVR XVHG VXOILGH DV GRQRU ZKLFK ZDVR[LGL]HGWRWKLRVXOIDWH=DYDU]LQHWDO ,Q D PLFURELDO PDW LQ D K\SHUVDOLQH R VDOLQLW\ ODJRRQ RQ 6DQ 6DOYDGRU ,VODQG %DKDPDV '&08 DPHQGHG VDPSOHVKDG RISKRWRV\QWKHWLF DFWLYLW\ RI WKH FRQWURO 7KLV DQR[\JHQLF SKRWRV\QWKHWLF DFWLYLW\ ZDV SRVVLEO\ GXH WR Microcoleus 3LQFNQH\ DQG3HDUO
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
&KDSWHU 6DOWVDQG%ULQHV RI WKH PHFKDQLVPV RI RVPRUHJXODWLRQ LQ F\DQREDFWHULD LQFUHDVHG JUHDWO\ LQ UHFHQW \HDUV HVSHFLDOO\ LQWKHFDVH RIJOXFRV\OJO\FHURO PHWDEROLVP LQ Synechocystis VWUDLQ 3&& WKH PROHFXODU DQDO\VLV RIZKLFKSURJUHVVHG UDSLGO\ +DJHPDQQ DQG =XWKHU +DJHPDQQ HW DO 7ZR UHFHQW UHYLHZ SDSHUV JLYH DQ RYHUYLHZ RI RXU SUHVHQW XQGHUVWDQGLQJ +DJHPDQQ HW DO -RVHW HW DO
$ ,RQ0HWDEROLVP %HIRUHWKHGLVFRYHU\RIRUJDQLFFRPSDWLEOHVROXWHV LW ZDVVXJJHVWHGWKDWLQWUDFHOOXODUVDOWFRQFHQWUDWLRQV LQ KDORSKLOLFF\DQREDFWHULDZHUHTXLWHKLJK0LOOHUHWDO UHSRUWHG LQWUDFHOOXODU SRWDVVLXP FRQFHQWUDWLRQV DV KLJK DV 0 LQ Aphanothece halophytica, ZLWK ORZ LQWUDFHOOXODU VRGLXP FRQFHQWUDWLRQV %OXH GH[WUDQ ZDV XVHG LQ WKHVH VWXGLHV WR HVWLPDWH WKH LQWUDFHOOXODU DQG H[WUDFHOOXODU VSDFH LQ FHOO SHOOHWV ,QWUDFHOOXODU SRWDVVLXP FRQFHQWUDWLRQV ZHUH IRXQG WR LQFUHDVH ZLWK WKH VDOW FRQFHQWUDWLRQ RI WKH JURZWK PHGLXP DQG WKXV SRWDVVLXP ZDV FODLPHG WR EHWKH PDLQ RVPRWLFXP LQ $ halophytica 0LOOHU HW DO
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ynechocystis 3&& XSRQD VKLIW IURP IUHVKZDWHU WR 0 1D&O D UDSLG HQWU\ RI1D&O ZDV REVHUYHG ZKLFK SHUPLWWHG D SDUWLDO UHFRYHU\ RI WKH YROXPH ZLWKLQ PLQXWHV IROORZHG E\ DQ H[FKDQJH RIVRGLXP E\ SRWDVVLXP LQ PLQXWHV WKHQ DFFXPXODWLRQ RI WKH RUJDQLF RVPRWLFVROXWHV JOXFRV\OJO\FHURODQG VXFURVHHQDEOHG WKH H[WUXVLRQ RISRWDVVLXP RYHU DSHULRG RI KRXUV 5HHG HWDO 7UDQVLHQW DFFXPXODWLRQ RI VDOW ZDV DOVR GRFXPHQWHG LQ Spirulina subsalsa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gmenellum quadruplicatum, D PDULQH FRFFRLG F\DQREDFWHULXP ZKLFK JURZV IURP WR RYHU 1D&O 8SRQ WUDQVIHUIURP WR1D&OSKRWRV\QWKHVLV ZDV VHYHUHO\ GHSUHVVHG DQG UHFRYHUHG RQO\ DIWHU VHYHUDO KRXUVGXULQJZKLFKWLPHH[FHVVVRGLXPZDVSXPSHG RXWRIWKHFHOOV%DWWHUWRQDQGYDQ%DDOHQ
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alothrix LVRODWHV 0DFND\HWDO
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ynechococcus 3&& %OXPZDOG HW DO :KHQ JURZQ DW ORZ VDOW FRQFHQWUDWLRQV DOVR Microcoleus chthonoplastes SURGXFHV WUHKDORVH DV RVPRWLF VROXWH UHSODFHG E\ JOXFRV\OJO\FHURODWKLJKHUVDOLQLWLHV.DUVWHQ
*OXFRV\OJO\FHURO *OXFRV\OJO\FHURO ZDV ILUVW LGHQWLILHG E\ PHDQV RI QDWXUDO DEXQGDQFH &105 LQ D Synechococcus VS LVRODWHG IURP LQWHUWLGDO URFNV LQ $XVWUDOLD JURZLQJ IURPP0WR 0WRWDOVDOWV%RURZLW]NDHWDO *OXFRV\OJO\FHURO ZDV GHWHFWHG LQ D ZLGH YDULHW\ RI VDOW WROHUDQW F\DQREDFWHULD 6RPH RI WKH
$2UHQ PRVW ZLGHVSUHDG KDORWROHUDQW VSHFLHV DFFXPXODWH WKH VROXWH VXFK DV Microcoleus chthonoplastes .HYEULQ HW DO GLIIHUHQW Synechococcus DQG Synechocystis LVRODWHV %RURZLW]ND HW DO 5LFKDUGVRQHWDO Spirulina platensis :DUUHW DO Agmenellum quadruplicatum 7HO2U HW DO DQG Microcystis firma (UGPDQQ HW DO /DUJH FRQFHQWUDWLRQV RI JOXFRV\OJO\FHURO ZHUH GHWHFWHG LQ QDWXUDO FRPPXQLWLHV RIMicrocoleus chthonoplastes LQ /DNH 6LYDVK (DVW &ULPHD =DYDU]LQHWDO DQGLQDK\SHUVDOLQHPLFURELDO PDW GHULYHG IURP 6RODU /DNH 6LQDL 2UHQ HW DO ,QWUDFHOOXODUFRQFHQWUDWLRQV RIJOXFRV\OJO\FHURO FDQ EHYHU\KLJK ,QMicrocoleus JURZQLQ J/ VDOW JOXFRV\OJO\FHURO DFFRXQWHG IRU RI WKH FHOO GU\ ZHLJKW =DYDU]LQ HW DO 8SRQ LQFUHDVH RI VDOLQLW\ E\ K\SHURVPRWLF VKRFN DGGLWLRQDO JOXFRV\OJO\FHURO ZDV SURGXFHG ,QWKH PDULQHVWUDLQ Synechococcus 1 &2 ZDV WKH PDLQ FDUERQ VRXUFH IRU JOXFRV\OJO\FHURO ELRV\QWKHVLV LQLWLDWHG WKURXJK DQ LQFUHDVH RI PHGLXP 1D&O FRQFHQWUDWLRQ IURP WR 0 +RZHYHU XS WR PD\ EH GHULYHG IURP DYDLODEOH LQWUDFHOOXODU RUJDQLF FDUERQ UHVHUYHV ,QWKHGDUN QRLQFUHDVHLQJOXFRV\OJO\FHURO ZDVREVHUYHG0DFND\DQG1RUWRQ 6\QWKHVLV RIJOXFRV\OJO\FHUROZDVDVORZSURFHVV DQG ZDVRQO\ WKH ILQDO SKDVH LQ D VHULHV RI HYHQWV ZKLFK RFFXU GXULQJ VDOWXSVKRFN 0XOWLSKDVLF RVPRWLF DGMXVWPHQW LQ WKH HXU\KDOLQH Synechocystis 3&& XSRQ D VXGGHQ VDOLQLW\ LQFUHDVH LQYROYHG UDSLG HQWU\ RI 1D&O IROORZHG E\H[FKDQJH RIVRGLXPE\SRWDVVLXP LRQV DQG PDQ\ KRXUV RI DGDSWDWLRQ IRU WKH FRPSOHWLRQ RI JOXFRV\OJO\FHURO DFFXPXODWLRQ WR WKH SURSHU QHZ OHYHO UHTXLUHG IRU RVPRWLF EDODQFH ZLWK WKH QHZ KLJKHU VDOLQLW\ 5HHG HW DO 6HFWLRQ 9$ 8SRQ VDOW GRZQVKRFN Synechocystis 3&& UHOHDVHG SDUW RI WKH H[FHVV JOXFRV\OJO\FHURO WR WKHPHGLXP)XOGDHWDO ,QSynechocystis XS WRRIWKHORZPROHFXODUZHLJKWFRPSRXQGV ZHUH H[FUHWHG DIWHU K\SRRVPRWLF VKRFN 7KLV K\SRRVPRWLFDOO\ LQGXFHG H[XGDWLRQ ZDV DWWULEXWHG WR WUDQVLHQW FKDQJHV LQ QRQVSHFLILF PHPEUDQH SHUPHDELOLW\ 5HHG HW DO +RZHYHU LQ WKH PDULQH Agmenellum quadruplicatum VDOW GRZQVKRFN LQGXFHG LQFUHDVHG FDUERK\GUDWH WXUQRYHU DQG H[FHVV JOXFRV\OJO\FHURO ZDV SUREDEO\ PHWDEROL]HG WR JO\FRJHQ7HO2UHWDO %LRV\QWKHVLV RI JOXFRV\OJO\FHURO ZDV VWXGLHG LQ GHSWK LQFOXGLQJ DW WKH PROHFXODU ELRORJLFDO OHYHO LQ Synechocystis VWUDLQ 3&& LQ D VHULHV RISDSHUV E\WKHJURXS RI+DJHPDQQ 7KHVH VWXGLHVSURYLGHG D GHWDLOHGSLFWXUHRIWKHPHFKDQLVPDQGWKHHYHQWV WKDW
&KDSWHU 6DOWVDQG%ULQHV GHWHUPLQH WKH UHJXODWLRQ RI JOXFRV\OJO\FHURO ELRV\QWKHVLV Synechocystis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stp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ynechocystis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ynechocystis 3&& KDV DQ DFWLYH WUDQVSRUW V\VWHP IRU JOXFRV\OJO\FHURO WKDW SHUPLWV XSWDNH RI JOXFRV\OJO\FHURO IURP WKH PHGLXP 8SWDNH DFWLYLW\ ZDV HQKDQFHG LQ FHOOV DGDSWHG WR LQFUHDVLQJ FRQFHQWUDWLRQV RI 1D&O 8SWDNH ZDV HQHUJ\ GHSHQGHQW DQG ZDV LQKLELWHG E\ XQFRXSOHUV 7KH DIILQLW\ RIWKHXSWDNH V\VWHPZDV UHODWLYHO\ ORZ .P
0 2QO\ VXFURVH DQG WUHKDORVH ZHUH IRXQG WR FRPSHWH IRU WKH WUDQVSRUW V\VWHP 7KH JOXFRV\OJO\FHURO WUDQVSRUW SURWHLQ LV HQFRGHG E\ WKH JHQH ggt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ynechococcus 3&& FHOOV ZKHQWKH\ZHUHJURZQDWKLJKVDOLQLWLHVWKHOHYHOVRI D IHZ RWKHU SURWHLQV ZHUH GHSUHVVHG +DJHPDQQ HW DO 8SRQ VDOW VKRFN WKH UDWH RI SURWHLQ V\QWKHVLV GURSSHG DW ILUVW DQG ZDV IROORZHG E\ V\QWKHVLV RI VSHFLILF SURWHLQV 6RPH RI WKHVH SURWHLQV ZHUH WUDQVLHQWO\ V\QWKHVL]HG GXULQJWKH ILUVW KRXUV RIWKH DGDSWDWLRQSKDVH ZKLOH RWKHUV UHPDLQHG WR EH V\QWKHVL]HG DW HQKDQFHG UDWHV LQ VDOWDGDSWHG FHOOV+DJHPDQQHWDO
*O\FLQH%HWDLQH *O\FLQH EHWDLQH LV FKDUDFWHULVWLFDOO\ IRXQG DV DQ RVPRWLF VROXWH LQ WKH PRVW VDOW WROHUDQW RI WKH F\DQREDFWHULD ([DPSOHV RI VWUDLQV WKDW DFFXPXODWH WKLV FRPSRXQG LQFOXGH Aphanothece halophytica 5HHGHWDO D Spirulina subsalsa LVRODWHGIURP %DUGDZLO /DJRRQ LQ 6LQDL (J\SW DQ HQYLURQPHQW ZLWKIOXFWXDWLQJVDOLQLW\*DEED\$]DULDHWDO Dactylococcopsis salina 0RRUH HW DO DQG Synechocystis '815HHGHWDO D 7KHODVW QDPHG VWUDLQ ZDV LVRODWHG IURP FDOFDUHRXV VWURPDWROLWHV RI LQWHUWLGDO IODWV LQ .XZDLW ,W ZDV LGHQWLILHG DV D Synechocystis UDWKHU WKDQ Aphanothece. %HORZ R VDOLQLW\ WKH FHOOV GLYLGHG LQPRUHWKDQRQHSODQH ,QWUDFHOOXODU JO\FLQH EHWDLQH FRQFHQWUDWLRQVDVKLJKDV DQG0ZHUH PHDVXUHG LQ FHOOV JURZQ DW R R DQG R VDOLQLW\ UHVSHFWLYHO\ 0RKDPPDG HW DO *O\FLQHEHWDLQH ZDV GHWHFWHG LQ PDVVLYH DPRXQWV LQ WKH Oscillatoria PDW FRYHULQJ WKH ERWWRP RI WKH K\SHUVDOLQH VXOIXU VSULQJ RI +DPHL 0D]RU 6HFWLRQ ,,& XVLQJ SURWRQ 105 VSHFWURVFRS\ 2UHQ HW DO ,QPRVWFDVHVJO\FLQHEHWDLQHZDVIRXQGDVWKHVROH RUJDQLF RVPRWLF VROXWH EXWLQ RQH Gloeocapsa VWUDLQ
JO\FLQH EHWDLQH ZDV UHSRUWHG WR RFFXU WRJHWKHU ZLWK WUHKDORVH 0DFND\ HW DO Synechocystis '81 FRQWDLQHG PLQRU DPRXQWV RI JOXFRV\OJO\FHURO LQ DGGLWLRQ WR JO\FLQH EHWDLQH 0RKDPPDGHWDO *O\FLQH EHWDLQH ZDV YHU\ HIIHFWLYH LQ SURWHFWLQJ HQ]\PDWLF DFWLYLWLHV DJDLQVW WKH LQKLELWRU\ DFWLRQ RI VDOW *O\FLQHEHWDLQHSURWHFWHGWKHVWUXFWXUDOLQWHJULW\ DQG DFWLYLW\ RI 5XELVFR DJDLQVW WKH GDPDJLQJ LQIOXHQFH RI VDOWV LQ Aphanothece halophytica ,QFKDURHQVDNGLDQG7DNDEH ,QFKDURHQVDNGLHW DO DQGUHVWRUHG. LQGXFHGORVVRIDFWLYLW\RI JOXFRVHSKRVSKDWH GHK\GURJHQDVH LQ WKH VDPH RUJDQLVP +DZNLQV HW DO 7KH JOXFRVH SKRVSKDWH GHK\GURJHQDVH RISpirulina subsalsa ZDV PDUNHGO\LQKLELWHGE\VDOW LQKLELWLRQ DW 0 1D&O EXW LQ WKH SUHVHQFH RI JO\FLQH EHWDLQH IXOO DFWLYLW\ ZDV REWDLQHG DW1D&O FRQFHQWUDWLRQV DV KLJK DV 0 *DEED\$]DULD HW DO ,Q Synechocystis '81 .&O ZDV VWLPXODWRU\ IRU JOXWDPLQH V\QWKHWDVH DW FRQFHQWUDWLRQV XS WR 0 EXWZDVLQKLELWRU\DERYH 0 *O\FLQHEHWDLQHGLG QRWLQKLELWXS WR 0 DQG WKHSUHVHQFH RI 0 EHWDLQH DOOHYLDWHG 1D&O LQKLELWLRQ E\ DW 1D&O FRQFHQWUDWLRQV EHWZHHQ 0 :DUU HW DO 8SRQ D VDOW XSVKRFN IURP WR 0 1D&O SODVPRO\VLV RFFXUUHG LQLWLDOO\ LQ Aphanothece halophytica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phanothece halophytica, Dactylococcopsis salina, Synechococcus 3&& DQG Synechocystis '81 7KHWUDQVSRUWVHUYHG DV DVFDYHQJLQJ V\VWHPIRU H[RJHQRXVJO\FLQHEHWDLQHDQGXVHRIJO\FLQHEHWDLQH SUHVHQWLQWKHPHGLXPPD\EHDQHIIHFWLYHVWUDWHJ\LQ HQYLURQPHQWV RI IOXFWXDWLQJ VDOLQLW\ 0RRUH HW DO *O\FLQH EHWDLQH WUDQVSRUW ZDV QRW IRXQG LQ KDORSKLOLF F\DQREDFWHULD WKDW DFFXPXODWHG VXFURVH RU JOXFRV\OJO\FHURO ,Q Aphanothece halophytica, JO\FLQH EHWDLQH ZDV WUDQVSRUWHG E\ DOLJKWGHSHQGHQW DFWLYH WUDQVSRUW V\VWHP ZKLFK ZDV VHQVLWLYH WR FKDQJHV LQ H[WHUQDO VDOW FRQFHQWUDWLRQ ERWK WUDQVLHQWO\DQGLQWKHORQJWHUP 8SWDNHFDQEHUDSLG
$2UHQ Aphanothece DFFXPXODWHG P0 JO\FLQH EHWDLQH LQWUDFHOOXODUO\ ZLWKLQ PLQXWHV ZKHQ P0 ZDV VXSSOLHG LQ WKH PHGLXP DQG KRXU RILQFXEDWLRQ RI Synechocystis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phanothece halophytica, D VORZ GHFUHDVH ZLWKLQ KRXUV RI LQWUDFHOOXODU JO\FLQH EHWDLQH FRQFHQWUDWLRQV ZDV REVHUYHGXSRQGLOXWLRQRIWKHPHGLXPIURP WR 0 1D&O ZLWKRXW DQ\ UHOHDVH WR WKH RXWVLGH PHGLXP OVKLWDQLHWDO
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
&KDSWHU 6DOWVDQG%ULQHV ZDV GLVFXVVHG E\ *DUFLD3LFKHO :KHQ PDWHULDO IURP WKH XSSHU OD\HU RI WKH (LODW J\SVXP FUXVW ZDV VXEMHFWHG WR VORZ GLOXWLRQ ZLWK GLVWLOOHG ZDWHU 0$$V DSSHDUHG UDSLGO\ LQ WKH RXWHU PHGLXP DQG WKH H[WHQW RI ORVV RI LQWUDFHOOXODU 0$$V ZDV DSSUR[LPDWHO\ SURSRUWLRQDO WR WKH H[WHQW RI WKH GLOXWLRQVWUHVVDSSOLHG2UHQ 7KHPHFKDQLVP RI UHOHDVH PD\ DJDLQ EH UHODWHG WR WUDQVLHQW SHUPHDELOLW\ FKDQJHV RI WKH FHOO PHPEUDQH DV ZDV VXJJHVWHG IRU JO\FLQH EHWDLQH UHOHDVH 5HHG HW DO
9, ,QWHUDFWLRQV %HWZHHQ &\DQREDFWHULD DQG2WKHU0LFURRUJDQLVPVLQ +\SHUVDOLQH (QYLURQPHQWV &\DQREDFWHULD DUH WKH PDLQ SULPDU\ SURGXFHUV LQ PDQ\ K\SHUVDOLQH HQYLURQPHQWV HVSHFLDOO\ LQ WKH XSSHU OD\HUV RI WKH VHGLPHQWV ZKHUH Microcoleus, Aphanothece, DQG RWKHU VSHFLHV PD\ IRUP GHQVH PDWV +HUH F\DQREDFWHULDO DFWLYLW\ GHWHUPLQHV WR D ODUJH H[WHQW WKH ELRORJLFDO SURSHUWLHV RI WKH HQWLUH HFRV\VWHPDQGF\DQREDFWHULDFDQWKXVEHH[SHFWHGWR LQWHUDFW LQ GLIIHUHQW ZD\V ZLWK WKH RWKHU PHPEHUV RI WKH ELRWD SUHVHQW ,Q WKLV VHFWLRQ WKUHH VSHFLILF LQWHUDFWLRQV DUH FRQVLGHUHG WKH UROH RI RVPRWLF VROXWHV SURGXFHG E\ F\DQREDFWHULD LQ WKH FDUERQ F\FOH RI K\SHUVDOLQH HQYLURQPHQWV WKH SUHVHQFH RI ILODPHQWRXV SKRWRV\QWKHWLF EDFWHULD DVVRFLDWHG ZLWK EXQGOHV RI Microcoleus chthonoplastes; DQG WKH IRUPDWLRQ DQG UHPRYDO RI SHUR[LGHV LQ Microcoleus FRPPXQLWLHV
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
DPLQHV 2UHQ 2QO\ UHFHQWO\ WKH ILUVW KDORSKLOLF GHJUDGLQJ EDFWHULD WKDW GHJUDGH JO\FLQH EHWDLQH ZHUH FKDUDFWHUL]HG Acetohalobium arabaticum DQG UHODWHG RUJDQLVPV 7KHVH PD\ EH UHVSRQVLEOH IRU WKH IRUPDWLRQ RI PHWK\ODWHG DPLQHV IURP JO\FLQH EHWDLQH DW KLJK VDOLQLWLHV =KLOLQD DQG =DYDU]LQ ,QWHUHVWLQJ LQWHUDFWLRQV ZHUH VKRZQ WR RFFXU EHWZHHQ WKH F\DQREDFWHULXP Microcoleus chthonoplastes DQG KDORSKLOLF DUFKDHD LQ WKH PLFURELDOPDWVRIWKH6LYDVKODJRRQ$UDEDW&ULPHD DW VDOLQLWLHV EHWZHHQ R =Y\DJLQWVHYD HW DO ,QWKHVHPDWV DWOHDVW RIWKH DHURELF KHWHURWURSKLF EDFWHULD UHFRYHUHG DV FRORQLHV ZHUH KDORSKLOLF DUFKDHD DQG WKLV SHUFHQWDJH LQFUHDVHG WR DW WKH KLJKHU VDOLQLWLHV H[DPLQHG 7KH VSHFLHV PRVW IUHTXHQWO\ LVRODWHG ZHUH Haloarcula japonica DQG Halorubrum distributum. 7KHVH DUFKDHD ZHUH WKXV FRQVLGHUHG DV HFRORJLFDOO\ VLJQLILFDQW FRPSRQHQWV RI WKH F\DQREDFWHULDO PDW FRPPXQLW\ &RPELQHG FXOWLYDWLRQ H[SHULPHQWV RI Microcoleus DQG KDORSKLOLF DUFKDHD DW R WKH XSSHU VDOW FRQFHQWUDWLRQ OLPLW DW ZKLFK Microcoleus JUHZ VKRZHG WKH H[LVWHQFH RILQWHUHVWLQJ PHWDEROLF LQWHUDFWLRQV Microcoleus VHFUHWHG D QXPEHU RI RUJDQLFDFLGVLQWRWKHPHGLXPDQGWKHVHZHUHXVHGDV FDUERQ VRXUFHV IRU WKH DUFKDHD 3DSHU FKURPDWRJUDSK\ RI WKH PHGLXP FRPSRQHQWV DIWHU LQFXEDWLRQ RI Microcoleus LQ WKH SUHVHQFH RI &2 VKRZHG WKH SUHVHQFH RI DFHWDWH EXW\UDWH aNHWRJOXWDUDWH VXFFLQDWH PDODWH IXPDUDWH DQG FLWUDWH =Y\DJLQWVHYDHW DO 7KH KDORSKLOLF DUFKDHD PD\ IXUWKHU VWLPXODWH JURZWK RIWKH Microcoleus E\ UHPRYLQJ R[\JHQ DQG UHVWULFWLQJ LWV DFFXPXODWLRQ WR WR[LFOHYHOV=Y\DJLQWVHYDHWDO
% )LODPHQWRXV3KRWRV\QWKHWLF%DFWHULD DVVRFLDWHGZLWK 0LFURFROHXV FKWKRQRSODVWHV $Q LQWHUHVWLQJ DVVRFLDWLRQ RIWKH DHURELFMicrocoleus chthonoplastes ZLWK SUHVXPDEO\ DQDHURELF ILODPHQWRXV SXUSOH EDFWHULD PD\ H[LVW ZLWKLQ Microcoleus EXQGOHV (OHFWURQ PLFURVFRSLF H[DPLQDWLRQ RI PLFURELDO PDWV RI 6RODU /DNH R VDOLQLW\ DQG *XHUUHUR 1HJUR R VDOLQLW\ VKRZHG WKH RFFXUUHQFH RI JOLGLQJ ILODPHQWRXV SKRWRWURSKLF SXUSOH EDFWHULD ZLWK VWDFNHG SKRWRV\QWKHWLF ODPHOODH EHWZHHQ WKH Microcoleus ILODPHQWVZLWKLQWKHFRPPRQVKHDWK' $PHOLRHWDO 2QH SRVVLELOLW\ LV WKDW WKHVH SXUSOH EDFWHULD XVHWR[LF VXOILGHWR ZKLFK WKH F\DQREDFWHULDO EXQGOHV ZHUHRIWHQH[SRVHG ZKLOHOLYLQJLQVKDUS JUDGLHQWVRI R[\JHQDQGVXOILGH6HFWLRQ,,,$ $OWHUQDWLYHO\WKH
EDFWHULDPD\ OLYHRQ HOHPHQWDO VXOIXU DQG WKLRVXOIDWH IRUPHG E\Microcoleus LQ WKH SURFHVV RIDQR[\JHQLF SKRWRV\QWKHVLV 6HFWLRQ ,9 RU JURZ SKRWR KHWHURWURSKLFDOO\ RQ RUJDQLF FRPSRXQGV H[FUHWHG E\ WKH F\DQREDFWHULXP 7KH ILODPHQWRXV SXUSOH EDFWHULXPZDV QRWFXOWXUHG DQG DIXUWKHU DQDO\VLV RI WKH LQWHUDFWLRQV EHWZHHQ WKH KDORSKLOLF F\DQREDFWHULXP DQG WKH DQR[\JHQLF SKRWRWURSKLF EDFWHULXPUHPDLQWREHFRPSOHWHG
& )RUPDILRQ DQG 5HPRYDO RI3HUR[LGHV LQ 0LFURFROHXV &RPPXQLWLHV Microcoleus DFFXPXODWHV SHUR[LGHV GXULQJ R[\JHQLF SKRWRV\QWKHVLV +\GURJHQSHUR[LGH ZDV SURGXFHG LQ WKH OLJKW DV D E\SURGXFW RI SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW DW WKH VWHS RIIHUUHGR[LQ R[LGDWLRQ DQG XS WR RI WKH HOHFWURQ IORZ WKURXJK WKH HOHFWURQ WUDQVSRUW FKDLQ ZDV XVHG IRU SHUR[LGH IRUPDWLRQ 'XELQLQHWDO E $WOHDVWSDUWRIWKHK\GURJHQ SHUR[LGH PD\ EH GHULYHG IURP GLVPXWDWLRQ RI VXSHUR[LGH UDGLFDOV E\ WKH )HFRQWDLQLQJ VXSHUR[LGH GLVPXWDVHSUHVHQWLQWKHF\WRSODVP $V Microcoleus ODFNVFDWDODVH SHUR[LGH DFFXPXODWLRQ FDQHDVLO\ OHDG WR JURZWK LQKLELWLRQ 7KH SUHVHQFH RI FDWDODVH VXSSOLHG E\ WKH KHWHURWURSKLF FRPSRQHQWV RIWKH PDW FRPPXQLW\ SUHYHQWHG DXWRLQKLELWLRQ RI Microcoleus E\SHUR[LGH'XELQLQHWDO E
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
$2UHQ XVHG LQ FDVHV ZKHUH LQVXIILFLHQW ELRWD GHYHORSHG 'DYLV *XLGHOLQHV IRU WKH HIIHFWLYH PDQDJHPHQW RIVDOWHUQ SRQGV ZLWK UHVSHFW WR RUJDQLF PDWWHUFRQWHQWZHUHGHYLVHG'H0HGHLURV5RFKDDQG &DPDUD6DPP\ Aphanothece UHOHDVHV PDVVLYH DPRXQWV RI VOLPH 'H 3KLOLSSLV HW DO -RQHV DQG
&KDSWHU 6DOWVDQG%ULQHV ODQG UXQRII 2WKHU PDQDJHPHQW SUDFWLFHV LQFOXGH DGMXVWPHQW RI EULQH GHSWK WR DOORZ OLJKW WR UHDFK EHQWKLF FRPPXQLWLHV 5HLQWURGXFWLRQ RI Dactylococcopsis DQG Synechococcus VS ZDVXVHGWR LPSURYH SHUIRUPDQFH RI VDOWHUQ V\VWHPV 'DYLV DQG *LRUGDQR ,QWURGXFWLRQ RI WKH EULQH VKULPS Artemia FRQWUROOHG H[FHVVLYH GHYHORSPHQW RI F\DQREDFWHULD 'H 0HGHLURV 5RFKD DQG &DPDUD 7KHXVH RIFKORULQDWLRQ XVLQJ FKORULQH JDV RU K\SRFKORULWH DOVR FRQWUROOHG WKH PXFLODJHSURGXFLQJ F\DQREDFWHULD 6HVKDGUL DQG %XFK 'DYLV +RZHYHU WKH PRVW HIIHFWLYH VWUDWHJ\ LV SUREDEO\ WR RSHUDWH VWDEOH HFRV\VWHPV ZLWK D ORZ QXWULHQWLQSXWDQGPLQLPDOGLVWULEXWLRQ&ROHPDQDQG :KLWH $QRWKHU LQWHUHVWLQJ SRWHQWLDO DSSOLFDWLRQ RI KDORSKLOLFF\DQREDFWHULDZDVGHVFULEHGE\&RKHQDQG ,GHVHV ([WUHPHO\ KLJK FRQFHQWUDWLRQV RI R[\FDURWHQRLGVHVSHFLDOO\FDQWKD[DQWKLQ ZHUHIRXQG XS WR RI WKH WRWDO RUJDQLF FDUERQ LQ K\SHUVDOLQH PDWV LQ WKH FRDVWDO DUHDV RI &KULVWPDV ,VODQG &HQWUDO 3DFLILF 7KHVH PDWV ZHUH FRPSRVHG PDLQO\ RI XQLFHOOXODU W\SHV Synechococcus, Synechocystis, Pleomorpha DQG WKLFN Oscillatoria ILODPHQWV &XOWLYDWLRQ RIWKHVH PDWV LQ D FRQWUROOHG V\VWHP ZDV SURSRVHG IRU WKH SURGXFWLRQ RI WKHVH YDOXDEOHFDURWHQRLGV
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
VSHFLHV EXW WKHUH DSSHDU WR EH QR UHSRUWV RISKDJHV ZKLFK DWWDFN WKRVH VSHFLHV LQ K\SHUVDOLQH HQYLURQPHQWV +RZHYHU WKHUHSRUW RIWKH RFFXUUHQFH RI WKUHH W\SHV RI UHVWULFWLRQ HQGRQXFOHDVHV LQ Aphanothece halophytica :KLWHKHDG DQG %URZQ PD\ LQGLFDWH WKDW GHIHQVH PHFKDQLVPV DJDLQVW IRUHLJQ EDFWHULRSKDJH '1$ PD\ EH LPSRUWDQW /LWWOH LV NQRZQ DERXW WKH in situ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ynechococcus VWUDLQ LVRODWHG IURP WKH %DMD &DOLIRUQLD VDOWIODWV ZDV DEOH WR VXUYLYH D ZHHN H[SRVXUHWRKLJKVRODU895DQG YDFXXPLQWKHVSDFH HQYLURQPHQW 0DQFLQHOOL HW DO )XWXUH LQ GHSWK VWXGLHV RI KDORSKLOLF DQG KDORWROHUDQW F\DQREDFWHULD LQ WKHLU QDWXUDO HQYLURQPHQW ZLOO XQGRXEWHGO\SUHVHQWDGGLWLRQDO VXUSULVHV
$FNQRZOHGJPHQWV 0\ SUHVHQW UHVHDUFK RQ KDORSKLOLF F\DQREDFWHULD LV IXQGHGE\WKH,VUDHO 6FLHQFH)RXQGDWLRQ DGPLQLVWHUHG E\WKH,VUDHO$FDGHP\RI6FLHQFHVDQG+XPDQLWLHV
5HIHUHQFHV $QGHUVRQ *& 6RPH OLPQRORJLFDO IHDWXUHV RI D VKDOORZ VDOLQHPHURPLFWLFODNH/LPQRO2FHDQRJU $ULHOL % 6KDKDN < 7DJOLFKW ' +DXVND * DQG 3DGDQ ( 3XULILFDWLRQDQG FKDUDFWHUL]DWLRQRIVXOILGHTXLQRQHUHGXFWDVH D QRYHOHQ]\PHGULYLQJDQR[\JHQLFSKRWRV\QWKHVLVLQ Oscillatoria limnetica. -%LRO&KHP $VDPL 6 7DNDEH 7 $ND]DZD 7 DQG &RGG *$ 5LEXORVH ELVSKRVSKDWH FDUER[\ODVH IURP WKH KDORSKLOLF F\DQRKDFWHULXP Aphanothece halophytica. $UFK %LRFKHP %LRSK\V
%DWWHUWRQ-&DQG9DQ%DDOHQ& *URZWKUHVSRQVHV RIEOXH JUHHQ DOJDH WR VRGLXP FKORULGH FRQFHQWUDWLRQ $UFK 0LNURELRO %DXOG - 2FFXUUHQFH RI EHQWKLF PLFURELDO PDWV LQ VDOLQH ODNHV+\GURELRORJLD %HUODQG% /H &DPSLRQ 7 DQG&DPSRV %DHWD 1HYHV 0+ ,QWHUDFWLRQGHODVDOLQLWpHWGHODWHPSHUDWXUHVXU ODPRUSKRORJLH OD FURLVVDQFH HW OD FRPSRVLWLRQ FHOOXODLUH G XQH &\DQREDFWpULH KDORWROpUDQWH (Aphanothece VS %RW0DU %OXPZDOG ( 0HKOKRP 5- DQG 3DFNHU / 6WXGLHV RI RVPRUHJXODWLRQ LQ VDOW DGDSWDWLRQ RI F\DQREDFWHULD ZLWK (65 VSLQSUREHWHFKQLTXHV3URF1DWO$FDG6FL86$ %OXPZDOG ( DQG 7HO2U ( 2VPRUHJXODWLRQ DQG FHOO FRPSRVLWLRQ LQ VDOWDGDSWDWLRQ RI Nostoc muscorum. $UFK 0LFURELRO %RURZLW]ND /- 7KH PLFURIORUD $GDSWDWLRQV WR OLIH LQ H[WUHPHO\VDOLQHODNHV+\GURELRORJLD %RURZLW]ND/- 'HPPHUOH6 0DFND\0$DQG1RUWRQ56 &DUERQ QXFOHDUPDJQHWLFUHVRQDQFHVWXG\RIRVPRUHJXODWLRQ LQDEOXHJUHHQDOJD6FLHQFH %UDLWKHZDLWH &-5 DQG :KLWWRQ %$ *\SVXP DQG KDOLWH DVVRFLDWHGZLWKWKHF\DQREDFWHULXPEntophysalis. *HRPLFURELRO - %URFN 7' /RZHUS+ OLPLW IRU WKH H[LVWHQFH RIEOXHJUHHQ DOJDH HYROXWLRQDU\ DQG HFRORJLFDO LPSOLFDWLRQV 6FLHQFH %URFN 7' +DORSKLOLF EOXHJUHHQ DOJDH $UFK 0LFURELRO %URFN7' 7KHUPRSKLOLF 0LFURRUJDQLVPV DQG /LIH DW +LJK 7HPSHUDWXUHV6SULQJHU9HUODJ1HZ
$2UHQ &RKHQ< .UXPEHLQ:(*ROGEHUJ 0 DQG 6KLOR0 D 6RODU /DNH 6LQDL 3K\VLFDO DQG FKHPLFDO OLPQRORJ\ /LPQRO 2FHDQRJU &RKHQ < .UXPEHLQ :( DQG 6KLOR 0 D 7KH 6RODU /DNH OLPQRORJ\ DQG PLFURELRORJ\ RI D K\SHUVDOLQH PRQRPLFWLF KHOLRWKHUPDOKHDWHG VHDPDUJLQDO SRQG JXOIRI$TXDED 6LQDL 5DSS&RPP,QW0HU0ÒGLW &RKHQ<.UXPEHLQ:(DQG6KLOR0 E 6RODU/DNH6LQDL 'LVWULEXWLRQ RI SKRWRV\QWKHWLF PLFURRUJDQLVPV DQG SULPDU\ SURGXFWLRQ/LPQRO2FHDQRJU &RKHQ < 3DGDQ ( DQG 6KLOR 0 E )DFXOWDWLYH DQR[\JHQLF SKRWRV\QWKHVLV LQ WKH F\DQREDFWHULXP Oscillatoria limnetica. %DFWHULRO &ROHPDQ 08 DQG :KLWH 0$ 7KH UROH RI ELRORJLFDO GLVWXUEDQFHV LQ WKH SURGXFWLRQ RI VRODU VDOW ,Q 6HYHQWK 6\PSRVLXP RQ 6DOW 9RO SS (OVHYLHU 6FLHQFH 3XEOLVKHUV$PVWHUGDP '¶$PHOLR('&RKHQ<DQG'HV0DUDLV '- $VVRFLDWLRQRI D QHZ W\SH RI JOLGLQJ ILODPHQWRXV SXUSOH SKRWRWURSKLF EDFWHULXP LQVLGH EXQGOHV RI Microcoleus chthonoplastes LQ K\SHUVDOLQHF\DQREDFWHULDOPDWV$UFK0LFURELRO 'DYLV -6 ,PSRUWDQFH RI PLFURRUJDQLVPV LQ VRODU VDOW SURGXFWLRQ,Q)RXUWK6\PSRVLXPRQ6DOW9RO SS 12KLR*HRORJLFDO6RFLHW\&OHYHODQG 'DYLV -6 %LRORJLFDO FRPPXQLWLHV LQ D QXWULHQW HQULFKHG VDOLQD$TXDW%RW 'DYLV -6 %LRORJLFDO PDQDJHPHQW IRUSUREOHP VROYLQJ DQG ELRORJLFDO FRQFHSWVIRUDQHZJHQHUDWLRQ RIVRODUVDOWZRUNV ,Q 6HYHQWK 6\PSRVLXP RQ 6DOW 9RO SS (OVHYLHU 6FLHQFH3XEOLVKHUV$PVWHUGDP 'DYLV -6 DQG *LRUGDQR 0 %LRORJLFDO DQG SK\VLFDO HYHQWV LQYROYHG LQ WKH RULJLQHIIHFWV DQG FRQWURO RIRUJDQLF PDWWHULQ VRODUVDOWZRUNV,QW-6DOW/DNH5HV 'H 0HGHLURV 5RFKD 5 DQG &DPDUD 05 3UHGLFWLRQ PRQLWRULQJ DQG PDQDJHPHQW RI GHWULPHQWDO DOJDO EORRPV RQ VRODU VDOW SURGXFWLRQ LQ QRUWKHDVW %UD]LO ,Q 6HYHQWK 6\PSRVLXP RQ 6DOW 9RO SS (OVHYLHU 6FLHQFH 3XEOLVKHUV$PVWHUGDP 'H 3KLOLSSLV 5 0DUJKHUL 0& 0DWHUDVVL 5 DQG 9LQFHQ]LQL 0 3RWHQWLDO RI XQLFHOOXODU F\DQREDFWHULD IURP VDOLQH HQYLURQPHQWV DV H[RSRO\VDFFKDULGH SURGXFHUV $SSO (QYLURQ 0LFURELRO 'H 3KLOLSSLV 5 0DUJKHUL 0& 3HORVL ( DQG 9HQWXUD 6 ([RSRO\VDFFKDULGH SURGXFWLRQ E\ D XQLFHOOXODU F\DQREDFWHULXP LVRODWHGIURPDK\SHUVDOLQHKDELWDW-$SSO3K\FRO 'H:LW5YDQ%RHNHO:+0 DQGYDQ*HPHUGHQ+ *URZWK RI WKH F\DQREDFWHULXP Microcoleus chtonoplastes RQ VXOILGH )(060LFURELRO(FRO 'HV 0DUDLV '- 7KH ELRJHRFKHPLVWU\ RI K\SHUVDOLQH PLFURELDO PDWV ,Q -RQHV -* HG $GYDQFHV LQ 0LFURELDO (FRORJ\9RO SS3OHQXP3UHVV1HZ
&KDSWHU 6DOWVDQG%ULQHV 3K\VLRORJFDO (FRORJ\ RI %HQWKLF 0LFURELDO &RPPXQLWLHV SS $PHULFDQ6RFLHW\IRU0LFURELRORJ\:DVKLQJWRQ'& 'URXHW ) DQG 'DLO\ : 5HYLVLRQ RI WKH FRFFRLG 0\[RSK\FHDH%XWOHU8QLY%RW6WXG 'XELQLQ $9 DQG *HUDVLPHQNR /0 'DUN DQDHURELF PHWDEROLVP RI KDORSKLOLF F\DQREDFWHULXP Microcoleus chtonoplastes. 0LNURELRORJL\D(QJOLVKWUDQVODWLRQ 'XELQLQ $9 *HUDVLPHQNR /0 DQG *XVHY 09 D 3K\VLRORJLFDOIHDWXUHV RIDVWUDLQRIMicrocoleus chthonoplastes IURP D K\SHUVDOLQH UHVHUYRLU 0LNURELRORJL\D (QJOLVK WUDQVODWLRQ 'XELQLQ$9=DVWUL]KQD\D20DQG*XVHY09E +\GURJHQ SHUR[LGH SURGXFWLRQ E\ WKH KDORSKLOLF F\DQREDFWHULXP Microcoleus chrhonoplustes. 0LNURELRORJL\D (QJOLVK WUDQVODWLRQ (KUOLFK $ DQG 'RU , 3KRWRV\QWKHWLF PLFURRUJDQLVPV RIWKH *DYLVK VDENKD ,Q )ULHGPDQQ*0 DQG .UXPEHLQ :( HGV +\SHUVDOLQH (FRV\VWHPV 7KH *DYLVK 6DENKD SS 6SULQJHU9HUODJ1HZ
SRWHQWLDO VWURPDWROLWHV RIWKH*DYLVK VDENKD ,Q )ULHGPDQQ*0 DQG .UXPEHLQ :( HG +\SHUVDOLQH (FRV\VWHPV 7KH *DYLVK 6DENKDSS6SULQJHU9HUODJ1HZ
OPKRII -) 6DKO +* 6ROLPDQ *6+ DQG 7USHU +* 7KH :DGL 1DWUXQ FKHPLFDO FRPSRVLWLRQ DQG PLFURELDO PDVV GHYHORSPHQWV LQ DONDOLQH EULQHV RI HXWURSKLF GHVHUW ODNHV *HRPLFURELRO- OQFKDURHQVDNGL $ DQG 7DNDEH 7 'HWHUPLQDWLRQ RI LQWUDFHOOXODU FKORULGH LRQ FRQFHQWUDWLRQ LQ D KDORWROHUDQW F\DQREDFWHULXP Aphanothece halophytica. 3ODQW &HOO 3K\VLRO ,QFKDURHQVDNGL $ 7DNDEH 7 DQG $ND]DZD 7 (IIHFW RI EHWDLQH RQ HQ]\PH DFWLYLW\ DQG VXEXQLW LQWHUDFWLRQ RIULEXORVH ELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH IURP Aphanothece halophytica. 3ODQW3K\VLRO ,VKLWDQL 07DNDEH7 .RMLPD.DQG7DNDEH7 5HJXODWLRQ RI JO\FLQHEHWDLQH DFFXPXODWLRQ LQ WKH KDORWROHUDQW F\DQREDFWHULXP Aphanothece hulophytica. $XVW -3ODQW 3K\VLRO -DKQNH// /HH % 6ZHHQH\ 0- DQG .OHLQ +3 $QDHURELF ELRV\QWKHVLV RI XQVDWXUDWHG IDWW\ DFLGV LQ WKH F\DQREDFWHULXP Oscillatoria limnetica. $UFK0LFURELRO -DYRU % +\SHUVDOLQH (QYLURQPHQWV 0LFURELRORJ\ DQG %LRJHRFKHPLVWU\6SULQJHU9HUODJ%HUOLQ -RQHV $* (ZLQJ &0 DQG 0HOYLQ 09 %LRWHFKQRORJ\ RI VRODUVDOWILHOGV+\GURELRORJLD -RQHV -+ DQG
$2UHQ F\DQREDFWHULXP Aphanorhece halophytica. 3ODQW 0RO %LRO 0DFND\ 0$ DQG 1RUWRQ 56 & QXFOHDU PDJQHWLF UHVRQDQFH VWXG\ RI ELRV\QWKHVLV RI JOXFRV\OJO\FHURO E\ D F\DQREDFWHULXP XQGHU RVPRWLF VWUHVV - *HQ 0LFURELRO 0DFND\ 0$ 1RUWRQ 56 DQG %RURZLW]ND /- 2UJDQLF RVPRUHJXODWRU\ VROXWHVLQF\DQREDFWHULD - *HQ 0LFURELRO 0DQFLQHOOL 5/ :KLWH 05 DQG 5RWKVFKLOG /- %LRSDQ VXUYLYDO , ([SRVXUH RI WKH RVPRSKLOHV Synechococcus VS 1DJHOL DQG Haloarcula VS WR WKH VSDFH HQYLURQPHQW $GY 6SDFH5HVLQSUHVV 0DUJKHUL 0&7UHGLFL05 %DUVDQWL /DQG%DOORQL : 7KH SKRWRV\QWKHWLF FRPPXQLW\ RI WKH 7UDSDQL VDOLQH ODJRRQV DQ DOWHUQDWLYH RSWLRQ IRU WKH H[SORLWDWLRQ RI DQ H[WUHPH HQYLURQPHQW$QQ0LFURELRO 0LNNDW 6 (IIPHUW8DQG+DJHPDQQ 0 8SWDNHDQG XVHRI WKH RVPRSURWHFWLYHFRPSRXQGV WUHKDORVH JOXFRV\OJO\FHURO DQG VXFURVH E\ WKH F\DQREDFWHULXP Synechococcus VS 3&& $UFK0LFURELRO 0LNNDW6 +DJHPDQQ 0 DQG 6FKRRU $ $FWLYH WUDQVSRUWRI JOXFRV\OJO\FHURO LV LQYROYHG LQ VDOW DGDSWDWLRQ RI WKH F\DQREDFWHULXP Synechocystis VS VWUDLQ 3&& 0LFURELRORJ\ 0LOOHU '0 -RQHV -+
&KDSWHU 6DOWVDQG%ULQHV 2UHQ$.RKO0DQG.DUVWHQ8 $QHQGHYDSRULWLFPLFURELDO PDW ZLWKLQ D J\SVXP FUXVW ]RQDWLRQ RI SKRWRWURSKV SKRWRSLJPHQWV DQG OLJKW SHQHWUDWLRQ 0DU(FRO 3URJ 6HU 2UHQ $ DQG 3DGDQ ( ,QGXFWLRQ RI DQDHURELF SKRWRDXWRWURSKLF JURZWK LQ WKH F\DQREDFWHULXUQ Oscillatoria limnetica. -%DFWHULRO 2UHQ $ 3DGDQ ( DQG $YURQ 0 4XDQWXP \LHOGV IRU R[\JHQLF DQGDQR[\JHQLFSKRWRV\QWKHVLV LQWKHF\DQREDFWHULXP Oscillatoria limnetica. 3URF1DWO$FDG6FL86$ 2UHQ $ 3DGDQ ( DQG 0DONLQ 6 6XOILGH LQKLELWLRQ RI SKRWRV\VWHP ,LQ F\DQREDFWHULD EOXHJUHHQ DOJDH DQG WREDFFR FKORURSODVWV%LRFKLP%LRSK\V$FWD 2UHQ $ DQG 6KLOR 0 $QDHURELF KHWHURWURSKLF GDUN PHWDEROLVPLQWKHF\DQREDFWHULXP Oscillatoria limnetica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¡UJHQVHQ%% %ODFNEXUQ7+ DQG&RKHQ< 0LFURHOHFWURGH VWXGLHV RISKRWRV\QWKHVLV DQG 2 +6 DQG S+ SURILOHVRIDPLFURELDOPDW/LPQRO2FHDQRJU 5LFKDUGVRQ'/ 5HHG5+DQG6WHZDUW:'3 Synechocystis 3&& DHXU\KDOLQHF\DQREDFWHULXP )(06 0LFURELRO /HWW 5LSSND 5 'HUXHOOHV -% :DWHUEXU\ -% +HUGPDQ 0 DQG 6WDQLHU 5< *HQHULF DVVLJQPHQWV VWUDLQ KLVWRULHV DQGSURSHUWLHV RISXUHFXOWXUHVRIF\DQREDFWHULD-*HQ0LFURELRO 5LWWHU'DQG
5RWKVFKLOG /- $ PRGHO IRU GLXUQDO SDWWHUQV RI FDUERQ IL[DWLRQ LQ D 3UHFDPEULDQ PLFURELDO PDW EDVHG RQ D PRGHP DQDORJ%LR6\VWHPV 5RWKVFKLOG /- *LYHU /- :KLWH 05 DQG 0DQFLQHOOL 5/ 0HWDEROLF DFWLYLW\ RI PLFURRUJDQLVPV LQ HYDSRULWHV - 3K\FRO 5RWKVFKLOG: DQG0DQFLQHOOL5/ 0RGHORIFDUERQIL[DWLRQ LQ PLFURELDO PDWV IURP 0\U DJR WR WKH SUHVHQW 1DWXUH 5RX[-0 3URGXFWLRQ RISRO\VDFFKDULGH VOLPH E\ PLFURELDO PDWV LQ WKH K\SHUVDOLQH HQYLURQPHQW RI D :HVWHUQ $XVWUDOLDQ VRODUVDOWILHOG,QW-6DOW/DNH5HV 5RXVVRPRXVWDNDNL 0 DQG $QDJQRVWLGLV . Cyanothece halobia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phanothece halophytica: DQDO\VLV RIZDOO SURWHLQV LQ PRWPXWDQWV - %DFWHULRO 6WDQLHU5< .XQLVDZD 5 0DQGHO 0 DQG &RKHQ%D]LUH * 3XULILFDWLRQ DQG SURSHUWLHV RI XQLFHOOXODU F\DQREDFWHULD RUGHU &KURRFRFFDOHV %DFWHULRO5HY 6WRO] -) 'LVWULEXWLRQ RISKRWRWURSKLF PLFUREHV LQ WKH IODW ODPLQDWHG PLFURELDO PDW DW /DJXQD )LJXHURD %DMD &DOLIRUQLD 0H[LFR%LR6\VWHPV 6XGR+%XUJHVV-* 7DNHPDVD+ 1DNDPXUD1DQG0DWVXQDJD7 6XOIDWHGH[RSRO\VDFFKDULGHSURGXFWLRQE\ WKHKDORSKLOLF F\DQREDFWHULXP Aphanocapsa halophytica. &XUU 0LFURELRO 7DKHU $* $EG HO :DKDE 6 3KLOLS * .UXPEHLQ :( DQG :DOL $0 (YDSRULWLF VHGLPHQWDWLRQ DQG PLFURELDO PDWV LQ D VDOLQDV\VWHP3RUW)RXDG(J\SW ,QW-6DOW/DNH5HV 7DNDEH7,QFKDURHQVDNGL$$UDNDZD.DQG
:DOVE\$(YDQ5LMQ-DQG&RKHQ< 7KHELRORJ\RIDQHZ JDVYDFXRODWH F\DQREDFWHULXP Dactylococcopsis salina VS QRYLQ6RODU/DNH3URF5R\6RF/RQGRQ% :DWHUEXU\ -% DQG 5LSSND 5 6XEVHFWLRQ , 2UGHU Chroococcales :HWWVWHLQ HPHQG 5LSSNDHWDO ,Q 6WDOH\ -7 %U\DQW 03 3IHQQLJ 1 DQG +ROW -* HGV %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ 9RO SS :LOOLDPV :LONLQV%DOWLPRUH :DUU 65& 5HHG 5+ &KXGHN -$ )RVWHU 5 DQG 6WHZDUW :'3 2VPRWLF DGMXVWPHQW LQ Spirulina platensis. 3ODQWD :DUU 65& 5HHG 5+ DQG 6WHZDUW :'3 2VPRWLF DGMXVWPHQWRIF\DQREDFWHULD WKHHIIHFWV RI1D&, .&, VXFURVH DQGJO\FLQHEHWDLQHRQJOXWDPLQHV\QWKHWDVHDFWLYLW\LQDPDULQH DQGDKDORWROHUDQWVWUDLQ-*HQ0LFURELRO :KDUWRQ 5$ -U 3DUNHU %& DQG 6LPPRQV *0 -U 'LVWULEXWLRQVSHFLHVFRPSRVLWLRQDQGPRUSKRORJ\RIDOJDOPDWV LQ$QWDUFWLF'U\9DOOH\ODNHV3K\FRORJLD :KLWHKHDG 35 DQG %URZQ 1/ Aha,,, $ UHVWULFWLRQ HQGRQXFOHDVH ZLWK D UHFRJQLWLRQ VHTXHQFH FRGLQJ RQO\ $7 EDVHSDLUV)(%6/HWW :KLWHKHDG35 DQG %URZQ 1/ $ VLPSOH DQG UDSLG PHWKRG IRU VFUHHQLQJ EDFWHULD IRU W\SH ,, UHVWULFWLRQ HQGRQXFOHDVHV HQ]\PHVLQAphunothece halophytica. $UFK0LFURELRO :LOPRWWH $ 0ROHFXODU HYROXWLRQ DQG WD[RQRP\ RI F\DQREDFWHULD ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 'RUGUHFKW :ULJKW6:DQG%XUWRQ+5 7KHELRORJ\RI$QWDUFWLFVDOLQH ODNHV+\GURELRORJLD
$2UHQ
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
, ,QWURGXFWLRQ
,, &RPSRVLWLRQRI&UXGH2LO
&\DQREDFWHULD QRW RQO\ SOD\HG D NH\ UROH LQ WKH IRUPDWLRQ RIVRPH LPSRUWDQW SHWUROHXP GHSRVLWV EXW WKH\ DUH RIWHQ IRXQG LQ DVVRFLDWLRQ ZLWK SUHVHQWGD\ RLOVHHSV DQGGHSRVLWV LQ ZHWDUHDV VHHEHORZ 7KH SXUSRVH RIWKLV FKDSWHU LV WR DVVHVV WKH SRVVLEOH UROHV RI F\DQREDFWHULD LQ ERWK RLO ELRV\QWKHVLV DQG GHJUDGDWLRQ ,I WKH\ DUH LPSRUWDQW LQ WKH ODWWHU FOHDUO\ WKH\ FRXOG EH XVHIXO WRROV LQ ELRUHPHGLDWLQJ RLOSROOXWHGHQYLURQPHQWV HVSHFLDOO\FRDVWDOUHJLRQV
&UXGH RLO LV D FRPSOH[ PL[WXUH RI WKRXVDQGV RI FRPSRXQGV EXWHDFK DFFXPXODWLRQ RIRLO WHQGV WR EH XQLTXH LQFRPSRVLWLRQ 6RPH EDFNJURXQG DERXW WKLV PL[WXUH LV QHHGHG WR XQGHUVWDQG WKLV FKDSWHU ,Q WHUPV RI HOHPHQW FRPSRVLWLRQ RLO LV FRPSRVHG SUHGRPLQDQWO\ RI FDUERQ DQG K\GURJHQ ZLWK PLQRU SURSRUWLRQV RI R[\JHQ QLWURJHQ DQG VXOIXU ZKLOH WUDFHV RI YDQDGLXP QLFNHO DQG RWKHU HOHPHQWV DUH DOVRXVXDOO\SUHVHQW 3HWUROHXP VDPSOHV LUUHVSHFWLYH
B. A. Whitton and M. Potts (eds), The Ecology of Cyunobacteriu, Kluwer Academic Publishers. Printed in the Netherlands.
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ediastrum DQGGLDWRPV (DUOLHU UHVHDUFK RQ WKHVH JURXSV SURYLGHG UHVXOWV RI RQO\ OLPLWHG XVH IRU LQWHUSUHWLQJ VWUDWLJUDSK\ DQG HQYLURQPHQW EHFDXVH RIWKHLUORQJDQGZLGHVSUHDG RFFXUUHQFHWKRXJKWKH\KDYHLPSRUWDQWLPSOLFDWLRQV IRU SHWUROHXP VWXGLHV DQG SRVVLEOH LQGXVWULDO GHYHORSPHQWV +RZHYHU DQ LQFUHDVLQJ QXPEHU RI VWXGLHVDUHSURYLGLQJPRUHGHWDLOHGLQIRUPDWLRQRQ W\SHV RI HQYLURQPHQWV DQG WKH F\DQREDFWHULDO SURGXFWV WKDW PD\ EH DVVRFLDWHG ZLWK WKHP 7KH
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
ZD[ HVWHUV DOFRKROV IDWW\ DFLGV RI WZR F\DQREDFWHULDO PDWV WZR DQR[\JHQLF SKRWRV\QWKHWLF EDFWHULDO PDWV DQG WKHLU FRPSRQHQW SKRWRV\QWKHWLF EDFWHULD 7KHUHVXOWV LQGLFDWHG FOHDUO\WKDWPDWOLSLGV UHIOHFWWKHLQSXWVRIWKHPDWEXLOGLQJSKRWRWURSKV 2WKHU VWXGLHV KDYH IRFXVVHG RQ SDUWLFXODU JHQHUD DQG VSHFLHV 7KH SUHGRPLQDQW VRXUFHV RI RUJDQLF PDWWHU DQG WKH PDLQ GLDJHQHWLF SURFHVVHV LQYROYHG KDYH EHHQ LQYHVWLJDWHG LQ Phormidium valderianum DQG Microcoleus chthonoplastes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cytonema DQG Schizothrix LQ WKH (YHUJODGHV )ORULGD JDYH PXFK PRUH FKDUDFWHULVWLF VLJQDOV 7KLHO HW DO ZLWK WKH VWURQJ SUHGRPLQDQFH RI DPRQJ RWKHUV QKHSWDGHFDQH Q KHSWDGHFHQHDQGWZRPRQRPHWKOKHSWDGHFDQHV
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ermocarpella, Mysosarcina, Chroococcidiopsis, Nodularia, Scytonema, Calothrix, Fischerella DQG Chlorogloeopsis. 7KH\ DOVR OLVWHG D QXPEHU RI
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ynechococcus DQG Synechocystis IRU H[DPSOH FDQ KDUGO\ OLYH KHWHURWURSKLFDOO\
&KDSWHU 2LO3ROOXWLRQ DUH XWLOL]HG YLDWKHLU R[LGDWLRQ WR IDWW\ DFLGV WKDW DUH IXUWKHUR[LGL]HGWRDFHWDWHXQLWV 7KHUH DUH VHYHUDO HDUO\ VWXGLHV RQ WKH PHWDEROLVP RI H[RJHQRXV RUJDQLF FRPSRXQGV E\ F\DQREDFWHULD ,WLV FRPPRQO\HVWDEOLVKHGWKDWH[RJHQRXV JOXFRVHLV PHWDEROL]HG YLD WKH R[LGDWLYH SHQWRVHSKRVSKDWH F\FOH 3HOUR\HWDO 5DER\HWDO -RVHI (VSDUGHOOLHUHWDO %RWWRPOH\ DQG9DQ%DDOHQ ,Q FHOOV RI Synechocystis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nabaena variabilis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nabaena doliolum. 7KH DURPDWLF DQG DVSKDOWLF IUDFWLRQV LQKLELWHG ERWK DFWLYLWLHV ZKHUHDV WKH SDUDIILQLF IUDFWLRQV ZHUH VWLPXODWRU\ 7UHDWPHQW ZLWKFUXGHRLOH[HUWVUHYHUVLEOHDVZHOO DVLUUHYHUVLEOH LQKLELWLRQ RI PLFURELDO SKRWRV\QWKHVLV 9DQ GHUPHXOHQ DQG $KHUQ :LQWHUV HW DO %DWWHUWRQ HW DO D E 0RUDOHV/RR DQG *RXW[ VXEMHFWHGVHYHUDOSK\WRSODQNWRQLVRODWHVWRWKH ZDWHU VROXEOH IUDFWLRQ RI WKH 0H[LFDQ FUXGH RLO ,VWKPXV &DFWXV DQG PHDVXUHG WKHLU JURZWK 7KH JURZWK RINitzschia closterium GLDWRP Asterionella glacialis GLDWRP Rhodomonas lens &U\SWR SK\FHDH DQG Dunaliella tertiolecta JUHHQ DOJD ZDV LQKLELWHG ZKHUHDV WKDW RI Skeletonema costatum GLDWRP DQG Agmenellum quadruplicatum F\DQREDFWHULXP ZDV QRW DIIHFWHG WKH JURZWK RI Prorocentrum minimum 'LQRSK\FHDH ZDV HQKDQFHG7DEOH 7KHUHDUH DQXPEHURIUHSRUWV RQWKH VWLPXODWLRQ RI SKRWRV\QWKHVLV GDUN UHVSLUDWLRQ DQG JURZWK RI SKRWRV\QWKHWLF PLFURRUJDQLVPV ZKHQ WKH\ DUH WUHDWHG ZLWKQDONDQHV 6RWRHWDO 9DQGHUPHXOHQ DQG $KHUQ *DXUDQG .XPDU 6FKURHGHUDQG 5HKP *DXUDQG 6LQJK $O+DVDQHWDO 1LWURJHQIL[LQJ F\DQREDFWHULD ZHUH LQFOXGHG E\ *DXU DQG 6LQJK 7DEOH SURYLGHV DQ H[DPSOH RI WKH UHVSRQVH RI F\DQREDFWHULD WR YDULRXV RLO DQG DONDQH IUDFWLRQV WR WZR QRQD[HQLF F\DQREDFWHULD Microcoleus chthonoplastes DQG Phormidium corium, LVRODWHG IURP DQ RLOULFK HQYLURQPHQW$O+DVDQHWDO *URZWKRIERWK F\DQREDFWHULD ZDV HQKDQFHG E\ WKH K\GURFDUERQV DQG WKH FKORURSK\OO WR FDURWHQH UDWLR RIWKH FHOOV GLG QRW GHFUHDVH LQ UHVSRQVH WR WKH K\GURFDUERQ WUHDWPHQW 7KH ODWWHU UHVXOW PD\ LPSO\ WKDW FUXGH RLO DQG QDONDQHV GR QRW FDXVH DQ\ VWUHVV WR WKHVH SDUWLFXODU VWUDLQV ,W LV FOHDU WKDW SKRWRV\QWKHWLF PLFURRUJDQLVPVUHVSRQG LQ YDULRXV ZD\V WR FUXGH RLO DQGLWVFRQVWLWXHQWIUDFWLRQV
66 5DGZDQDQG5+ $O+DVDQ
Table 1. (IIHFWVRIZDWHUVROXEOHIUDFWLRQ:6) RIFUXGHRLO,VWKPXV&DFWXVRQJURZWKUDWHDQGPD[LPXPFHOOGHQVLW\RIPDULQHPLFURDOJDH IURP0RUGHV/RRDQG*RXW[
63(&,(6
:6)
GLDWRP
Nitzschia closterium
GLDWRP
Asterionella glacialis
FU\SWRSK\WH
Rhodomonas lens
JUHHQ DOJD
Dunaliella tertiolecta
Agmenellum quadruplicatum F\DQREDFWHULXP
GLDWRP
Skeletonema costatum
Prorocentrum minimum
GLQRIODJHOODWH
0D[ VSHFLILF JURZWK UDWH GLYLVLRQV G 0.80
6WLPXODWLRQ RU UHGXFWLRQ LQ JURZWK UDWH RI FRQWURO
5HGXFWLRQ LQ PD[ FHOO GHQVLW\
6SHFLILFJURZWKH[SUHVVHGDV RUJDQLFFDUERQ
Table (IIHFWV RI FUXGH RLO DQG nDONDQHV RQ JURZWK DQG SLJPHQWVRIQRQD[HQLFF\DQREDFWHULDOFXOWXUHV IURP $O+DVDQHWDO
+\GURFDUERQV
1RQHFRQWURO
&UXGHRLO Q'HFDQH & Q8QGHFDQH& Q'RGHFDQH& Q7ULGHFDQH & Q7HWUDGHFDQH& Q+H[DGHFDQH & Q1RQDGHFDQH &
'U\ZHLJKWLQPJ/
,QPJ/
Microcoleus chthonoplastes
Phormidium corium
%LRPDVV
%LOLSURWHLQ
&KORURSK\OOO FDURWHQH
%LRPDVV
%LOLSURWHLQ
&KORURSK\OO FDURWHQH
:HLJKWZHLJKW
&KDSWHU 2LO3ROOXWLRQ & 2LO7R[LFLW\WR3KRWRV\QWKHWLF 0LFURRUJDQLVPV
' +\GURFDUERQ 2[LGDWLRQE\ &\DQREDFWHULD
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gmenellum quadruplicatum, %DWWHUWRQ HW DO E IRXQG WKDW IXHO RLOV VLJQLILFDQWO\ LQKLELWHG WKHLU JURZWK EXW FUXGH RLOV ZHUH PXFK OHVV WR[LF 7KH WR[LFLW\ RIWKH IXHORLOVZDVSUREDEO\GXH PDLQO\WRKLJKHUDURPDWLF FRPSRXQGV :LQWHUV HW DO VKRZHG WKDW WKH ZDWHUVROXEOH IUDFWLRQV RI IRXU WHVWHG IXHO RLOV H[KLELWHG GLIIHUHQWWR[LFHIIHFWV RQWZRF\DQREDFWHULD DQG IRXU PLFURDOJDH 7KH ZDWHUVROXEOH IUDFWLRQV RI WZR RIWKH IXHO RLOV NLOOHG ERWK WKH F\DQREDFWHULD A. quadruplicatum DQG Coccochloris elabens, DQG WKH WR[LF HIIHFW ZDV LQ SDUW GXH WR WKH FRQVWLWXHQW S WROXLGLQH 0RUH WKDQ SXUH DURPDWLF FRPSRXQGV RFFXUULQJ LQ RLO KDYH EHHQ WHVWHG E\ VHYHUDO LQYHVWLJDWRUV:LQWHUVHWDO &HUQLJOLDHW DO 1DUUR IRUWKHLUWR[LFLW\ WR WKH F\DQREDFWHULXP A. quadruplicatum GDWD WDEXODWHG E\ 1DUUR 7KH IROORZLQJ FRPSRXQGV H[KLELWHG KLJK WR[LFLW\ GLPHWK\O QDSKWKDOHQHV PHWK\O DQG HWK\O DQLOLQHV GL DQG WULPHWK\O SKHQROV GLPHWK\O S\UUROH LQGROH DQG LWV PHWK\O GHULYDWLYHV TXLQROLQHV DQG PHWK\O GHULYDWLYHV WROXLGLQH DQG PHWK\O DQG HWK\O GHULYDWHV DQGSDQLVLGLQH,WZDV DOVRFRQILUPHGE\%DWWHUWRQHW DO D WKDW PDULQH F\DQREDFWHULD DUH YHU\ VHQVLWLYH WR DQLOLQH DQG SWROXLGLQH FRPSRXQGV WKDW DUHEHOLHYHGWREHVHOHFWLYHO\WR[LFWRF\DQREDFWHULD
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rototheca zopfi :DONHUHWDO GRHVXWLOL]HFUXGHRLOQDONDQHV LVRDONDQHV DQG DURPDWLF K\GURFDUERQV PRVW RI WKH ODWHU VWXGLHV ZLWK SKRWRWURSKLF PLFURRUJDQLVPV GHDO HLWKHU ZLWK DOLSKDWLF RU DURPDWLF FRPSRXQGV 7KHUHIRUHVWXGLHVRQWKHWZRFKHPLFDOJURXSVZLOOEH UHYLHZHGVHSDUDWHO\
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hlamydomonas ulvaensis, Chlorella pyrenoidosa DQG Scenedesmus brasiliensis, WKH SK\WRIODJHOODWH
Euglena gracilis DQG WKH F\DQREDFWHULD Anabaena cylindrica DQG Phormidium foveolarum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scillatoria VS DQG Agmenellum quadruplicatum, ZKLFK ZHUH IRXQG WR R[LGL]H QDSKWKDOHQH XQGHU SKRWRDXWRWURSKLF FRQGLWLRQV WR QDSKWKRO &HUQLJLOD HW DO D 7KH DXWKRUV DOVR WHQWDWLYHO\ LGHQWLILHG DPRQJWKHR[LGDWLRQSURGXFWV FLVGLK\GUR[\ GLK\GURQDSKWKDOHQH DQG K\GUR[\WHWUDORQHQ 7KH\ FRQFOXGHG WKDW F\DQREDFWHULD SUREDEO\ SRVVHVV ERWK WKH PRQRR[\JHQDVH DQG GLR[\JHQDVH V\VWHPV WKDW FDWDO\]H WKH YHU\ LQLWLDO VWHS RI QDSKWKDOHQH R[LGDWLRQ :RUNLQJ RQ Oscillatoria VS &HUQLJOLD HW DO F GHPRQVWUDWHG WKDW 2 LV LQFRUSRUDWHG LQWR QDSKWKROUHVXOWLQJIURPQDSKWKDOHQHR[LGDWLRQ 1DUUR VKRZHG WKDW Oscillatoria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gmenellum quadruplicatum, Oscillatoria VS DQG Anabaena VS FRXOG R[LGL]H DQG PHWK\O QDSKWKDOHQH DW WKH PHWK\O JURXS SURGXFLQJ UHVSHFWLYHO\ DQG K\GUR[\PHWK\O QDSKWKDOHQH 7KH DXWKRUV DOVR GHPRQVWUDWHG WKDW 2 ZDV LQFRUSRUDWHG LQWR PHWK\O QDSKWKDOHQH ZKLFK SURGXFHG K\GUR[\PHWK\O QDSKWKDOHQH 7KH R[LGDWLRQ RI PHWK\OQDSKWKDOHQHV E\ F\DQREDFWHULD LV SUREDEO\ VLPLODU WR WKDW PHGLDWHG E\ IXQJDO DQG PDPPDOLDQ HQ]\PHV\VWHPV&HUQLJLODHWDO E ,WVKRXOGEH
665DGZDQDQG5+$O+DVDQ HPSKDVL]HG WKDW DOWKRXJK F\DQREDFWHULD DFKLHYH DURPDWLF ULQJ R[LGDWLRQ RI QDSKWKDOHQH SURGXFLQJ QDSKWKRO WKH\ DSSHDU WR SUHIHU WKH K\GUR[\ODWLRQ RI WKH DON\O VLGHFKDLQ ZKHQ R[LGL]LQJ PHWK\O QDSKWKDOHQHV 7KH R[LGDWLRQ RI ELSKHQ\O E\ F\DQREDFWHULD KDV DOVR EHHQ LQYHVWLJDWHG &HUQLJOLD HW DO E VKRZHG WKDW Oscillatoria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gmenellum quadruplicatum FXOWLYDWHG XQGHU SKRWRWURSKLF FRQGLWLRQV R[LGL]HG SKHQDQWKUHQH SURGXFLQJ WUDQVGLK\GUR[\GLK\GURSKHQ DQWKUHQHDQG PHWKR[\SKHQDQWKUHQH LQDGGLWLRQWR VPDOO DPRXQWV RI SKHQDQWKUROV 8VLQJ 2 LW KDV EHHQ IXUWKHU FRQILUPHG WKDW RQO\ RQH DWRP RI WKH R[\JHQ PROHFXOH ZDV LQFRUSRUDWHG LQWR WKH WUDQV SKHQDQWKUHQH GLK\GURGLRO SURGXFHG E\ WKH F\DQREDFWHULXP ,Q RWKHU ZRUGV D PRQRR[\JHQDVH V\VWHPLVLQYROYHGLQWKHLQLWLDOR[LGDWLYHDWWDFNE\A. quadrupilicatum RQSKHQDQWKUHQH 7KLV ZDV SUREDEO\ WKH ILUVWUHSRUW RIDSURNDU\RWH R[LGL]LQJ DQ DURPDWLF K\GURFDUERQWRDGLK\GURGLRORIWKHWUDQVIRUP $QRWKHU DURPDWLF K\GURFDUERQV JURXS ZKLFK UHFHLYHG WKH DWWHQWLRQ RIUHVHDUFK ZRUNHUV LV WKDW RI DQLOLQH DQ DURPDWLF DPLQH DQG LWV GHULYDWLYHV FRPSRXQGV ZKLFK DUH XVHG LQ LQGXVWU\ IRU WKH SURGXFWLRQ RI G\HV SHVWLFLGHV DQG SKDUPDFHXWLFDOV &HUQLJOLDHWDO IRXQG WKDWWZR DXWRWURSKLFDOO\ JURZQ F\DQREDFWHULD Agmenellum quadruplicatum DQG Oscillutoria VS PHWDEROL]H DQLOLQH SURGXFLQJ IRUPDQLOLGH DFHWDQLOLGH DQG SDPLQRSKHQRO ,W LV NQRZQWKDWEDFWHULD$OH[DQGHU DQGPDPPDOV *RWKRVNDU HW DO DOVR PHWDEROL]H DURPDWLF DPLQHVE\1IRUP\ODWLQJWKHP
$ONDQHV :KLOH PRVW RI WKH ZRUN RQ DURPDWLF K\GURFDUERQV ZDV WDUJHWWHG DW VWXG\LQJ WKH SRWHQWLDO RI F\DQREDFWHULDIRU GHWR[LILFDWLRQ RIWKHVH FRPSRXQGV ZRUN RQ DOLSKDWLF K\GURFDUERQV KDG WKH PDMRU REMHFWLYH RI VWXG\LQJ WKH SRWHQWLDO RI WKHVH
&KDSWHU 2LO3ROOXWLRQ SKRWRWURSKV IRU WKH FRPSOHWH XWLOL]DWLRQ RI WKHVH FRPSRXQGV $V LQ WKH UHVHDUFK RQ DURPDWLF K\GURFDUERQV WKHUH ZDV DOVR QR JXDUDQWHH IRU WKH D[HQLW\ RIWKHF\DQREDFWHULDO FXOWXUHV XVHG LQ VWXGLHV RQ WKH R[LGDWLRQ RIDOLSKDWLF FRPSRXQGV +RZHYHU WKH IHZ VWXGLHV GRQH VR IDU RIIHU LQWHUHVWLQJ UHVXOWV WKDWPHULWDEULHIGHVFULSWLRQ 7KH LQIOXHQFH RI FUXGH RLO DQG LQGLYLGXDO DONDQHV KDV EHHQ WHVWHG RQ WKH QRQD[HQLF VWUDLQV RI Microcoleus chthonoplastes DQG Phormidium corium LVRODWHG IURP RLOULFK VHGLPHQWV $O+DVDQ HW DO 7KH WRWDO OLSLGV IURP M. chthonoplastes JURZQ SKRWRWURSKLFDOO\ ZLWKRXW K\GURFDUERQV FRQWDLQHG PDLQO\ VDWXUDWHG IDWW\ DFLGV ZLWK KH[DGHFDQRLF DFLG DQG RFWDGHFDQRLF DFLG FDUERQ DWRPV 7KH VDPH ZDV WUXH IRU OLSLGV IURP P. corium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nabaena VS DQG Nostoc ellipsosporum, KDYH WKH SRWHQWLDO IRU GHJUDGLQJ WKH KLJKO\ FKORULQDWHG SHVWLFLGH OLQGDQH 7KH WZR F\DQREDFWHULDFRXOG DOVREH JHQHWLFDOO\ HQJLQHHUHGWR GHJUDGHFKORUREHQ]RDWH
( &\DQREDFWHULDDV,PPRELOL]HUVRI2LO 'HJUDGHUV 7KHUH DUH D IHZ UHSRUWV LQGLFDWLQJ WKDW F\DQREDFWHULD PD\ SOD\ DQ LPSRUWDQW DOEHLW LQGLUHFW UROH LQ K\GURFDUERQ GHJUDGDWLRQ LQ QDWXUH E\ LPPRELOL]LQJ RLOGHJUDGLQJ EDFWHULD DQG IXQJL LQ WKHLU PXFLODJH /LJKW KDV EHHQ VKHG RQ WKLV VXEMHFW WKURXJK WKH GLVFRYHU\ RI D QDWXUDO SKHQRPHQRQ DVVRFLDWHG ZLWK H[WHQVLYH RLOSROOXWLRQ DORQJWKH ZHVWHUQ FRDVWRIWKH $UDELDQ *XOI 7KH UHOHDVH RI RLO IURP D .XZDLWL WHUPLQDO LQWR WKH *XOIE\ WKH ,UDTL IRUFHV GXULQJ WKH RFFXSDWLRQ RI.XZDLW $XJXVW )HEUXDU\
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icrocoleus chthonoplastes DQG Phormidium corium, SUHGRPLQDWH ,Q WKH H[WHQVLYH VKHDWK OD\HUV RI ERWK F\DQREDFWHULD XS WR RQH PLOOLRQ EDFWHULDO FHOOV SHU JUDPRIIUHVK PDWZHUH FRXQWHG 7KHVH EDFWHULD FDQ XWLOL]H FUXGH RLO DQG EHORQJ SUHGRPLQDQWO\ WR WKH JHQHUD Rhodococcus, Arthrobacter, Pseudomonas DQG Bacillus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
665DGZDQDQG5+$O+DVDQ SRWHQWLDO RI F\DQREDFWHULD EHFRPHV UHGXFHG XQWLO WR[LF FRQVWLWXHQWV YRODWLOL]H RU DUH ZDVKHG LQWR WKH RSHQ VHD )XUWKHU WKH IDFW WKDW F\DQREDFWHULD LPPRELOL]H LQ WKHLU HQYHORSH OD\HUV RLOGHJUDGLQJ EDFWHULD DQGSUREDEO\KDYHGRQHIRUJHRORJLFDO DJHV SURYLGHV WKH DWWUDFWLYH SRVVLELOLW\ WKDW JHQHWLF LQWHUDFWLRQ FRXOG SRVVLEO\ KDYH RFFXUUHG QDWXUDOO\ EHWZHHQWKH RUJDQRWURSKLF DQGSKRWRWURSKLFSDUWQHUV 7KURXJK WKH FRGLQJ RI WKLV JHQH IRU FDWDO\]LQJ HQ]\PH V\VWHPV K\GURFDUERQ GHJUDGDWLRQ PD\ KDYH EHHQ WUDQVIHUUHG IURP WKH EDFWHULD WR WKH F\DQREDFWHULD &REOH\ HW DO UHSRUWHG RQ WKH FRQVWUXFWLRQ RI D VKXWWOH SODVPLG ZKLFK FDQ EH HIIHFWLYHO\ PRELOL]HG IURP Escherichia coli LQWR WKH F\DQREDFWHULXP Fremyella diplosiphon. ,Q WKLV FRQWH[W ZHKDYHFROOHFWHGWHQVRIF\DQREDFWHULDIURP WKH$UDELDQ*XOIFRDVWVDQGIRXQGWKHPDOODVVRFLDWHG ZLWK RLOGHJUDGLQJ EDFWHULD HYHQ WKRVH VDPSOHV FROOHFWHG IURP QRQSROOXWHG DUHDV )XUWKHU UHVHDUFK LV VWLOO QHHGHG WR JLYH D ILQDO DQVZHU WR WKH TXHVWLRQ ZKHWKHU RU QRW F\DQREDFWHULD PD\ WKHPVHOYHV FRQWULEXWH GLUHFWO\ WR K\GURFDUERQ R[LGDWLRQ
$FNQRZOHGJPHQWV :H WKDQN RXU FRZRUNHUV 0 .KDQDIHU , (O 1HPU $ (O'HVRXN\ DQG $ .KDQ 7KH ILQDQFLDO VXSSRUW E\ .XZDLW 8QLYHUVLW\ WKURXJK WKH UHVHDUFK SURMHFWV6DQG6OLVDFNQRZOHGJHG
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ostoc muscorum. -%LRO&KHP $QGHUVRQ6/DQG0F,QWRVK/ /LJKWDFWLYDWHGKHWHURWURSKLF JURZWK RI WKH F\DQREDFWHULXP Synechocystis VS VWUDLQ 3&& DEOXHOLJKWUHTXLULQJSURFHVV-%DFWHULRO %DWWHUWRQ - :LQWHUV . DQG 9DQ %DDOHQ & D $QLOLQHV VHOHFWLYHWR[LFLW\WREOXHJUHHQDOJDH6FLHQFH %DWWHUWRQ - :LQWHUV . DQG 9DQ %DDOHQ & E 6HQVLWLYLW\ RI WKUHHPLFURDOJDHWRFUXGHRLOVDQGIXHORLOV0DULQH(QYLURQ5HV %RRQ--DQG'H/HHXZ-: 2UJDQLFJHRFKHPLFDODVSHFWVRI F\DQREDFWHULDO PDWV ,Q )D\ 3 DQG 9DQ %DDOHQ & HGV 7KH
&KDSWHU 2LO3ROOXWLRQ &\DQREDFWHULD SS (OVHYLHU 6FLHQFH 3XEOLVKHUV $PVWHUGDP1HZ
)XVH\3DQG2XGRW- 5HODWLYHLQIOXHQFHRISK\VLFDOUHPRYDO DQGELRGHJUDGDWLRQLQ WKHGHSXUDWLRQ RISHWUROHXP FRQWDPLQDWHG VHDVKRUHVHGLPHQWV0DU3ROOXW%XOO *DXU -3 DQG .XPDU +' *URZWK UHVSRQVH RI IRXU PLFURDOJDH WRWKUHH FUXGH RLOV DQG DIXUQDFH RLO (QYLURQ 3ROOXW 6HU$ *DXU -3 DQG 6LQJK $. *URZWK SKRWRV\QWKHVLV DQG QLWURJHQIL[DWLRQ RIAnabaena doliolum H[SRVHGWR$VVDPFUXGH H[WUDFW%XOO (QYLURQ&RQWDP7R[LFRO *LEVRQ '7 0LFURELDO GHJUDGDWLRQ RI DURPDWLF K\GURFDUERQV ,Q *LEVRQ '7 HG 0LFURELDO 'HJUDGDWLRQ RI 2UJDQLF&RPSRXQGVSS0DUFHO'HNNHU1HZ
.OXJ 0- DQG 0DUNRYHW] $- 'HJUDGDWLRQ RIK\GURFDUERQV E\ PHPEHUV RIWKH JHQXV Candida. ,, 2[LGDWLRQ RI QDONDQHV DQGDONDQHVE\Candida lipolytica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
66 5DGZDQDQG5+$O+DVDQ 3HDUFH - /HDFK &. DQG &DUU 1* 7KH LQFRPSOHWH WULFDUER[\OLF DFLG F\FOH LQ WKH EOXHJUHHQ DOJD Anabaena variabilis. -*HQ0LFURELRO 3HOUR\ 5$ DQG %DVVKDP -$ (IILFLHQF\ RI HQHUJ\ FRQYHUVLRQRIDHURELFJOXFRVHPHWDEROLVPLQAphanocapsa -%DFWHULRO 3HOUR\ 5$ 5LSSND 5 DQG 6WDQLHU 5< 7KH PHWDEROLVP RI JOXFRVHE\XQLFHOOXODUEOXHJUHHQDOJDH $UFK0LNURELRO 3HQLJXHO * &RXGHUF 5 DQG 6H\YH & /HV PLFURDOJXHV DFWXHOOHVHWIRVVLOHVLQOHUHWVVWUDWLJUDSKLTXHHWSHWUROLHU %XOOHWLQ &HQWUHV GH5HFKHUFKH([SORUDWLRQ3URGXFWLRQ (OI$TXLWDLQH 3HUU\-- 0LFURELDO PHWDEROLVPRIF\FOLF DONDQHV ,Q $WODV 50 HG 3HWUROHXP 0LFURELRORJ\ SS 0DFPLOODQ 3XEO &R1HZ
&KDSWHU 2LO3ROOXWLRQ 6LNNHPD- 'H %RQW-$0 DQG 3RROPDQ % 0HFKDQLVPV RI PHPEUDQHWR[LFLW\RIK\GURFDUERQV0LFURELRO5HY 6LQJHU 0( DQG )LQQHUW\ :5 0LFURELDO PHWDEROLVP RI VWUDLJKW FKDLQ DQG EUDQFKHG DONDQHV ,Q $WODV 50 HG 3HWUROHXP 0LFURELRORJ\ SS 0DFPLOODQ 3XE &R 1HZ
FKDUDFWHUL]DWLRQ DQGHIIHFWV RQ JURZWK RIPLFURDOJDH 0DU %LRO :RG]LQVNL56DQG&R\OH-( 3K\VLFDOVWDWHRISKHQDQWKUHQH IRUXWLOL]DWLRQE\EDFWHULD$SSO0LFURELRO :RON &3 DQG 6KDIIHU 3: +HWHURWURSKLF PLFUR DQG PDFURFXOWXUHV RI QLWURJHQIL[LQJ F\DQREDFWHULXP $UFK 0LNURELRO
&KDSWHU &\DQREDFWHULDO 'RPLQDQFH LQ WKH 3RODU 5HJLRQV :DUZLFN)9LQFHQW 'pSDUWHPHQW GH ELRORJLH &HQWUH G¶pWXGHV QRUGLTXHV 8QLYHUVLWp /DYDO 6DLQWH)R\ 4XHEHF &DQDGD *. 3
6XPPDU\ , ,QWURGXFWLRQ ,, +DELWDWVDQG&RPPXQLWLHV $ 0DULQH (QYLURQPHQWV % ,FHDQG 6QRZ & 5RFNDQG 6RLOV ' 5LYHUV DQG 6WUHDPV ( /DNHV DQG 3RQGV 3ODQNWRQ %HQWKRV ,O, %LRGLYHUVLW\DQG (QGHPLVP ,9 (FRSK\VLRORJ\RI3RODU&\DQREDFWHULD $ 7HPSHUDWXUH5HODWLRQV % 'HVLFFDWLRQ)UHH]LQJDQG6DOLQLW\7ROHUDQFH & 'HIHQVHV$JDLQVW895 ' /LJKW+DUYHVWLQJDQG3KRWRV\QWKHVLV ( 1LWURJHQ)L[DWLRQ 9 :K\'R&\DQREDFWHULD'RPLQDWH2U1RW " $ 7HPSHUDWXUH(IIHFWV % 6DOLQLW\(IIHFWV & 1XWULHQW 5HTXLUHPHQWV ' *URZWK 5DWHV ( /RVV 5DWHV
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ostoc RURVFLOODWRULDQV 2VFLOODWRULDFHDH DUHDIHDWXUH RIVWUHDPV ODNHV DQG SRQGV LQERWKSRODUUHJLRQV ZLWKH[WUHPH DFFXPXODWLRQV XS WR FPWKLFN DQG ! J &KOD FP DW VRPH VLWHV 3LFRF\DQREDFWHULD RIWHQ GRPLQDWH WKH SK\WRSODQNWRQRISRODUDQG VXESRODUODNHV ,Q WKH FRDVWDO VDOLQH ODNHV RI$QWDUFWLFD SLFRF\DQREDFWHULD DFKLHYH VRPHRIWKHKLJKHVWQDWXUDO FRQFHQWUDWLRQVRQUHFRUG XSWR [ FHOOV P/ +RZHYHU SLFRF\DQREDFWHULDDUH FRQVSLFXRXVO\DEVHQWRUUDUH LQ WKH DGMDFHQWSRODU RFHDQV 7KH HFRSK\VLRORJLFDO FKDUDFWHULVWLFV RIKLJKODWLWXGH F\DQREDFWHULDWKDWFRQWULEXWHWRWKHLUVXFFHVVDQGGRPLQDQFHLQFOXGH DQDELOLW\WRJURZRYHUDZLGHWHPSHUDWXUH B $ Whitton and M Potts (eds). The Ecology of Cyanobacteria, © Kluwer Academic Publishers. Printed in the Netherlands
:) 9LQFHQW
UDQJH EXW DW VORZ UDWHV WROHUDQFH RIGHVLFFDWLRQ IUHH]LQJ DQG VDOLQLW\ VWUHVV D YDULHW\ RI DGDSWLYH VWUDWHJLHV DJDLQVW KLJK OHYHOV RI VRODU UDGLDWLRQ LQFOXGLQJ XOWUDYLROHW UDGLDWLRQ LQ H[SRVHG KDELWDWV DQG DFFOLPDWLRQ WR VKDGHDOORZLQJQHWJURZWKLQSURWHFWHGGLPOLJKWHQYLURQPHQWV ,QPDQ\SRODUKDELWDWVWKHODUJHVWDQGLQJVWRFNV RIF\DQREDFWHULDO ELRPDVV DUHWKHUHVXOWRIJUDGXDO DFFXPXODWLRQ RYHUPDQ\ VHDVRQV ZLWKRQO\ PLQRUORVVHV YLD ELRWLF DQG DELRWLF UHPRYDO SURFHVVHV &\DQREDFWHULD DUH QRW VXFFHVVIXO LQ WKH SRODU RFHDQV ZKHUH VORZ WHPSHUDWXUHGHSUHVVHGDQG OLJKWOLPLWHGJURZWKUDWHV DUHXQDEOH WRNHHS SDFH ZLWK WKH FRQWLQXRXV ORVVHV GXHWR JUD]LQJDGYHFWLRQDQGPL[LQJ
, ,QWURGXFWLRQ “When I persuaded our botanist, Dr. Berggren, to accompany me in the journey over the ice, I joked with him on the singularity of a botanist making an excursion into a tract, perhaps the only one in the world, that was a perject desert as regards botany. This expectation was, however, not confirmed. Dr. Berggren’s keen eye soon discovered, partly on the surface of the ice, partly in the above mentioned [cryoconite] powder, a brown, polycellular alga, which small as it is, together with the powder and certain other microscopic organisms by which it is accompanied, is the most dangerous enemy to the mass of ice, so many thousand feet in height and hundreds of miles in extent.” (p.163 in Leslie, 1879) 7KHVH REVHUYDWLRQV E\ WKH 6ZHGLVK H[SORUHU $GROI (ULN 1RUGHQVNLROG GHVFULEH D UHPDUNDEOH GLVFRYHU\ PDGH GXULQJ KLV H[SHGLWLRQ DFURVV WKH *UHHQODQG ,FH 6KHHW ,Q FHUWDLQ UHJLRQV RI WKH LQWHULRU WKH VXUIDFH RI WKH LFH VKHHW LV FRDWHG E\ ³FU\RFRQLWH´ OLWHUDOO\ ³FROG URFN GXVW´ 7KHVH SDUWLFXODWHVDUHWUDSSHGE\EODFNPXFLODJHSURGXFLQJ F\DQREDFWHULD DQG WKH UHVXOWDQW GDUN SDWFKHV DEVRUE UDGLDWLRQ 7KLV LQ WXUQ LQIOXHQFHV WKH ORFDO KHDW EDODQFH SURGXFLQJ KROHV SRQGV DQG VWUHDPV WKDW DFFHOHUDWH ORFDO PHOWLQJ DQG WKH GHJUDGDWLRQ RI LFH 1RUGHQVNL|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³DQG KHUH DQG WKHUH WKH EHDXWLIXO Phalaropus >QRUWKHUQ
SKDODURSH@ ZDV VHHQ WR SOXFN WKH DOJD Nostoc commune 6HH&KDSWHU ZKLFKLVSOHQWLIXOLQWKHVH ZDWHUV´ S LQ/HVOLH -DPHV 0XUUD\ WKH ELRORJLVW RQ 6KDFNOHWRQ¶V H[SHGLWLRQ WR 5RVV ,VODQG $QWDUFWLFD GXJ WKURXJK WKH LFH RI D IUR]HQ ODNH DQG IRXQG EHQWKLF PDW PDWHULDO ³WKDW RQ FDUHIXO WKDZLQJ UHOHDVHG D PXOWLWXGH RI OLYLQJ WKLQJV IRU VWXG\´ 0XUUD\ QRWDEO\ ILODPHQWRXV F\DQREDFWHULD DQG WKH URWLIHUV ZKLFK IHHG RQ WKHP 7KHVH DQG HDUOLHU FROOHFWLRQV IURP WKH 5RVV 6HD UHJLRQUHYHDOHGPDQ\VSHFLHVRIF\DQREDFWHULDDVZHOO DV GLDWRPV DQG RWKHU RUJDQLVPV )LJ 6RPHWZR \HDUV ODWHU *ULIILWK 7D\ORU D JHRJUDSKHU RQ WKH %ULWLVK $QWDUFWLF ([SHGLWLRQ ZDV VHQW E\ &DSWDLQ 5REHUW)DOFRQ 6FRWWWRH[SORUHWKH ³'U\9DOOH\´ RQH RID VHULHV RIODUJHO\ LFHIUHH YDOOH\V DW ODWLWXGH V LQWKH0F0XUGR6RXQGUHJLRQ $V 7D\ORU FURVVHG RQH RI WKH IUR]HQ ODNHV KH ORRNHG GRZQ DQG PDGH QRWH RIWKH ³H[WHQVLYH ZDWHU SODQWV´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
&KDSWHU 3RODU&\DQREDFWHULD
Fig. 1. 'UDZLQJV RI F\DQREDFWHULD FROOHFWHG IURP 5RVV ,VODQG $QWDUFWLFD ODWLWXGH 6 GXULQJ 6FRWW V 7HUUD 1RYD H[SHGLWLRQ7KHQHZO\ GHVFULEHG WD[D ZHUH Phormidium priestleyi Srhizothrix antarctica DQG 1RVWRFfuscescens YDU mixta )URP3ODWH, )ULWVFK
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
,, +DELWDWVDQG&RPPXQLWLHV $ 0DULQH(QYLURQPHQWV &KURRFRFFRLG IRUPV RI F\DQREDFWHULD JHQHUDOO\ DVFULEHG WR WKH JHQXV Synechococcus DUH ZLGHO\ GLVWULEXWHGWKURXJKRXWWKHZRUOGRFHDQV DQGLQPDQ\ WHPSHUDWH DQG WURSLFDO UHJLRQV WKH\ FRQWULEXWH D PDMRU VRPHWLPHV GRPLQDQW IUDFWLRQ RI WRWDO SK\WRSODQNWRQ ELRPDVV DQG SURGXFWLYLW\ :DWHUEXU\ HW DO 7KH SRODU RFHDQV DUH D QRWDEOH
H[FHSWLRQ ,Q WKH $UFWLF DV ZHOO DV $QWDUFWLF FRQFHQWUDWLRQV RI Synechococcus IDOO WR ORZ YDOXHV RIWHQ EHORZ FHOOV P/ +LJKHU FRQFHQWUDWLRQV RFFXU LQ VHD LFH DOWKRXJK VXFK SRSXODWLRQV PD\ VLPSO\ UHSUHVHQW FHOOV ZKLFK ZHUH WUDSSHG LQ WKH LFH GXULQJIUHH]LQJZLWKOLWWOHin situ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
:)9LQFHQW
Fig. 2. 5HODWLRQVKLSEHWZHHQSLFRF\DQREDFWHULDODEXQGDQFHVROLG FLUFOHV DQGODWLWXGHLQWKH6RXWKHUQ2FHDQ 7KH GDWDDUHIRUWUDQVHFWV RYHU WKH SHULRG 6HSW )HE IURP 7DEOH LQ 0DUFKDQW HW DO $OVR VKRZQ DUH WKH FRUUHVSRQGLQJ ZDWHU WHPSHUDWXUH PHDVXUHPHQWVRSHQFLUFOHV DQGUYDOXHVIRUWKHUHODWLRQVKLSVEHWZHHQYDULDEOHV
VRPH FRQWURYHUV\ RYHU WR ZKDW H[WHQW WKHVH ORZ GHQVLW\ SRSXODWLRQV UHSUHVHQW FRPPXQLWLHV WKDW DUH DFWLYHO\ JURZLQJ in situ YHUVXV SRSXODWLRQV WKDW DUH WUDQVSRUWHG IURP ORZHU ODWLWXGHV E\ DGYHFWLRQ 0LVKLWLQD HW DO UHSRUWHG WKDW SK\FRHU\WKULQ ULFKSRSXODWLRQV ZHUHSUHVHQWDWDOOWLPHVGXULQJWKHLU VDPSOLQJ RI WKH RSHQ ZDWHUV RI WKH %DUHQWV 1RUZHJLDQ DQG *UHHQODQG 6HDV LPSO\LQJ WKDW WKHUH ZHUHUHVLGHQWFRPPXQLWLHV RIDFWLYHO\ JURZLQJ FHOOV Synechococcus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
FHOOV ZLWK LQFUHDVLQJ ODWLWXGH IRU H[DPSOH LQ WKH 1RUWK $WODQWLF 0XUSK\ DQG +DXJHQ DQG WKH *UHHQODQG6HD/HJHQGUHHWDO
% ,FHDQG6QRZ 7KH FU\RFRQLWH IHDWXUHV WKDW ZHUH ILUVW GLVFRYHUHG GXULQJ 1RUGHQVNLROGV H[SHGLWLRQ RYHU WKH *UHHQODQG ,FH&DS DUH QRZNQRZQ WR RFFXU LQWKH ORZHUSDUW RI PDQ\JODFLHUVLQWKHSRODUUHJLRQVDVZHOODVRQDOSLQH JODFLHUV RIWKHWHPSHUDWH]RQH:KDUWRQHWDO 7KH KROHV SRQGV DQG VWUHDPV IRUP LQ WKH ZDVWDJH ]RQH DQG DUH UHDGLO\ FRORQL]HG E\ PLFURRUJDQLVPV SDUWLFXODUO\F\DQREDFWHULD 7KH*UHHQODQGFU\RFRQLWH FRPPXQLWLHV DUHGRPLQDWHG E\ Calothrix parietina, D F\DQREDFWHULXPNQRZQ IURP$QWDUFWLFDDV DSURGXFHU RI WKH EODFN 895DEVRUELQJ SLJPHQW VF\WRQHPLQ 6HH &KDSWHU $OO RIWKH VDPSOHV RI FU\RFRQLWH H[DPLQHG E\ *HQGHO DQG 'URXHW IURP WKH 7KXOH DUHDFRQWDLQHG ILQHPLQHUDOSDUWLFXODWHVORRVHO\ ERXQGE\WKHILODPHQWVRIWKLVF\DQREDFWHULXP &RPSOHPHQWDU\ VWXGLHV RQFU\RFRQLWH FRPPXQLWLHV LQ WKH VRXWK SRODU UHJLRQ VSHFLILFDOO\ RQ WKH &DQDGD *ODFLHULQWKH0F0XUGR'U\9DOOH\VVKRZHGWKDWWKH FRPPXQLWLHV ZHUH GRPLQDWHG E\ ILODPHQWRXV F\DQREDFWHULD VSHFLILFDOO\ Phormidium frigidum, Lyngbya martensiana, Microcoleus paludosus YDU
&KDSWHU 3RODU&\DQREDFWHULD acuminatus and Nodularia harveyana. &\OLQGULFDO FHOOV RI WKH FKURRFRFFDOHDQ Synechococcus aeruginosa ZHUHDOVRFRPPRQ 7KHVHWD[DDUHIRXQG WKURXJKRXWWKHYDOOH\V LQRWKHUWHUUHVWULDO DQG DTXDWLF KDELWDWVDQGWKHFRPPXQLWLHVDUHOLNHO\WREHGHULYHG IURP ZLQG EORZQ PLFURELDO PDWV :KDUWRQ HW DO &\DQREDFWHULD DSSHDU WR EH UHODWLYHO\ UDUH LQ PHOWLQJ VQRZ EDQNV E\ FRPSDULVRQ ZLWK RWKHU SKRWRWURSKV QRWDEO\ WKH FKORURSK\WH Chlamydomonas nivalis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³HVFDSH PHFKDQLVP´ RI ELRPDVV DQG PLQHUDO PDWHULDOV FRXOG UHVXOW LQ DTXDQWLWDWLYHO\ VLJQLILFDQWORVV RIQXWULHQWV IURP WKH ODNHV HDFK \HDU $QRWKHU Phormidium-dominated FRPPXQLW\ LV DOVR IRXQG JURZLQJ ZLWKLQ WKH OLTXLG ZDWHU LQFOXVLRQV RI WKH ODNH LFH DQG DSSHDUV WR EH LQRFXODWHG IURP WKH VXUIDFHE\ZLQGEORZQPDWHULDO3ULVFXHWDO 2QH RI WKH PRVW H[WHQVLYH GHYHORSPHQWV RI F\DQREDFWHULDLQWKHSRODUUHJLRQVRFFXUVRYHUDQDUHD RI NP RQ WKH 0F0XUGR ,FH 6KHOIODW 6 7KLV DEODWLRQ ]RQH FRQVLVWV RI DQ LQWHUFRQQHFWHG FRPSOH[RIPHOWZDWHUODNHVSRQGVDQGVWUHDPVZKLFK FRQWDLQ WKLFN VHYHUDO PP WR FP PLFURELDO PDWV GRPLQDWHG E\ RVFLOODWRULDQV RU OHVV FRPPRQO\ E\ Nostoc commune DQG N. microscopicum ZLWK VXEGRPLQDQW GLDWRP DVVHPEODJHV 9LQFHQW +RZDUG:LOOLDPV HW DO +DZHV HW DO 7KH SRQGV DUH FKHPLFDOO\ GLYHUVH UDQJLQJ IURP GLOXWH ORZ FRQGXFWLYLW\ PHOWZDWHUV 6 FP WR VXOIDWHULFK EULQHV 6HH &KDSWHU ZLWK VHYHUDOWLPHVWKHVDOLQLW\RIVHDZDWHUGH0RUDHWDO 6RPHRIWKH SRQGV DUH LQXQGDWHG E\ D VLQJOH GDLO\WLGH +DZHV HWDO 7KH PLFURELDO PDWV DUHFRUUHVSRQGLQJO\ YDULHG EXWW\SLFDOO\FRQVLVWRIDQ RUDQJHRUUHGVXUIDFHOD\HUZKLFKLVULFKLQFDURWHQRLG
SLJPHQWV DQGDGHHSEOXHJUHHQOD\HUZKLFKKDV KLJK FRQFHQWUDWLRQV RI FKORURSK\OO a &KOa DQG SK\FRF\DQLQ)LJ 6WXGLHVRQWKHSK\VLRORJLFDOHFRORJ\RIWKHLFHVKHOI PDWV UHYHDOHG WKDW WKH\ DFW DV ³FRPSUHVVHG HXSKRWLF ]RQHV´ ZKHUH WKHUH DUH ODUJH FKDQJHV LQ VSHFWUDO TXDOLW\ GRZQ WKURXJK WKH YHUWLFDO SURILOH 4XHVDGD DQG9LQFHQW 8OWUDYLROHWUDGLDWLRQ 895 DQG KLJKHQHUJ\SKRWRV\QWKHWLFDOO\DFWLYHUDGLDWLRQ3$5 DUH VWURQJO\ DWWHQXDWHG E\ WKH VXUIDFH OD\HU RI WKH PDWDQGPRVWRIWKHSKRWRV\QWKHVLV WDNHVSODFHLQWKH ³GHHS &KOa PD[LPXP´ OD\HU '&0 JURZLQJ WRZDUGV WKH EDVH RI WKH SURILOH LQ DQ RUDQJH RU UHG GLPOLJKWUHJLPH 6RPHRIWKHPLFURELDOPDWVFRQWDLQ VSHFLHV FDSDEOH RI JOLGLQJ PRWLOLW\ WKDW YHUWLFDOO\ PLJUDWH WKURXJK WKH OLJKW JUDGLHQW RI WKH PDW LQ UHVSRQVH WR FKDQJHV LQ DPELHQW VRODU UDGLDWLRQ 9LQFHQW HW DO F 9LQFHQW DQG 4XHVDGD 4XHVDGDDQG9LQFHQW
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loeocapsa VSS 7KHGDUNFRORUDWLRQLVGXHWRWKH895VFUHHQLQJ SLJPHQW VF\WRQHPLQ VHH &KDSWHU $OWKRXJKWKLV FRPPXQLW\ FDQ ZLWKVWDQG EULJKW VRODU UDGLDWLRQ DQG
:)9LQFHQW
Fig 3. 6FKHPDWLF GLDJUDP RI YHUWLFDO VHFWLRQV WKURXJK $QWDUFWLF PLFURELDO PDWV GRPLQDWHG E\ F\DQREDFWHULD 7KH VKDGHG DUHD LV WKH EOXH JUHHQFRORUHG'&0$OOSURILOHVKDYHEHHQGUDZQWRWKHVDPHVFDOHIURP9LQFHQWHWDOE
GHVLFFDWLRQ LW DSSHDUV WR EH LQWROHUDQW RI VDOLQH FRQGLWLRQV%URDG\ 7KHFRPPXQLW\LVDEVHQW IRU H[DPSOH IURP 6LJQ\ ,VODQG LQ WKH PDULWLPH $QWDUFWLFUHJLRQDQGIURPFRDVWDODUHDVLQIOXHQFHGE\ VHDVSUD\ +\SROLWKLF FRPPXQLWLHV DUH FRPPRQO\ IRXQG ZKHUHYHU WUDQVOXFHQW VWRQHV RFFXU RYHU WKH VRLO VXUIDFHDVWKH\GR IRUH[DPSOH LQWKH9HVWIROG+LOOV DQG 6FKLUPDFKHU 2DVLV UHJLRQV LQ WKH $QWDUFWLF FRQWLQHQW $W WKH ODWWHU VLWH WKH K\SROLWKLF HQYLURQPHQW LV FRORQL]HG H[FOXVLYHO\ E\ Aphanocapsa endolithica. 7KH 9HVWIROG +LOOV FRPPXQLWLHV DUH PRUH GLYHUVH ZLWK Chroococcidiopsis, Plectonema DQG WZR JUHHQ DOJDO VSHFLHV (Desmococcus DQG Prusiococcus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
FRPPXQLWLHV DUHW\SLFDOO\FRPSRVHG RIF\DQREDFWHULD LQ SDUWLFXODU WKH JHQXV Chroococcidiopsis PLFURVFRSLF OLFKHQV RU WKH JUHHQ DOJD +HPLFKORULV antarctica 1LHQRZ DQG )ULHGPDQQ DQG UHIV WKHUHLQ 7KH FKDVPRHQGROLWKLF FRPPXQLWLHV DSSHDU WR EH PRUH GLYHUVH DQG Chroococcidiopsis FRH[LVWV ZLWKDYDULHW\RIHXNDU\RWLFWD[D 9HUWLFDO ]RQDWLRQLV D IHDWXUH RIPDQ\ RIWKH HQGROLWKLF FRPPXQLWLHV DQG F\DQREDFWHULD W\SLFDOO\ IRUP WKH ORZHUPRVW VWUDWXP ,Q FRDVWDO FKDVPRHQGROLWKLF FRPPXQLWLHV WKH RXWHU JUHHQ OD\HU RI Desmococcus RU Prusiococcus LV XQGHUODLQ E\ DQ LQWHULRU EOXHJUHHQ VWUDWXP RI Chroococcidiopsis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
&KDSWHU 3RODU&\DQREDFWHULD YDULDELOLW\ EHWZHHQ DQG ZLWKLQIURVWSRO\JRQV ZLWK D VHDVRQDO VXFFHVVLRQ IURP FKORURSK\WHV LQ VSULQJ WR ILODPHQWRXVF\DQREDFWHULDLQVXPPHU'DYH\ 7KH PRVW FRPPRQ IRUP LQ WKH 6LJQ\ ,VODQG IHOOILHOG VRLOV LV Phormidium autumnale DQG LW RFFXUV WKURXJKRXWWKHXSSHU PPRIWKHVRLOSURILOH 'DYH\ DQG &ODUNH IRXQG WKDW Nostoc KDG D PHGLDQ GHSWK RI P ZKLOH P. autumnale KDG D PHGLDQ GHSWK RI P 7KH\ LQWHUSUHWHG WKLV GHHSHU ORFDWLRQ RI P. autumnale, DV ZHOO DV WKH PLJUDWLRQ EHKDYLRU RIWKHVH DVVHPEODJHV DV DGDSWLYH UHVSRQVHV WRDYRLGGHVLFFDWLRQ 6RLOVRQWKH$QWDUFWLFFRQWLQHQWWKDWDUHIOXVKHG E\ PHOWZDWHU RIWHQ VXSSRUW YLVLEOH JURZWKV RI F\DQREDFWHULD %URDG\ &RPPRQ IRUPV LQFOXGH Nostoc commune VHH &KDSWHU DQG PHPEHUV RI WKH 2VFLOODWRULDOHV ZLWK VXEGRPLQDQW JHQHUDVXFK DV Nodularia, Calothrix, Scytonema DQG Tolypothrix. :DWHUDYDLODELOLW\DQGVDOLQLW\DSSHDUWR EH WKH PDMRU GHWHUPLQDQWV WKDW LQIOXHQFH GLVWULEXWLRQ RIWKHVHF\DQREDFWHULD )RUH[DPSOH LQWKH9HVWIROG +LOOV KHWHURF\VWRXV F\DQREDFWHULD DUH DEVHQW IURP VRLOVZKLFKDUHPRLVWEXWQRWVXEMHFWWRIORZLQJZDWHU ZKLOH Gloeocapsa EHFRPHV ORFDOO\ DEXQGDQW RQO\ ZKHUHWKHVDOLQLWLHVDUHORZ $QWDUFWLF EU\RSK\WH FRPPXQLWLHV RIWHQ FRQWDLQ DQ DVVRFLDWHG HSLSK\WLF IORUD GRPLQDWHG E\ ILODPHQWRXV DQGFRFFRLGF\DQREDFWHULD 6RPHVSHFLHVIRUPFUXVWV RYHUWKHPRVVHV IRUH[DPSOH Stigonema minutum DW 6FKLUPDFKHU 2DVLV DQG 1RVWRF DQG RVFLOODWRULDQV DW WKH9HVWIROG+LOOVLQDUHDVVXEMHFWWRVHDVSUD\ZKHUH ELUGV DUH SUHYDOHQW WKHVH VXUIDFH FRPPXQLWLHV DUH UHSODFHG E\ JUHHQ FUXVWV RI FKORURSK\WHV %URDG\
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loeocapsa, Schizothrix DQGDWVRPHORFDOLWLHV HJ WKH$OSK5LYHUV\VWHPLQVRXWKHUQ9LFWRULD/DQG WKH
KHWHURF\VWRXV JHQXV Calothrix. ,Q WKH ODWWHU FRPPXQLW\ *ORHRFDSVD LV VXEGRPLQDQW DQG JURZV HSLSK\WLFDOO\ RYHU WKH Calothrix DV ZHOO DV GLUHFWO\ RYHUWKHURFNVXEVWUDWH 7ZRW\SHVRIPLFURELDOPDWVDUHIRXQGLQ$QWDUFWLF VWUHDPV 7KH PRVW FRPPRQ PDW LV GRPLQDWHG E\ RVFLOODWRULDQV SDUWLFXODUO\ Phormidium, Oscillatoria DQG Schizothrix LQZKLFKWKHILODPHQWVDUHHPEHGGHG LQ PXFLODJH ZKHUH WKH\ ELQG WRJHWKHU VLOW DQG VDQG JUDLQV 7KHPDWVRFFXUXSWRVHYHUDOPPLQWKLFNQHVV XS WR J &KOD FP DQG DUH RIWHQ UHG RU RUDQJH FRORUHGZLWKDEOXHJUHHQERWWRPOD\HUWKDWLVULFKLQ SK\FRELOLQ SLJPHQWV HJ (OOLV(YDQV DQG %D\OLVV VHH DOVR )LJ 7KH VHFRQG JURXS RIPDWV LV FRPSRVHG SULPDULO\ RI Nostoc commune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tigonema minutum DQG Plectonema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ivularia minutula, Nostoc commune DQG Tolypothrix tenuis. 2VFLOODWRULDQ FRPPXQLWHV DOVR RFFXU LQ $UFWLF VWUHDPV &URDVGDOH +DPLOWRQ DQG (GOXQG 9p]LQDDQG9LQFHQW 6KHDWKDQG
0OOHU 7KHPDWVDUHUDUHO\DVZHOO GHYHORSHG DQG H[WHQVLYH DV WKRVH LQ $QWDUFWLFD SHUKDSV UHIOHFWLQJWKHLQFUHDVHGJUD]LQJSUHVVXUHLQWKH$UFWLF ,QVHFWV IRU H[DPSOH DUH DEVHQW IURP FRQWLQHQWDO $QWDUFWLFD EXW FDQ DFKLHYH KLJK SRSXODWLRQ GHQVLWLHV LQWKH$UFWLF 'LIIHUHQFHVLQQXWULHQWVXSSO\PD\DOVR FRQWULEXWH WR WKH FRQWUDVW EHWZHHQ $UFWLF DQG $QWDUFWLFPDWFRPPXQLWLHV
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ynechococcus DQG Synechocystis ZHUH QRWHG LQ WKH SODQNWRQ RI WKH SHUPDQHQWO\ LFHFRYHUHG 0F0XUGR 'U\ 9DOOH\ ODNHV GXULQJ WKH ILUVW LQYHVWLJDWLRQ RI WKHLU ELRORJLFDO OLPQRORJ\ *ROGPDQ HW DO $ VXEVHTXHQW VWXG\ GHWHUPLQHG WKDW Synechocystis ZDV D GRPLQDQW FRPSRQHQW RI WKH GHHS &KOa PD[LPXP '&0 LQ /DNH 9DQGD WKH GHHSHVW PRVW ROLJRWURSKLF RI WKH 'U\ 9DOOH\ ODNH VHULHV 9LQFHQWDQG 9LQFHQW )XUWKHUWRWKHQRUWKEXWVWLOOZLWKLQWKH$QWDUFWLF5RVV 6HDVHFWRUVWXGLHVRQWKHODNHVRIWKH7HUUD1RYD%D\ UHJLRQLQGLFDWHG WKDWXSWR RIWKH SK\WRSODQNWRQ &KOD ZDV LQ WKH P IUDFWLRQ &XOWXUHV RI SLFRSODQNWRQ VDPSOHV IURP WKHVH ODNHV FRQWDLQHG Synechococcus DV ZHOODVHXNDU\RWLF VSHFLHV DQGWZR XQLGHQWLILHG WD[D WKDW ZHUH WHQWDWLYHO\ LGHQWLILHG DV SURFKORURSK\WHV $QGUHROL HW DO 3LFRF\DQREDFWHULD DOVR DSSHDUHG WR EH LPSRUWDQW HOHPHQWV RIODNHV RIWKHPDULWLPH ]RQHRI$QWDUFWLFD ,Q ILYH ODNHV RQ 6LJQ\ ,VODQG Synechococcus-OLNH FHOOV ZHUH WKH GRPLQDQW SK\WRSODQNWRQ LQ WHUPV RI FHOO FRQFHQWUDWLRQV DQG &Kla, EXW QDQRSODQNWRQ
:) 9LQFHQW PRVWO\ IODJHOODWHV DQG FKORURSK\WHV GRPLQDWHG WKH WRWDO SULPDU\ SURGXFWLRQ +DZHV (OOLV(YDQV 7KH PRVW GHWDLOHG VWXG\ WR GDWH RQ SLFRF\DQREDFWHULDLQ$QWDUFWLFLQODQGZDWHUVLVIURPD PHURPLFWLF PDULQHGHULYHGODNHLQWKH9HVWIROG+LOOV $FH/DNH ODW 6 Synechococcus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b , EXW DOVR VXJJHVWHG JHQHWLF DIILQLWLHV ZLWK Prochlorococcus 5DQNLQ DQG %RZPDQXQSXEOLVKHGGDWD 3LFRF\DQREDFWHULD DUH DOVR LPSRUWDQW FRPSRQHQWV LQ ROLJRWURSKLF ODNHV RIWKH VXEDUFWLF DQG $UFWLF 6HH &KDSWHUV DQG ,Q WKH FROG VXEDUFWLF ODNHV RI QRUWKHUQ 4XpEHF WKH P IUDFWLRQ W\SLFDOO\ DFFRXQWV IRU RI WKH WRWDO SODQNWRQLF &KOD %HUJHURQ DQG 9LQFHQW DQG LV GRPLQDWHG E\ SLFRF\DQREDFWHULD 6LPLODUO\ LQ $UFWLF ODNHV SLFRF\DQREDFWHULD DUH RIWHQ WKH SK\WRSODQNWRQ GRPLQDQWV $ FRPSDULVRQ RI WKH JURZWK UDWHV DQG SLJPHQW FKDUDFWHULVWLFV RI ILYH LVRODWHV RI Synechococcus IURP $UFWLF ODNHV VKRZHG GLIIHUHQFHV EHWZHHQVWUDLQV DQGHYLGHQFHRIFRQVLGHUDEOHJHQHWLF GLYHUVLW\ ZLWKLQ WKLV FRPSRQHQW RI WKH PLFURELRWD 9p]LQDDQG9LQFHQW 2VFLOODWRULDQ F\DQREDFWHULD DUH RIWHQ IRXQG DV D PDMRU HOHPHQW RI WKH SK\WRSODQNWRQ LQ SRODU ODNHV 0DQ\ RI WKHVH WD[D KDYH H[WUHPHO\ WKLQ WULFKRPHV FD P LQ GLDPHWHU DQG EHFDXVH RI WKHVH PRUSKRORJLFDO FKDUDFWHULVWLFV WKHLU OLJKW DEVRUELQJ SURSHUWLHV DUH DV HIILFLHQW DV VPDOO FRFFRLG RU RYRLG FHOOV VXFK DV WKRVH RI Synechococcus. 7KHVH ILODPHQWRXV SRSXODWLRQV ZHUH IRXQG LQ 'U\ 9DOOH\ ODNHV 9LQFHQWDQG-DPHV DQG LQ$UFWLFSRQGV
&KDSWHU 3RODU&\DQREDFWHULD
Fig 4. 7KH VHDVRQDO GLVWULEXWLRQ RI SLFRF\DQREDFWHULD DW P GHSWK LQ $FH /DNH $QWDUFWLFD 7KH QHW SRSXODWLRQ ORVV RU JURZWK UDWHV ZHUH FDOFXODWHGIURPQDWXUDOORJUHJUHVVLRQVRIWKHFHOOFRXQWGDWDYHUVXVWLPH 5HGUDZQIURP5DQNLQHWDO
9p]LQD DQG 9LQFHQW ,Q SDUW KRZHYHU WKHVH SRSXODWLRQV PD\ UHSUHVHQW ILODPHQWV ZDVKHG LQ IURP VWUHDP EHGV RU WKH OLWWRUDO ]RQH ZKHUH WKLFN PDWV RI F\DQREDFWHULDRFFXU
%HQWKRV 7KH ULFKHVW ELRPDVV DFFXPXODWLRQV RI F\DQREDFWHULD LQ WKH SRODU UHJLRQV RFFXU LQ WKH EHQWKLF KDELWDWV RI ODNHV DQG SRQGV 7KH FRPPXQLWLHV IRUP KLJKO\ SLJPHQWHG OD\HUV RYHU WKH ERWWRP VXEVWUDWD DQG PD\ JUDGXDOO\DFFXPXODWHDVPXFLODJLQRXVILOPVDQGPDWV XS WR VHYHUDO FP RU HYHQ VHYHUDO WHQV RI FP LQ WKLFNQHVV ,Q WKH ODNHV RIWKH0F0XUGR 'U\9DOOH\V UHJLRQ ILYH W\SHV RI EHQWKLF PDW ZHUH GLVWLQJXLVKHG 3DUNHUDQG:KDUWRQ Moat mats ± WKHVH IRUP DURXQG WKH HGJH RIWKH ODNH ZKHUH WKH LFH PHOWV HDFK VHDVRQ 7KH\ DUH FKDUDFWHUL]HG E\ WKLFN VSRQJ\ OD\HUV DQG DUH RIWHQ SLJPHQWHGEULJKWRUDQJHRUEURZQ Columnar lift-off mats ± WKHVH DUH SURGXFHG E\ WKH WUDSSLQJ RI QLWURJHQ DQG R[\JHQ EXEEOHV ZLWKLQ WKH VXUIDFH PDW DQG WKH\ JURZ DV XSULJKW FROXPQDU VWUXFWXUHV 3RUWLRQV RI WKH PDW PD\ EUHDN RII DQG IORDWXSXQGHUWKHSHUPDQHQWLFHFRYHURIWKH ODNH VR WKDW WKH\ EHFRPH LQFRUSRUDWHG LQWR WKH LFH ZKHQ LW
IUHH]HV VHHDERYH ,Q RQH ODNH RIWKH 'U\ 9DOOH\V /DNH )U\[HOO WKH FROXPQDU PDWV SUHFLSLWDWH FDOFLXP FDUERQDWH DQG UHPDLQ LQ SODFH ZLWK D KDUG FDOFLWH LQWHULRU Pinnacle mats ± WKHVH RFFXU WR DW OHDVW P GHSWK ZLWKLQ WKH UHODWLYHO\ ZHOO LOOXPLQDWHG HQYLURQPHQW RI /DNH9DQGDDQG IRUPFRQLFDO VWUXFWXUHV FPKLJK DQGFPZLGH 7KHVH VWUXFWXUHV LQFRUSRUDWH VDQG JUDLQV DQG FDOFLWH FU\VWDOV DQG UHVHPEOH WKH IRVVLO 3UHFDPEULDQVWURPDWROLWHConophyton &KDSWHU Aerobic prostrate mats ±WKHVH RFFXU RYHUWKH VXUIDFH VHGLPHQWVDWGHSWKDQGSUHFLSLWDWHFDOFLWH Anaerobic prostrate mats ± WKHVH RFFXU LQ GHHS DQR[LFEDVLQVRIWKHODNHV 7KHPDWV ZHUH W\SLFDOO\ GRPLQDWHG E\ ILODPHQWRXV F\DQREDFWHULD XVXDOO\ PHPEHUV RI WKH 2VFLOODWRULDFHDH ,Q ODNHV RI WKH 'U\ 9DOOH\V WKH GRPLQDQW IRUP ZDV Phormidium frigidum LQ DOO PDW W\SHV VRPHWLPHV LQ DVVRFLDWLRQ ZLWK Lyngbya PDUWHQVLDQD 0RUH UHFHQW VWXGLHV LQ WKH /DUVHPDQQ +LOOV RI HDVWHUQ $QWDUFWLFDUHYHDOHG DVLPLODUO\ ZLGHGLYHUVLW\ RIPDWW\SHV(OOLV(YDQV DQGUHIVWKHUHLQ D ,Q GHHSHU ODNHV ! P GHSWK EODFN Nostoc PDWV VHYHUDO PPLQ WKLFNQHVV RFFXUUHG WR DGHSWK RI FP
:) 9LQFHQW
E $WJUHDWHUGHSWKNostoc ZDVUHSODFHGE\RUDQJH SLJPHQWHG PDWV XS WR PPWKLFNDQG GRPLQDWHG E\ P. frigidum. 6PDOO VXUIDFH SURWUXVLRQV RI WKH PDW ZHUH DVVRFLDWHG ZLWK RVFLOODWRULDQ ILODPHQWV WKDW RYHUJUHZ VSKHULFDO Nostoc FRORQLHV 7KHVH PL[HG DVVHPEODJHV DUH PRUSKRORJLFDOO\ VLPLODU WR FRPPXQLWLHV UHSRUWHG IURP HOVHZKHUH LQ $QWDUFWLFD VSHFLILFDOO\ DW $EODWLRQ 3RLQW $OH[DQGHU ,VODQG DQG LQSRQGVRQWKH0F0XUGR,FH6KHOIZKLFK FRQWDLQHG Nostoc microscopicum. 7KH VXUIDFH FDURWHQRLG SLJPHQWDWLRQRIWKHPDWVGLPLQLVKHGZLWKGHSWKLQWKH ODNH DQG ZDV DEVHQW IURP FRPPXQLWLHV LQ ZDWHU GHHSHU WKDQ P FRQVLVWHQW ZLWK WKH UROH RI WKHVH SLJPHQWV DV D GHIHQVH DJDLQVW EULJKW VRODU UDGLDWLRQ VHHEHORZ F ,Q ODNHV ZLWK D PD[LPXP GHSWK RI P WKH PDWV IRUPHG FLUFXODU SODWHV FP WKLFN DQG FPLQGLDPHWHU7KHSODWHVDFFXPXODWHGLQWKHERWWRP ZDWHUVIRUPHGDOD\HUDWOHDVWFPLQWKLFNQHVVDQG ZHUH GRPLQDWHG E\ RVFLOODWRULDQV ZLWK Nostoc, Gloeocapsa DQG Chroococcus DV VXEGRPLQDQWV 6LPLODU SODWH RU GLVFVKDSHG JURZWKV RI F\DQREDFWHULDDUHNQRZQWRRFFXULQVKDOORZSRQGVLQ WKH VRXWKHUQPRVW SDUW RI9LFWRULD/DQG WKH 3\UDPLG 7URXJKDUHD G ,Q VKDOORZ SRQGV RI KLJKHU VDOLQLW\ UHG SLJPHQWHG PDWV FRYHUHG WKH ERWWRP DQG ZHUH SHULRGLFDOO\ GHWDFKHG E\ ZLQG 7KHVH PDWV DOVR FRQWDLQHG VSKHULFDO Nostoc FRORQLHV RQ WKHLU XSSHU VXUIDFHEXWZLWKRXWWKHILODPHQWRXVRYHUJURZWK 7KLFN EHQWKLF PDWV RIF\DQREDFWHULD ZHUH UHSRUWHG IURP PDQ\ RWKHU $QWDUFWLF ORFDWLRQV )RU H[DPSOH GLYHUV ZKR H[DPLQHG WKH ERWWRP RI 3UL\DGDUVKDQL /DNH PD[LPXPGHSWK RI P LQ WKH 6FKLUPDFKHU 2DVLV UHJLRQ IRXQG FP WKLFN F\DQREDFWHULDO PDWVWKDWZHUHGRPLQDWHGE\RVFLOODWRULDQVDVZHOO DV Synechocystis DQG Chroococcus ,QJROH DQG 3DUXOHNDU %HQWKLF F\DQREDFWHULD RFFXU LQ DVVRFLDWLRQ ZLWK PRVVHV LQ WKH PDULWLPH ODNHV RI 6LJQ\ ,VODQG 8QOLNH WKH ODNH FRPPXQLWLHV RI FRQWLQHQWDO $QWDUFWLFD WKHVH PDULWLPH F\DQREDFWHULDO FRPPXQLWLHV DUH GRPLQDWHG E\ Phormidium LQ VKDOORZ ZDWHUV DQG E\ Tolypothrix tenuis DQG Plectonema DQG VRPHWLPHV WKH [DQWKRSK\WH Tribonema DWGHSWK(OOLV(YDQV
SRLQWHG WR WKH ODFN RI FRQVLVWHQW WD[RQRPLF FULWHULD LQ DQDO\VHV RI WKH $QWDUFWLF PLFURIORUD LQ JHQHUDO EXW SDUWLFXODUO\ ZLWK UHVSHFW WR WKH F\DQREDFWHULD 0RVW LQYHVWLJDWRUV XVH WKH WUDGLWLRQDO WD[RQRPLF VFKHPH RI *HLWOHU DQG ZRUN ZLWK VDPSOHV RI IUHVKO\ FROOHFWHG RU SUHVHUYHG ILHOG PDWHULDO 6RPHDXWKRUVDGRSWHGWKHKLJKO\VLPSOLILHG 'URXHWFODVVLILFDWLRQLQZKLFKPXFKLQIRUPDWLRQZDV VDFULILFHG )HZVWXGLHV RISRODUUHJLRQV WR GDWHKDYH XVHG WKH FULWHULD RI $QDJQRVWLGLV DQG .RPiUHN WKDW DUH EDVHG RQ PRUSKRORJLFDO IHDWXUHV DQG DWWULEXWHV RIVWUDLQV LQFXOWXUH $ QRWDEOH H[DPSOH RI WKLV DSSURDFK LQ $QWDUFWLFD ZDV WKH ZRUN RI%URDG\ DQG .LEEOHZKLWH VLPLODU VWXGLHV DUH RQO\MXVW EHJLQQLQJLQWKH$UFWLF9p]LQDDQG9LQFHQW 7KH VORZ UDWHV RI VSHFLDWLRQ E\ F\DQREDFWHULD LQ JHQHUDO &DVWHQKRO] LQ FRPELQDWLRQ ZLWK WKH HIILFLHQW GLVSHUVDO DELOLWLHV RI WKLV JURXS RI PLFUR RUJDQLVPV DQG WKH UHODWLYHO\ \RXQJ DJH RI LFHIUHH HQYLURQPHQWVLQWKH$UFWLFDQG$QWDUFWLFVXJJHVWWKDW HQGHPLVP LV OLNHO\ WR EH UDUH DPRQJVW SRODU F\DQREDFWHULD 0RVW RIWKH IRUPV LGHQWLILHG WR GDWH DSSHDU WR EH FRVPRSROLWDQ WD[D +RZHYHU WKH PRUSKRORJLFDO VLPSOLFLW\ RI F\DQREDFWHULD PDVNV D KLJK OHYHO RI JHQHWLF YDULDELOLW\ )RU H[DPSOH Phormidium autumnale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
,,, %LRGLYHUVLW\DQG(QGHPLVP
0DQ\ REVHUYDWLRQV IURP WHPSHUDWH ODNHV DQG ULYHUV VXSSRUW WKH WHQHW WKDW F\DQREDFWHULD SUHIHU ZDUP WHPSHUDWXUHV )RU H[DPSOH EORRPIRUPLQJ VSHFLHV KDYH WHPSHUDWXUH RSWLPD IRU JURZWK DQG SKRWRV\QWKHVLV DERYH & DQG JHQHUDOO\ DFKLHYH
7KHELRGLYHUVLW\RISRODUF\DQREDFWHULDLV DWSUHVHQWD VXEMHFW RI FRQVLGHUDEOH GHEDWH DQG XQFHUWDLQW\ ,Q ODUJH SDUW WKLV UHIOHFWV WKH LQDGHTXDF\ RI FXUUHQW WD[RQRPLF FULWHULD IRU WKHVH RUJDQLVPV %URDG\
,9 (FRSK\VLRORJ\RI3RODU &\DQREDFWHULD $ 7HPSHUDWXUH 5HODWLRQV
&KDSWHU 3RODU&\DQREDFWHULD
:DYHOHQJWKQP Fig. 5. ,Q YLYR DEVRUEDQFH FKDUDFWHULVWLFV RI WKUHH LVRODWHV RI $UFWLF F\DQREDFWHULD ( DQG ( DUH ERWK RVFLOODWRULDQV LGHQWLILHG DV Phormidium autumnale, EXW( FRQWDLQVERWKSK\FRHU\WKULQDQGSK\FRF\DQLQZKLOH( FRQWDLQV RQO\SK\FRF\DQLQ 3LV DSLFRSODQNWRQLF LVRODWHFRQWDLQLQJSK\FRF\DQLQEXWQRWSK\FRHU\WKULQ9p]LQDDQG9LQFHQW
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
UDWHV DW & +RZHYHU UHVSLUDWLRQ UDWHV DOVR LQFUHDVHG DQG LQ WKH GDUN WKH PDWV ZHUH UDSLGO\ GHVWUR\HG E\ EDFWHULDO GHFRPSRVLWLRQ XQGHU ZDUPHU WHPSHUDWXUHV 9LQFHQW DQG +RZDUG:LOOLDPV 6LPLODU DVVD\V IRU SKRWRV\QWKHVLV LQ Phormidium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scillatoria priestleyii LVRODWHG IURP WKH 0F0XUGR ,FH 6KHOIKDG D JURZWK RSWLPXP LQ WKH UDQJH & &DVWHQKRO] DQG 6FKQHLGHU
:)9LQFHQW
Fig. 6. *URZWK YHUVXV WHPSHUDWXUH UHODWLRQVKLSV 7KH EODFN V\PEROV DUH PD[LPXP JURZWK UDWHV PD[ DW RSWLPXP WHPSHUDWXUHV IRU SRODU PDWIRUPLQJF\DQREDFWHULDZLWK7RSW DWRUEHORZ&IURP7DQJHWDO 7KHOLQHLVWKHXSSHUERXQGIRUPDULQHSK\WRSODQNWRQJLYHQ E\ (SSOH\ 7KH RSHQ V\PEROV DUH IRU D PDULQH EDFWHULXP LVRODWHG IURP FRQJHODWLRQ LFH LQ (DVW $QWDUFWLFD IURP %RZPDQ HW DO 7KHRSHQWULDQJOHV DUHIRUSLFRF\DQREDFWHULDOLVRODWHV IURP$UFWLFODNHV JURZQDW &DQGRSWLPDOOLJKW9p]LQD 9LQFHQW 7KHFORVHGWULDQJOHVDUHPD[ DQG7RSW YDOXHVIRUPDULQHGLDWRPVIURPWKH$UFWLFDQG$QWDUFWLF-DFTXHV
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ostoc commune SODFHG LQ WKH GU\ DWPRVSKHUH RQ WKH 0F0XUGR ,FH 6KHOI GHK\GUDWHG UDSLGO\ DQG ZHUH FRPSOHWHO\ GHVLFFDWHG ZLWKLQ KRXUV VHH &KDSWHU 3KRWRV\QWKHVLV DQG UHVSLUDWLRQ ZDV PHDVXUHG LQ WKH PDWV ZLWKLQ PLQXWHV RI UHZHWWLQJ Phormidium-dominated PDWV DSSHDUHG WR EH OHVV WROHUDQW RIGHK\GUDWLRQ DQG WRRN PXFK ORQJHU WR UHFRYHU WKHLU SK\VLRORJLFDO DFWLYLW\ DIWHUUHZHWWLQJ
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
F\DQREDFWHULDO VSHFLHV DUH PRWLOH DQG DUH FDSDEOH RI PLJUDWLRQXSDQGGRZQWKHPDWSURILOHWR DYRLG895 H[SRVXUHZKLOHHQVXULQJDGHTXDWH3$5 IRUH[DPSOH Oscillatoria priestleyii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ostoc FRORQLHVDQG VKHHWV LQWKHVKDOORZZDWHUV RIODNHV DQG SRQGV )LJ VWUHDP FRPPXQLWLHV RIRivularia, Stigonema DQG Scytonema 6KHDWK HW DO EODFN ILOPV DQG FUXVWV RYHU GHVHUW DQG VHPLGHVHUW VRLOV $OHNVDQGURYD DQG Gloeocapsa FUXVWV RQ H[SRVHGURFNIDFHV .RQKDXVHUHWDO 6LPLODU FRPPXQLWLHV RFFXU LQ $QWDUFWLFD DQG LQFOXGH 1RVWRF DWWKH HGJH RIVWUHDPEHGV RU LQ VORZO\ IORZLQJ IOXVK HQYLURQPHQWV Calothrix DQG Gloeocapsa DW WKH HGJH RIVWUHDPV DQG EODFN FUXVWV RYHU VRLOV $ GDUNUHG SLJPHQW RFFXUV LQ WKH VKHDWKV RI Gloeocapsa WKDW IRUPVEURZQFUXVWVRYHUWKHURFNVLQ$UFWLF6KHDWKHW DO DV ZHOO DV $QWDUFWLF VWUHDPV WKLV SLJPHQWDWLRQ ZDV DVFULEHG WR JORHRFDSVLQ EXW LWV :5DEVRUELQJ DQG RWKHU PROHFXODU SURSHUWLHV KDYH \HWWREHVWXGLHG :DWHUVROXEOHSLJPHQWVWKDWDEVRUE 89$% DUH DOVR ZHOO NQRZQ LQ F\DQREDFWHULD SDUWLFXODUO\ P\FRVSRULQHOLNH DPLQR DFLGV VXFK DV DVWHULQD VKLQRULQH SRUSK\UD DQG SDO\WKHQH ZKLFK DEVRUE PD[LPDOO\ LQ WKH UDQJH QP 9LQFHQW DQG 5R\ $TXHRXV H[WUDFWV RI Nostoc commune IURP $ODVND )LJ DQG RVFLOODWRULDQ LVRODWHV IURP WKH 0F0XUGR ,FH 6KHOI 4XHVDGD DQG 9LQFHQW KDYH VSHFWUD WKDW VXJJHVW WKH SUHVHQFH RI WKLV JURXS RI VXQVFUHHQV 7KHVH FRPSRXQGV PD\ EH UHVSRQVLEOH IRU WKH VWURQJ 895 DWWHQXDWLRQ LQ WKH XSSHU IHZ KXQGUHG P RI $QWDUFWLF Phormidium PDWV 4XHVDGD DQG 9LQFHQW F 895 GDPDJH RIELRORJLFDO V\VWHPV FDQ EH WKH UHVXOW RIGLUHFW SKRWRFKHPLFDO GHJUDGDWLRQ RIFHOOXODU
:) 9LQFHQW
Fig. 7. 89VFUHHQLQJFDSDELOLW\RINostoc commune IURP7RROLN/DNH $ODVND &XUYH$LVWKH in vivo DEVRUEDQFHVSHFWUXPFXUYH% LV D VFDQRID PHWKDQROZDWHUH[WUDFWVKRZLQJ WKHSUHVHQFH RIFRPSRXQGV ZLWK WKH DEVRUSWLRQ FKDUDFWHULVWLFV RIP\FRVSRULQHOLNH DPLQR DFLGV FXUYH& LV D VFDQ RID DFHWRQHZDWHUH[WUDFW VKRZLQJ WKHFKDUDFWHULVWLF VSHFWUXP RIVF\WRQHPLQ 9LQFHQW DQG 4XHVDGD 3ODWHH
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
WKH SKRWRV\QWKHWLF DSSDUDWXV 6HH &KDSWHUV DQG 7KHVHUHSDLUPHFKDQLVPVDUHVWLPXODWHGE\ORQJ ZDYHOHQJWK 895 RU EOXH OLJKW 6WXGLHV RQ WZR LVRODWHV RI RVFLOODWRULDQ F\DQREDFWHULD IURP WKH 0F0XUGR ,FH 6KHOI VKRZHG WKDW WKHUH ZHUH ODUJH GLIIHUHQFHV LQ 89% VHQVLWLYLW\ EHWZHHQ VSHFLHV EXW IRU WKH JURZWK RI ERWK VWUDLQV 89$ H[SRVXUH VXEVWDQWLDOO\ RIIVHW WKH LQKLELWRU\ HIIHFWV RI :% 4XHVDGDHWDO
' /LJKW+DUYHVWLQJ DQG3KRWRV\QWKHVLV $UFWLF DQG $QWDUFWLF F\DQREDFWHULD H[SHULHQFH H[WUHPH YDULDWLRQV LQ 3$5 VXSSO\ IURP FRQWLQXRXV OLJKWLQVXPPHUWRFRQWLQXRXVZLQWHUGDUNQHVV 0DQ\ RI WKH FRPPXQLWLHV LQKDELW VXEVXUIDFH HQYLURQPHQWV WKDW UHPDLQ KLJKO\ VKDGHG HYHQ LQ PLGVXPPHU IRU
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p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
JURZWK ZLWK UHODWLYHO\ OLWWOH ELRPDVV ORVV WKURXJK SURFHVVHVVXFKDVJUD]LQJRUVORXJKLQJ
( 1LWURJHQ)L[DWLRQ 1LWURJHQIL[LQJ F\DQREDFWHULD SDUWLFXODUO\ Nostoc commune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ostoc, YDULHG JUHDWO\ EHWZHHQ EU\RSK\WH FRPPXQLWLHV DQG WKHUHZHUHFRQFRPLWDQWYDULDWLRQVLQQLWURJHQIL[DWLRQ UDWHV DV PHDVXUHG E\ WKH DFHW\OHQH UHGXFWLRQ DVVD\ WHFKQLTXH6ROKHLPHWDO 1LWURJHQDVHDFWLYLW\ ZDV VWLPXODWHG JUHDWO\ SUREDEO\ E\ SKRVSKRUXV LQ UHJLRQV JUD]HG E\ JHHVH DQG WKH UDWHV LQ WKHVH DUHDV ZHUH DVPXFK DV RQH RUGHURIPDJQLWXGH KLJKHUWKDQ WKRVHPHDVXUHG LQF\DQREDFWHULDOPDWV LQ WKH ZDUPHU ORZODWLWXGHPLUHV RIWKH VXE $UFWLF .DUDJDW]LGHV HW DO DQG VXE $QWDUFWLF 6PLWK DQG 5XVVHOO
9 :K\'R &\DQREDFWHULD 'RPLQDWH2U 1RW " 7KH ODUJH DQG UDSLGO\ H[SDQGLQJ OLWHUDWXUH RQ KLJK ODWLWXGH F\DQREDFWHULD FRQWLQXHV WR UHLQIRUFH D VWULNLQJGLFKRWRP\LQWKHLUSDWWHUQ RIGLVWULEXWLRQ DQG DEXQGDQFH ,QFROG QRQPDULQHKDELWDWV VXFKDV LFH
:) 9LQFHQW
FDSV SRODU GHVHUW VRLOV JODFLDO VWUHDPV DQG LFH FDSSHG ODNHV F\DQREDFWHULD DUH RIWHQ WKH GRPLQDQW SKRWRWURSKV ,QHQYLURQPHQWVVXFKDV VKDOORZSRQGV DQG URFNIDFHV WKH\ PD\ FRQWULEXWH PRVW RIWKH WRWDO HFRV\VWHP VWDQGLQJ VWRFN ZLWK VSHFWDFXODU ELRPDVV DFFXPXODWLRQV DW VRPH VLWHV
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
% 6DOLQLW\5HVSRQVHV 7KH JUHDWHU SUHVHQFH RI F\DQREDFWHULD LQ SRODU IUHVKZDWHUV YHUVXV PDULQH KDELWDWV PLJKW VLPSO\ UHIOHFWWKHLUOLPLWVRIWROHUDQFHWRVDOLQLW\ :ULJKWDQG %XUWRQ QRWHG WKDW WKH FRPELQHG H[WUHPH RI VDOLQLW\ SOXV ORZ WHPSHUDWXUHV H[HUWV D VHYHUH
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
& 1XWULHQW5HTXLUHPHQWV %ORRPIRUPLQJ DQG SLFRSODQNWRQLF VSHFLHV RI F\DQREDFWHULD DUH H[WUHPH LQ WHUPV RI WKHLU QXWULHQW UHTXLUHPHQWV 7KH IRUPHU JURXS RFFXUV PRVWO\ LQ QXWULHQWULFKHQYLURQPHQWV ZKLOHWKHODWWHUKDVDKLJK QXWULHQWVFDYHQJLQJ DELOLW\ DQG LV PRUH W\SLFDO RI ROLJRWURSKLF FRQGLWLRQV &RQVLVWHQWZLWKWKLV JHQHUDO SDWWHUQ SLFRF\DQREDFWHULD RFFXU WKURXJKRXW WKH ROLJRWURSKLF ODNHV RI ERWK SRODU UHJLRQV KRZHYHU HYHQ LQ HQULFKHG ODNHV RIWKH $UFWLF DQG $QWDUFWLFD EORRPIRUPLQJ VSHFLHV DUH UDUHO\ SUHVHQW 7KHVH ODWWHUIRUPVDUHW\SLFDOO\JDVYDFXRODWHDQGFDQDGMXVW WKHLUSRVLWLRQLQWKHZDWHUFROXPQWKLVVWUDWHJ\ZRXOG QRW EH IDYRUHG LQ WKH FROG XQVWDEOH ZDWHUV RISRODU ODNHV $ FRQVLGHUDWLRQ RI QXWULHQWVL]H UHODWLRQVKLSV VXJJHVWV WKDW SLFRF\DQREDFWHULD ZRXOG KDYH OHVV RID FRPSHWLWLYH DGYDQWDJH XQGHU QXWULHQWUHSOHWH FRQGLWLRQV WKDQ XQGHU FRQGLWLRQV RI QXWULHQW GHILFLHQF\ 1HYHUWKHOHVV SLFRSODQNWRQLF WD[D EXW HXNDU\RWHV DUH RIWHQ D GRPLQDQW FRPSRQHQW RI WKH SK\WRSODQNWRQ LQ WKH QXWULHQWULFK $UFWLF 2FHDQ *UDGLQJHU DQG /HQ] DQG 6RXWKHUQ 2FHDQ 9LQFHQW LQGLFDWLQJWKDWSLFRF\DQREDFWHULDDUH QRWSUHFOXGHG VLPSO\ RQWKHEDVLV RIWKHLU VPDOO VL]H ,Q WKHVH HQYLURQPHQWV WKH VXFFHVV RI SLFRSODQNWRQ VL]HG SKRWRWURSKV LV PRUH OLNHO\ WR EH WKH UHVXOW RI VHOHFWLRQ IRU RUJDQLVPV ZKLFK KDYH D VXSHULRU OLJKW FDSWXULQJ DELOLW\ DQG WKDW DUH OHVV SURQH WR JUD]LQJ ORVVHVWKDQODUJHUFHOOV 7KH DYDLODELOLW\ RI LURQ ZDV LGHQWLILHG DV D SRWHQWLDOO\ LPSRUWDQW FRQWURO RQ SK\WRSODQNWRQ SURGXFWLRQ LQ FHUWDLQ SDUWV RI WKH ZRUOG RFHDQ LQFOXGLQJ WKH VHDV DURXQG $QWDUFWLFD &\DQREDFWHULD DV DJURXS DUHNQRZQ WR KDYH DKLJKUHTXLUHPHQWIRU
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ostoc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
( /RVV5DWHV 6ORZJURZWKFDQRQO\EHRQHFRPSRQHQWRIDQRYHUDOO VWUDWHJ\ IRU PLFURELDO VXFFHVV LW PXVW DOVR EH
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
$FNQRZOHGJPHQWV 7KLV SDSHU ZDV ZULWWHQ ZKLOH RQ VDEEDWLFDO DW $QWDUFWLF &5& 8QLYHUVLW\ RI 7DVPDQLD +REDUW $XVWUDOLD , WKDQN WKH GLUHFWRU DQG VWDII RI WKLV LQVWLWXWH IRU WKHLU VXSSRUW DQG KRVSLWDOLW\ 0\ SRODU UHVHDUFKJURXSLVIXQGHGE\WKH1DWXUDO 6FLHQFHV DQG (QJLQHHULQJ 5HVHDUFK &RXQFLO RI &DQDGD ZLWK ORJLVWLF VXSSRUW IURP &HQWUH G¶pWXGHV QRUGLTXHV DQG 3RODU &RQWLQHQWDO 6KHOI 3URMHFW WKLV LV SXEOLFDWLRQ 3&63e3&3 , WKDQN-RKQ %RZPDQ (YRQQH
7DQJDQG/\QQH3RZHOOIRUWKHLUUHYLHZFRPPHQWVRQ WKHPDQXVFULSW
5HIHUHQFHV $OHNVDQGURYD 9' 9HJHWDWLRQ RI WKH 6RYLHW SRODU GHVHUWV &DPEULGJH 8QLYHUVLW\3UHVV&DPEULGJH $OH[DQGHU 9 6WDQOH\ '0 'DOH\ 5- DQG 0F5R\ &3 3ULPDU\ SURGXFHUV ,Q +REELH -( HG /LPQRORJ\ RI 7XQGUD 3RQGV SS 'RZGHQ +XWFKLQVRQ DQG 5RVV ,QF 6WURXGVEXUJ3$ $OH[DQGHU9 :KDOHQ 6& DQG .OLQJHQVPLWK.0 1LWURJHQ F\FOLQJLQ$UFWLFODNHVDQGSRQGV+\GURELRORJLD $QDJQRVWLGLV . DQG .RPi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sychrobacter glacincola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
:) 9LQFHQW 'DYH\ 0& 7KH HIIHFWV RI IUHH]LQJ DQG GHVLFFDWLRQ RQ SKRWRV\QWKHVLV DQG VXUYLYDO RI WHUUHVWULDO $QWDUFWLF DOJDH DQG F\DQREDFWHULD3RODU%LRO 'DYH\ 0& 7KH VHDVRQDO SHULRGLFLW\ RIDOJDH RQ $QWDUFWLF IHOOILHOGVRLOV+RODUFWLF(FRO 'DYH\ 0& DQG &ODUNH .- 7KH VSDWLDO GLVWULEXWLRQ RI PLFURDOJDH RQ $QWDUFWLF IHOOILHOG VRLOV $QWDUFWLF 6FLHQFH 'DYH\ $ DQG 0DUFKDQW +- 6HDVRQDO YDULDWLRQ LQ QLWURJHQ IL[DWLRQ E\ Nostoc commune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³7HUUD 1RYD´ ([SHGLWLRQ 1DWXUDO +LVWRU\ 5HSRUW %ULWLVK 0XVHXP 1DWXUDO +LVWRU\ S *HLWOHU / &\DQRSK\FHDH ,Q .RONZLW] 5 HG 'U / 5DEHQKRUVW¶V .U\SWRQJDPHQIORUD YRQ 'HXWVFKODQG 2VWHUUHLFK XQG GHU6FKZHL]$NDGHPLVFKH9HUODJVJHVHOOVFKDIW/HLS]LJ *HQGHO 5: DQG 'URXHW ) 7KH F\DQREDFWHULD RIWKH 7KXOH $UHD *UHHQODQG7UDQV$P0LFURVFRS6RF *ROGPDQ &5 0DVRQ '7 DQG +REELH -( 7ZR $QWDUFWLF GHVHUWODNHV/LPQRO2FHDQRJU *UDGLQJHU 5 DQG /HQ] - 6HDVRQDO RFFXUUHQFH RI SLFRF\DQREDFWHULD LQ WKH *UHHQODQG 6HD DQG FHQWUDO $UFWLF 2FHDQ 3RODU%LRO +DPLOWRQ 3% DQG (GOXQG 6$ 2FFXUUHQFH RI Prasiola fluviatilis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
&KDSWHU 3RODU&\DQREDFWHULD +HQU\ *+5 DQG 6YRERGD- 'LQLWURJHQ IL[DWLRQ DFHW\OHQH UHGXFWLRQ LQ KLJK DUFWLF VHGJH PHDGRZ FRPPXQLWLHV $UFWLF $OSLQH5HV +RZDUG:LOOLDPV & 3ULGPRUH 5' %URDG\ 3$ DQG 9LQFHQW:) (QYLURQPHQWDO DQG ELRORJLFDO YDULDELOLW\ LQ WKH 0F0XUGR ,FH 6KHOI HFRV\VWHP ,Q .HUU\ .5 DQG +HPSHO * HGV $QWDUFWLF (FRV\VWHPV (FRORJLFDO &KDQJH DQG &RQVHUYDWLRQSS6SULQJHU9HUODJ1HZ
$QWDUFWLFD 3URFHHGLQJV RI WKH 1DWLRQDO ,QVWLWXWH RI 3RODU 5HVHDUFK6\PSRVLXPRQ3RODU %LRORJ\ 3DUNHU%&6LPPRQV*0:KDUWRQ5$6HDEXUJ.*DQG/RYH)* 5HPRYDORIRUJDQLFDQGLQRUJDQLFPDWWHUIURP $QWDUFWLF ODNHVE\ DHULDOHVFDSHRIEOXHJUHHQDOJDOPDWV -3K\FRO 3DUNHU%&DQG:KDUWRQ5$ 3K\VLRORJLFDOHFRORJ\RIEOXH JUHHQDOJDOPDWVPRGHPVWURPDWROLWHV LQ$QWDUFWLFRDVLVODNHV $UFK+\GURELRO6XSSO 3HUVVRQ 3( &\DQREDFWHULD DQG RIIIODYRXUV 3K\FRORJLD 3ULVFX-&)ULWVHQ&+ $GDPV (( *LRYDQQRQL 6- 3DHUO +: 0F.D\&3'RUDQ 37*RUGRQ'$/DQRLO %'DQG3LQFNQH\ -/ 3HUHQQLDO$QWDUFWLFODNHLFHDQRDVLVIRUOLIHLQDSRODU GHVHUW6FLHQFH 3URWHDX 3- *HQYLFN :+ *DUFLD3LFKHO ) DQG &DVWHQKRO] 5 7KH VWUXFWXUH RI VF\WRQHPLQ DQ XOWUDYLROHW VXQVFUHHQ SLJPHQW IURP WKH VKHDWKV RI F\DQREDFWHULD ([SHULHQWLD 4XHVDGD $ 0RXJHW -/ DQG 9LQFHQW :) *URZWK RI $QWDUFWLF F\DQREDFWHULD XQGHU XOWUDYLROHW UDGLDWLRQ 89$ FRXQWHUDFWV89%LQKLELWLRQ-3K\FRO 4XHVDGD$DQG9LQFHQW:) $GDSWDWLRQRIF\DQREDFWHULDWR WKH OLJKW UHJLPH ZLWKLQ $QWDUFWLF PDWV 9HUK ,QWHUQDW 9HUHLQ /LPQRO 4XHVDGD $ DQG 9LQFHQW :) 6WUDWHJLHV RI DGDSWDWLRQ E\ $QWDUFWLFF\DQREDFWHULDWRXOWUDYLROHWUDGLDWLRQ(XU-3K\FRO 5DQNLQ /0 )UDQ]PDQQ 3' 0F0HHNLQ 7$ DQG %XUWRQ +5 6HDVRQDOGLVWULEXWLRQRISLFRF\DQREDFWHULDLQ$FHODNH D PDULQHGHULYHG $QWDUFWLF ODNH ,Q %DWWDJOLD % 9DOHQFLD - DQG :DOWRQ'HGV $QWDUFWLF&RPPXQLWLHV 6SHFLHV 6WUXFWXUHDQG 6XUYLYDO SS &DPEULGJH 8QLYHUVLW\ 3UHVV 8QLWHG .LQJGRP 5REDUWV 5' DQG =RKDU\ 7 7HPSHUDWXUH HIIHFWV RQ SKRWRV\QWKHWLFFDSDFLW\UHVSLUDWLRQ DQG JURZWKUDWHV RIEORRP IRUPLQJ F\DQREDFWHULD 1HZ =HDODQG -RXUQDO RI 0DULQH DQG )UHVKZDWHU5HVHDUFK 6FKPLGW 6 0RVNDO : GH 0RUD 6- +RZDUG:LOOLDPV & DQG 9LQFHQW :) /LPQRORJLFDOSURSHUWLHV RI$QWDUFWLF SRQGV GXULQJZLQWHUIUHH]LQJ $QW6FL 6HDEXUJ .* 3DUNHU%& :KDUWRQ 5$ DQG 6LPPRQV *0 7HPSHUDWXUH JURZWK UHVSRQVHV RI DOJDO LVRODWHV IURP $QWDUFWLF RDVHV-3K\FRO 6KHDWK 5* DQG 0XOOHU .0 'LVWULEXWLRQ RI VWUHDP PDFURDOJDHLQIRXUKLJK$UFWLFGUDLQDJHEDVLQV $UFWLF 6KHDWK 5* 9LV 0/ +DPEURRN-$ DQG &ROH .0 7XQGUD VWUHDP PDFURDOJDH RI 1RUWK $PHULFD FRPSRVLWLRQ GLVWULEXWLRQ DQG SK\VLRORJLFDO DGDSWDWLRQV +\GURELRORJLD 6PLWK 9 DQG 5XVVHOO $FHW\OHQH UHGXFWLRQ E\ EU\RSK\WH F\DQREDFWHULDDVVRFLDWLRQV3RODU%LRO 6ROKHLP % (QGDO $ DQG 9LJVWDG + 1LWURJHQ IL[DWLRQ LQ $UFWLF YHJHWDWLRQ DQGVRLOV IURP 6YDOEDUG 1RUZD\ 3RODU%LRO 7DQJ (3< 7UHPEOD\ 5 DQG 9LQFHQW :) &\DQREDFWHULDO GRPLQDQFH RI SRODU IUHVKZDWHU HFRV\VWHPV DUH KLJKODWLWXGH PDWIRUPHUVDGDSWHGWRORZWHPSHUDWXUH"-3K\FRO 7D\ORU* :LWK 6FRWW 7KH 6LOYHU/LQLQJ 'RGG 0HDG DQG &R1HZ
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
:) 9LQFHQW
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³VXQVFUHHQ´ ELRFKHPLFDOV ZKLOVWUHWDLQLQJ D FDSDFLW\ IRU VKDGHDGDSWDWLRQ PDNHV WKHP HPLQHQWO\ VXLWHG IRU FRORQLVDWLRQ RI B A Whitton and M Potts (eds). The Ecology of Cyanobacteria, Kluwer Academic Publishers Printed LQ the Netherlands
'':\QQ:LOOLDPV
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± WKHXOWLPDWHGHVHUW
, $+LVWRULFDO 3HUVSHFWLYH &\DQREDFWHULD DUH DQ DQFLHQW JURXS RI SURNDU\RWHV GDWLQJ IURP WKH HDUOLHVW HYROXWLRQ RI SULPLWLYH SKRWRV\QWKHWLF V\VWHPV LQ WKH SULPRUGLDO VRXS RIWKH 3URWHUR]RLF HUD 6FKRSI DQG :DOWHU &KDSWHU (OHYHQ WD[D UHVHPEOLQJ 2VFLOODWRUHDFHDQ ILODPHQWRXV F\DQREDFWHULD KDYH EHHQ GHVFULEHG LQ FKHUW GDWLQJ IURP *\D LQ HDUO\ $UFKHDQ $SH[ %DVDOW LQ WKH :DUUDZRRQD *URXS RI QRUWK ZHVWHUQ$XVWUDOLD6FKRSI 7KH RFFXUUHQFH RI WKHVH IRVVLO F\DQREDFWHULD ZKLFK DUH PLOOLRQ \HDUV ROGHU WKDQ DQ\ FRPSDUDEOH VXLWH RI IRVVLO SURNDU\RWHV NQRZQ DW WKH WLPH VXJJHVWV WKDW 2 ± SURGXFLQJ F\DQREDFWHULDO SKRWRDXWRWURSK\ ZDV IXQGDPHQWDOWROLIHSURFHVVHVRQ(DUWKDWDYHU\HDUO\ VWDJHRILWVHYROXWLRQ 7KHVHPLFURELDOIRVVLOVDUHQRW RQO\ IRXQGDVSUHVHUYHGVWURPDWROLWHVLQVHGLPHQWDU\ GHVHUWURFNVEXWDOVRDVOLYHDQDORJXHVLQEHQWKLFPDWV RI LFHFRYHUHG ODNHV LQ WKH $QWDUFWLF 'U\ 9DOOH\V :DOWHU DQG %DXOG :KDUWRQ 6WDO WKLV YROXPH FKDSWHU &DPSEHOO IRXQG WKDW VRLO DFFUHWLRQ VWDELOLVDWLRQ DQG ELRJHQLF PRGLILFDWLRQ E\ FDOFLXP FDUERQDWHSUHFLSLWDWLQJ WULFKRPHV RI Microcoleus vaginatus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chizothrix, Microcoleus DQG Oscillatoria. 2QH VSHFLHV Schizothrix calcicola, ZDV IUHTXHQWO\ IRXQG DV D VLQJOH SRSXODWLRQ ZKHUH QR RWKHU VSHFLHV ZDV GHWHFWHG E\ WKH LVRODWLRQ SURFHGXUHV XVHG
&KDSWHU &\DQREDFWHULDLQ 'HVHUWV
Fig. 1. $QWDUFWLFD VKRZLQJLFHIUHHDUHDVDQGWKH FDWFKPHQWDQGWLPHDYHUDJHGIORZRIGHVLFFDWLQJNDWDEDWLFZLQGV IURPWKHSRODUSODWHDX DIWHU3DULVK ,FHIUHHUHJLRQVDUHVWLSSOHGDQGDUHDVH[FHHGLQJ NP DUHPDUNHGE\ILOOHGWULDQJOHV 6FDOHEDU NP >DIWHU :\QQ:LOOLDPV@
+RZHYHU FRFFRLG JHQHUD LQFOXGLQJ Anacystis DQG Coccochloris ZHUHDOVRIRXQG&DPHURQ D 7KH WD[RQRP\ RI F\DQREDFWHULD GHVFULEHG E\ &DPHURQ F ZDV VXPPDULVHG UHODWLYH WR RWKHU PLFURELDOJURXSV&DPHURQ D +RZHYHULWZDV RI OLPLWHG YDOXH DV LW ZDV EDVHG RQ WKH JUHDWO\ VLPSOLILHG FODVVLILFDWLRQ SURSRVHG E\ 'URXHW DQG 'DLO\ DQG GHYHORSHG LQ 'URXHW ZKHUHE\ WUDGLWLRQDO ³VSHFLHV´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
SUREOHPV PD\ UHPDLQ XQUHVROYHG ZLWK PDQ\ DPELJXLWLHV XQWLO WKH IXUWKHU DSSOLFDWLRQ RI QHZ PROHFXODU WHFKQLTXHV VXFK DV '1$'1$ K\EULGLVDWLRQ 7KLV KDV EHHQ VXFFHVVIXOO\ XVHG E\ 6WXOS DQG RWKHUV 1LHQRZ DQG )ULHGPDQQ WR GHPRQVWUDWH WKDW HQGROLWKLF VWUDLQV RI Chroococcidiopsis IURP ERWK KRW DQG FROG GHVHUWV EHORQJWRDVLQJOHVSHFLHV &RPSDULVRQ RI7DEOHV DQGIRUVSHFLHVLQKRWDQGFROGGHVHUWVUHVSHFWLYHO\ VKRZV PDQ\ FRPPRQ JHQHUD DQG VHYHUDO FRPPRQ VSHFLHVGHVSLWHKLJKO\GLVSDUDWHWHPSHUDWXUHUHJLPHV 0RUH UHFHQWO\ 7XUQHU KDV UHYLHZHG WKH PROHFXODU V\VWHPDWLFV RI R[\JHQLF SKRWRV\QWKHVLVLQJ EDFWHULD LQFOXGLQJ F\DQREDFWHULD ZLWK VSHFLDO UHIHUHQFH WR WKH XVH RI 6 U51$ DQDO\VLV IRU Chroococcidiopsis DQG UHODWHG JHQHUD %LOOL HW DO KDYH H[WUDFWHG '1$ IURP GHVHUW VWUDLQV RI Chroococcidiopsis WR FRPSDUH WKHLU JHQRPLF VLPLODULW\ ZLWK VSHFLDO UHIHUHQFH WR WKH jtsZ JHQH FRQFHUQHGZLWKFHOOGLYLVLRQ 7KH HFRSK\VLRORJ\ RI F\DQREDFWHULD KDV EHHQ VWXGLHG LQ D ZLGH GLYHUVLW\ RI GHVHUWV &DPHURQ
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loeocapsa FRPPXQLW\ ³PD\ UHSUHVHQW DQ DQDORJRXV VLWXDWLRQ WR FHUWDLQ H[WUDWHUUHVWULDO FRQGLWLRQV´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
'':\QQ:LOOLDPV /LQQDHXV7HUUDFH%ULWLVK$QWDUFWLF6XUYH\ $ /RQJWHUP (FRORJLFDO 5HVHDUFK VLWH QHDU DQRWKHU 666, KDV EHHQ HVWDEOLVKHG LQ 7D\ORU 9DOOH\ DW /DNH +RDUH WR SURPRWH ELRORJLFDO UHVHDUFK LQWR GHVHUW DQG ODNH FRPPXQLWLHV GRPLQDWHG E\ F\DQREDFWHULD :KDUWRQ 7KLVUHYLHZZLOOFRQVLGHUWKHVWUXFWXUHDQGIXQFWLRQ RI GHVHUW F\DQREDFWHULDO FRPPXQLWLHV DQG WKHLU FRQWLQXHGLPSRUWDQFHDVH[RELRORJLFDODQDORJXHV
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hroococcidiopsis GRPLQDWHV FHUWDLQ HQGROLWKLF FRPPXQLWLHV LQ ERWK KRW DQG FROG GHVHUWV )ULHGPDQQ :HVVHOV DQG %XGHO DQG LV VSHFLDOO\UHPDUNDEOH IRU LWV KHDWWROHUDQFH DQG UDWH RIUHK\GUDWLRQUHFRYHU\ DIWHU GHVLFFDWLRQ &DLROD HW DO 'ULHG VSKHULFDO PLFURFRORQLHV FDQ UHPDLQ YLDEOH DIWHU EDNLQJ LQ D KRW DULG HQYLURQPHQW DW VXEVWUDWXP WHPSHUDWXUHV H[FHHGLQJ & 3RWWV +RZHYHU GHVSLWH WKH VWUHVVWROHUDQFH RI
&KDSWHU &\DQREDFWHULDLQ 'HVHUWV Chroococcidiopsis, WKH PRVW IUHTXHQWO\ UHFRUGHG F\DQREDFWHULDOJHQHUDLQKRWGHVHUWVDUHSchizothrix, Plectonema, Microcoleus DQG Nostoc :KLWWRQ 7KHXELTXLW\DQGVXUYLYDOFKDUDFWHULVWLFVRI WKHJHQXVNostoc PHULWLWVLQFOXVLRQDVDVSHFLDOWRSLF E\3RWWVLQWKLVYROXPH&KDSWHU &\DQREDFWHULDFRPSULVHWKH YDVWPDMRULW\ RIWKH PLFURIORUD RI D KRW GHVHUW FUXVW )ULHGPDQQ DQG *DOXQ /DQJH HW DO GHVFULEH WKH VWUXFWXUH DQG WD[RQRPLF FRPSRVLWLRQ RI D W\SLFDO FUXVWFRPPXQLW\ PPWKLFNRQVDQGGXQHVLQWKH 1L]]DQDGLVWULFW RIWKH 1HJHY 'HVHUW ,VUDHO ,W LV GRPLQDWHGE\IRXUJHQHUDRIF\DQREDFWHULDDOWKRXJK WKHPRVWFRPPRQVSHFLHVLV Microcoleus sociatus. 7KLVVSHFLHVWRJHWKHUZLWKVSDUVHUGloeocapsa VSS ZDV IRXQG ZLWKLQ WKH FUXVW ZKLOVW Nostoc VS DQG Calothrix parietina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icrocoleus sociatus XQGHU GLIIHUHQW OHYHOV RI KXPLGLW\ 0D]RU HW DO 1HWSKRWRV\QWKHVLVZDVQRWGHWHFWHGDW UHODWLYHKXPLGLW\5+ EXWZDVVLJQLILFDQWDW 5+ +RZHYHUZDWHUUHWHQWLRQE\M. sociatus ZDV PXFKJUHDWHUWKDQWKDWE\Nostoc, VRFDXWLRQPXVWEH DSSOLHG ZKHQ DVVHVVLQJ GHVLFFDWLRQ SURWHFWLRQ E\ F\DQREDFWHULDO FUXVWV RI GLIIHUHQW VSHFLHV FRPSRVLWLRQ 6FKHUHUDQG=KRQJ VKRZHGWKDW WKH SURSRUWLRQ RI WZR GLVWLQFW HFRW\SHV RI Nostoc commune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
WRFUHDWHDOLIHVXSSRUWLQJQLFKH0D]RUHWDO &DPSEHOO VKRZHG H[SHULPHQWDOO\ WKDW UH ZHWWHG WULFKRPHV RI Microcoleus vaginatus LQ D PLFURSK\WLFFUXVWLQWKH8WDKGHVHUWFRXOGJOLGH WLPHVWKHLURZQOHQJWKLQDGD\OHDYLQJDFRQWLQXRXV (36VKHDWKWKUHDGSRWHQWLDOO\FDSDEOHRIWUDSSLQJDQG ELQGLQJVDQGJUDLQVWRFUHDWHWKHFUXVWIRXQGLQWKH ILHOG )URPVXEVHTXHQWPLFURVFDOHVWXGLHVRIYHUWLFDO VSDWLDO GLVWULEXWLRQ RIF\DQREDFWHULD LQ WKHVH 8WDK FUXVWV *DUFLD3LFKHO DQG%HOQDS FRQFOXGHG WKDW WKH PRWLOLW\ DQG PRUSKRORJ\ RIM. vaginatus KHOSHG WR PDNH LW WKH SULPDU\ FRORQLVHU RI WKHVH GHVHUWVRLOV ,QKRWGHVHUWHQGROLWKLFQLFKHV F\DQREDFWHULDDUH H[SRVHGWRPRUHVXGGHQHQYLURQPHQWDOFKDQJHVWKDQ LQFROGGHVHUWVEHFDXVHRIGDLO\WUDQVLWLRQVIURPKRW GU\FRQGLWLRQVWRZDUPKXPLGRQHVZKHQIRJRUGHZ PD\ FRQGHQVH DW QLJKW 0D]RU HW DO ,QWROHUDQFH RIVXFK IOXFWXDWLRQV OLPLWV WKHVSHFLHV FRPSRVLWLRQ RI HXNDU\RWLF DOJDH LQ 1L]]DQD VDQG 1HJHY 'HVHUW WR VLQJOH PHPEHUV RI WKH JHQHUD Chlorococcum DQG Stichococcus. 3DUDGR[LFDOO\ FKORURSK\WHVDOVRGRPLQDWHPXFKFROGHUGULHUQLFKHV LQWKH%HDFRQ VDQGVWRQH RIVRXWKHUQ9LFWRULD/DQG ZKHUHGHZQHYHURFFXUV)ULHGPDQQ 1LHQRZ DQG)ULHGPDQQ 3DOPHUDQG)ULHGPDQQ
% &ROG'HVHUW(FRV\VWHPV 7KHPRVWH[WHQVLYHDUHDVRIFROGGHVHUWVRLOVRQWKH $QWDUFWLF FRQWLQHQW OLH SULQFLSDOO\ ZLWKLQ WKH NP LFHIUHHDULG]RQH*XQQDQG:DUUHQ LQ VRXWKHUQ 9LFWRULD /DQG 3ODWH D 7KLV DUHD H[WHQGVIURPVHDOHYHOWRFPDOWLWXGH %HFDXVH RI WKH GRPLQDQW ORZWHPSHUDWXUH R[LGDWLRQ DQG VDOLQLVDWLRQDVSHFWVRIWKHLUSHGRJHQHVLV7HGURZDQG 8JROLQL UHFRPPHQGHG WKDW FRQWLQHQWDO $QWDUFWLF VRLOV VKRXOG EH GHILQHG DV ³FROGGHVHUW VRLOV´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
'' :\QQ:LOOLDPV
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nacystis marina, Anacystis montana, Schizothrix calcicola DQG Nostoc commune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ostoc commune DQG Schizothrix calcicola DW WKH SHULSKHU\ RI IUR]HQSRQGV&DPHURQ D 0XFKRIWKH VXUIDFHRIWKH $QWDUFWLF LFHIUHHGHVHUW )LJ LV KLJKO\ PRELOH DQG GLVWXUEHG E\ IUHTXHQW VWURQJ NDWDEDWLF ZLQGV IORZLQJ GRZQKLOO IURP WKH P KLJK SRODU SODWHDX 3DULVK 7KHVH PDLQO\ JUDYLW\GULYHQ ZLQGV PDLQWDLQ WKH DULGLW\ RI LFHIUHH DUHDV E\ IOXVKLQJ WKHP ZLWK FROG GU\ DLU :\QQ:LOOLDPV $V LQ 9LFWRULD9DOOH\ WKLV
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chizothrix calcicola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nabaena, Lyngbya, Microcoleus, Nostoc, Phormidium, Plectonema DQG Scytonema ZHUHWKHPRVWIUHTXHQWO\GRPLQDQWJHQHUD EXW FDXWLRQHG DJDLQVW JHQHUDOLVDWLRQ EHFDXVH RILQWHU VLWHYDULDWLRQV7DEOH &RFFRLGF\DQREDFWHULDVXFK DV Synechococcus DQG Synechocystis ZHUH OHVV
&KDSWHU &\DQREDFWHULDLQ 'HVHUWV Table 1. &\DQREDFWHULDOVSHFLHVFRPPRQO\IRXQGLQVHOHFWHGKRWGHVHUWV
6HH$SSHQGL[LQ&DPHURQDQG%ODQN IRUP\V\QRQ\P 6RLO OLWKRVRO$&KD FKDVPROLWKLF(QG HQGROLWKLF 1RPHQFODWXUHRI'URXHWDQG'DLO\
Table 2. &\DQREDFWHULDOVSHFLHVFRPPRQO\IRXQGLQ$QWDUWLFFROGGHVHUWV
'' :\QQ:LOOLDPV
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hroococcidiopsis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icrocoleus vaginatus (Phormidium autumnale), Nostoc commune, Oscillatoria lutea, Oscillatoria submembranacea DQG Schizothrix calcicola. &RFFRLG VSHFLHV FRPSULVHG Anacystis dimidiata, Anacystis marina, Anacystis montana, Coccochloris aeruginosa, Coccochloris elabens DQG Coccochloris stagnina. 6HDEXUJHWDO JUHDWO\H[SDQGHGWKLV OLVW IRU VRXWKHUQ 9LFWRULD /DQG JLYLQJ DOWHUQDWLYH QDPHV IRUVRPH RIWKH VSHFLHV UHSRUWHG E\&DPHURQ 7DEOH 7KH\ JDYH LOOXVWUDWHG GHVFULSWLRQV RI VSHFLHV DQG GHWDLOV RI WKH KDELWDWV IURP ZKLFK WKH\ ZHUH REWDLQHG DOWKRXJK WKH QRQDTXDWLF KDELWDWV DUH
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¡Q 5\DQ HW DO $ GHWDLOHG VXUYH\ RI WHUUHVWULDO PLFURDOJDH DW (GZDUG 9,, 3HQLQVXOD 0DULH %\UG /DQG 6 : \LHOGHG VSHFLHV RIF\DQREDFWHULD %URDG\ E 2IWKHVH WZHOYHVSHFLHVZHUHORFDOO\DEXQGDQW +RZHYHUPRVW ZHUH DVVRFLDWHG ZLWK SRQGV RU RWKHU VRXUFHV RIPHOW ZDWHU 1HYHUWKHOHVV KH GHVFULEHG WKLQ EODFN FUXVWV RYHU OLWKRVROV GRPLQDWHG E\ WKH F\DQREDFWHULXP Cyanothece aeruginosa DQG WKH FKORURSK\WHV Stichococcus bacillaris ZLWK RFFDVLRQDO Coenocystis VS ,QODQG DW P DVO RQ *MHOYLNVIMHOOD 6 ( (QJHOVNM¡Q GHVFULEHG HQFUXVWHG F\DQRSK\FHDQFKORURSK\FHDQ VXE IRUPDWLRQV EXW RQO\ DVVRFLDWHG ZLWK VHHSDJH WUDLOV IURP PHOWZDWHU 'DUN ILOPV RI Gloeocapsa DQG Phormidium ZHUH IRXQG RQ JQHLVV GHEULV DQG H[WHQVLYH VKHHWV RI Nostoc VS DQG Oscillatoria/Phormidium ZHUH IRXQG RQ ORFDOO\ ZHWWHG PRUDLQLF JUDYHO +RZHYHU WKHUH ZDV QR UHSRUW RI GHVHUW ELRWLF FUXVWV $W QHDUE\
)227127(6 72 7$%/( 23326,7(
LQFOXGLQJVSHFLHVUHSRUWHGQHDUE\E\(QJHOVNM¡Q (SL( HSLOLWKLFFKDVPR FKDVPROLWKLFHQGR HQGROLWKLFVRLO6 OLWKRVRO (,) OLVWHGDVOLWKRELRQWLFLQ)ULHGPDQQHWDO EXWQRWLQ6HDEXUJ 5(& OLVWHGLQ&DPHURQDDQG EXWQRWLQ6HDEXUJ &DPHURQUHSRUWVPhormidium autumnale DV0LFURFROHXV YDJLQDWXV 1RPHQFODWXUHRI'URXHWDQG'DLO\
5REHUWVNROOHQ 6 : NP IURP WKH VHDDW PDVO5\DQHWDO GHVFULEHG PRLVW FOLIIIDFH VLWHV EHDULQJ D ILOP RI Gloeocapsa VS Phormidium VSS DQGNostoc commune. 1RVRLO FUXVWV ZHUH QRWHG DQG PRUH SUROLILF F\DQREDFWHULDO FRPPXQLWLHVZHUHUHVWULFWHGWRVWUHDPVRQO\
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loeocapsa VSS Homoeothrix FI rivularis DQG Stigonema minutum) LQ 3ULQFHVV (OL]DEHWK /DQG DQG 0DF 5REHUWVRQ/DQG%URDG\& Gloeocapsa VSS Calothrix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¡Q DQG GRZQ WR RQO\ ILYH JHQHUD NP LQODQG DW WKH 3HQVDFROD 0RXQWDLQV QHDU 'XIHN 0DVVLI &DPHURQ D 3DUNHUHWDO &ODULGJH
& (QGROLWKLF6SHFLHV $QWDUFWLFHQGROLWKLFF\DQREDFWHULD7DEOH DUHPRUH DEXQGDQW LQ PRLVWHU PRUH QRUWKHUO\ SDUWV RI WKH 0F0XUGR 'U\ YDOOH\V VXFK DV UHSRUWHG DW%DWWOHVKLS 3URPRQWRU\ 3ODWH E LQ WKH &RQYR\ 5DQJH 6 )ULHGPDQQ HW DO DQG LQ FHQWUDO 9LFWRULD /DQG DW 7LPEHU 3HDN 3ULHVWOH\ *ODFLHU
'' :\QQ:LOOLDPV 3ODWH E 5XVVHOO HW DO VXEPLWWHG $OWKRXJK %DWWOHVKLS 3URPRQWRU\ DOVR VXSSRUWV OLFKHQEDVHG FRPPXQLWLHV DQG D VSDUVH RFFXUUHQFH RIHemichloris antarctica JUHHQ DOJD WKUHH F\DQREDFWHULDO FRPPXQLWLHV SUHGRPLQDWH LQ FRQWUDVW WR WKH SUHYDOHQFH RIJUHHQ DOJDH LQ WKH GULHUHQYLURQPHQW DW /LQQDHXV 7HUUDFH 6 1LHQRZ DQG )ULHGPDQQ )ULHGPDQQ HW DO 7KH PRVW FRPPRQ RQH DW %DWWOHVKLS 3URPRQWRU\ LV D HormathonemaGloeocapsa FRPPXQLW\ ZLWK OHVVHU SUHYDOHQFH RI Gloeocapsa DQG Chroococcidiopsis FRPPXQLWLHV LOOXVWUDWHG DQG GHVFULEHG FRPSUHKHQVLYHO\ E\ )ULHGPDQQ HW DO 7KH Hormathonema Gloeocapsa FRPPXQLW\ FKDUDFWHULVWLFDOO\ RFFXUV LQ PRLVW %HDFRQ VDQGVWRQH DOPRVW GHYRLG RISULPDU\ LURQVWDLQLQJ)H2 DWS+ FRPSDUHGZLWK )H2 DWS+ LQGU\LURQVWDLQHG VDQGVWRQH DW /LQQDHXV 7HUUDFH DQG UHVXOWV LQ H[IROLDWLYH ZHDWKHULQJ ,W FDQ DOVR FRQWDLQ Anabaena LQ WKH XSSHU ]RQH DQG LV FKDUDFWHULVHG E\ Aphanocapsa WRJHWKHU ZLWK Gloeocapsa XS WR HLJKW VSHFLHV LQ WKH UHJLRQ Anabaena WKUHH VSHFLHV DQG Lyngbya. 7KH UHG Gloeocapsa FRPPXQLW\ ZKLFK DOVR FRQWDLQV Eucapsis DQG Microchaete LVPDLQO\IRXQGRQYHUWLFDO IDFHV RI ODUJH ERXOGHUV ZKHUH WKH KRUL]RQWDO IDFH LV XVXDOO\ RFFXSLHG E\ OLFKHQGRPLQDWHG FRPPXQLWLHV 7KHLQIUHTXHQW Chroococcidiopsis FRPPXQLW\LVPRUH FKDUDFWHULVWLF RI GULHU H[WUHPH KDELWDWV DOWKRXJK Chroococcidiopsis FDQ DOVREHIRXQG DV DSK\FRELRQW F\DQRELRQW LQ OLFKHQ FRPPXQLWLHV %GHO DQG +HQVVHQ
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
Table 3. 5HTXLUHPHQWVDQGVWUDWHJLHVIRUFRORQLVDWLRQRIGHVHUWVE\F\DQREDFWHULD
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
'' :\QQ:LOOLDPV
% (SLOLWKLF 6XEOLWKLF DQG &KDVPROLWKLF +DELWDWV %\ YLUWXH RI WKHLU PXFLODJLQRXV JURZWK IRUP F\DQREDFWHULD DUH DEOH WR DGKHUH WR WKH VXUIDFH RI URFNV DQG UHWDLQ VSDUVH ZDWHU UHVRXUFHV IURP GHZ LQWHUPLWWHQW UDLQ RU VQRZ DQG PHOWZDWHU LQ FROG UHJLRQV 7KH\FDQORGJHZLWKLQWKHUXJRVLWLHV RIURFN VXUIDFHVLQWKHVXEDHULDOHSLOLWKLFKDELWDWZKLFKDIIRUG VRPHSURWHFWLRQDVWKHPHVRVFDOH 7KLVJURZWKIRUP KDV EHHQGHVFULEHG E\ %URDG\ F DW WKH FRDVWDO VHPLGHVHUW UHJLRQ RI 0DZVRQ 5RFN ZKHUH WKH PLFURIORUD ZDV SUHGRPLQDQWO\ Chroococcidiopsis, Gloeocapsa, Myxosarcina DQG Calothrix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hroococcidiopsis DQG Plectonema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hroococcidiopsis VS RU Gloeocapsa VS +RZHYHU WKH\ DOVR RFFXU LQ WKH $QWDUFWLF LQWHULRU DQG D FKDVPROLWKLF DVVRFLDWLRQ RI Gloeocapsa VS DQGRWKHUF\DQREDFWHULDZDVIRXQGLQ DQRUWKRVLWHLQWKH'XIHN0DVVLI)LJ E\)ULHGPDQQ +H DOVR IRXQG DQ H[DPSOH RI D OLFKHQ GRPLQDWHG FRPPXQLW\ ZLWK D IXQJDO DQG DOJDO ]RQH
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hamaesiphon VS Chroococcus minutus, Dichothrix baueriana, Gloeocapsa sanguinea, Gloeocapsa muralis, Gloeocapsa biformis, Nostoc FI minutum, Oscillatoria FI irrigua DQG Scytonema VS +RZHYHU VSHFLHV GLYHUVLW\ RI HQGROLWKLF F\DQREDFWHULD ZLWKLQ WKH VDQGVWRQH0RVHVFDYH ZDV UHVWULFWHG WR GRPLQDQW Chroococcidiopsis VS ZLWKVSDUVH Synechococcus VS DQG Gloeothece VS
& 7KH(QGROLWKLF+DELWDW +RW'HVHUWV /LJKWPLFURVFRSLF DQG 6(0 VWXGLHV RIWKH HQGROLWKLF KDELWDWEHJDQ LQ KRWGHVHUWV )ULHGPDQQ HW DO )ULHGPDQQ )ULHGPDQQ DQG 2FDPSR )ULHGPDQQ 7KLV UHYHDOHG WKH GRPLQDQW UROH RI WKH F\DQREDFWHULD Gloeocapsa, Chroococcus turgidus DQG Chroococcidiopsis 3ODWH H $ZLGH YDULHW\ RI GHVHUWV DQG WUDQVOXFHQW URFN W\SHV ZHUH IRXQG WR FRQWDLQ FU\SWRHQGROLWKLF F\DQREDFWHULD )ULHGPDQQ DQG .LEOHU 7KHVH LQFOXGHG VDQGVWRQH IURP 6LQDL 'HVHUW 8WDK DQG &DOLIRUQLD 86$ DQG 1DPLELD DQG JUDQLWH IURP 6LQDL DQG &KLOH 3RWWV DQG )ULHGPDQQ LQLWLDWHG VWXGLHV RIWKHHFRSK\VLRORJ\ RIGHVHUWFRPPXQLWLHV VKRZLQJ WKDW VWUDLQV RI GRPLQDQW Chroococcidiopsis )ULHGPDQQ ZHUH OHVV WROHUDQW RIZDWHU VWUHVV WKDQ Chroococcus VWUDLQV $WWKHEHJLQQLQJRIDORQJWHUPVWXG\RIWKHIDFWRUV UHJXODWLQJ PLFURELDO DFWLYLW\ DQG GLYHUVLW\ LQ HQGROLWKLF FRPPXQLWLHV LQ GLYHUVHSDUWV RIWKH ZRUOG )ULHGPDQQ VKRZHG GDLO\ WHPSHUDWXUH YDULDWLRQV IURP WR & LQ JUDQLWH LQ WKH 6RQRUDQ'HVHUW 7KHUH ZDV OLWWOH GLIIHUHQFH EHWZHHQ WKH VXUIDFH DQG PPGHSWK EXW SHDN WHPSHUDWXUHV ZHUHPXFKKLJKHUWKDQWKRVHRIWKHDLU&DW P DERYH WKH URFN 5HODWLYH KXPLGLW\ RI WKH DLU P
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hroococcidiopsis VS DQG Nostochopsis lobatus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ostoc VSS RFFXUUHG QHDU WKH VXUIDFH RI KRW GHVHUW PLFURSK\WLF FUXVWV EXWUHTXLUHG VXQVFUHHQV VXFK DV VF\WRQHPLQ WR VXUYLYH WKH LQWHQVH 89% IOX[ 7KLV LV QRW VXFK D SUREOHP IRU VKHOWHUHG F\DQREDFWHULD LQ HQGROLWKLF FRPPXQLWLHVIURPKRWDQGFROGGHVHUWVLQWKH 0LGGOH (DVW0H[LFR 1RUWK$PHULFD 6RXWK$PHULFD$IULFD DQG $QWDUFWLFD ZKRVH QLWURJHQ EXGJHWV ZHUH VWXGLHG E\ )ULHGPDQQ DQG .LEOHU 7KH\ FRQFOXGHG WKDWELRORJLFDOQLWURJHQIL[DWLRQZDVQRWVLJQLILFDQWLQ HQGROLWKLFFRPPXQLWLHVXQOLNHDOJDOVRLOFUXVWVLQKRW
GHVHUWV VXFK DV *UHDW%DVLQ 'HVHUW ZKLFK IL[HG J 1 P\ 5\FKHUW DQG 6NXMLQV *UHHQILHOG VLPLODUO\ IRXQG 1IL[DWLRQ WR EH QHJOLJLEOH LQ VDQGVWRQH DW %HDFRQ +HLJKWV ZKLOVW LWV WRWDO RUJDQLF FRQWHQW ZDV J P UDQJH J )ULHGPDQQ DQG .LEOHU UHSRUWHG WKH WRWDO RUJDQLF QLWURJHQ FRQWHQW RI URFNV IURP 1RUWK $PHULFDQ DQG 0LGGOH (DVWHUQ GHVHUWV WR UDQJH IURP WR JP ZKLOVW WKDW RI $QWDUFWLF 'U\ 9DOOH\ URFNV IURP D YDULHW\ RI VLWHV ZDV J P )ULHGPDQQ )ULHGPDQQ HW DO VR WKDW QLWURJHQ LV SUREDEO\ QRW UDWHOLPLWLQJ IRU WKH SUHYDLOLQJYHU\VORZ&WXUQRYHU-RKQVWRQDQG9HVWDO 1LWURJHQ IRU GHVHUW HQGROLWKV LV SULPDULO\ GHULYHG DELRWLFDOO\ DV QLWUDWHV DQG DPPRQLD IURP DWPRVSKHULFHOHFWULFDOGLVFKDUJHVDQGFRQYH\HGWRWKH URFN E\ DWPRVSKHULF SUHFLSLWDWLRQ 7KH LQFRPSOHWH 1F\FOH DOVR LQFOXGHV GHFRPSRVLWLRQ RI RUJDQLF PDWWHU WR DPPRQLD UHDVVLPLODWLRQ RI DPPRQLD DPPRQLD YRODWLOLVDWLRQ ORVV RI QLWURJHQ GXULQJ H[IROLDWLRQ EXW UDUHO\ ELRORJLFDO QLWURJHQ IL[DWLRQ )ULHGPDQQDQG.LEOHU
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rebouxia DQG Pseudotrebouxia. 7KHUH LV RIWHQ DOVR D ORZHU ]RQH FRQWDLQLQJ WKH F\DQREDFWHULD Gloeocapsa DQG Chroococcidiopsis LQ DGGLWLRQ WR WKH GRPLQDQW HXNDU\RWLF DOJDH Hemichloris antarctica, Stichococcus VS DQG Heterococcus endolithicus SOXV K\DOLQH IXQJL 3ODWH D 7KH ELRPDVV RIOLFKHQGRPLQDWHG FRPPXQLWLHV GHULYHGIURPDQDO\VLV RIOLSLGSKRVSKDWH LQ VDQGVWRQH IURP /LQQDHXV 7HUUDFH 9HVWDO D LV ORFDOO\ VXEVWDQWLDOJ&PUDQJH WR 7KLVLV D VLJQLILFDQW SRWHQWLDO FRQWULEXWLRQ RI FDUERQ DYDLODEOHIRUKHWHURWURSKLF DFWLYLW\LQWKHROLJRWURSKLF GHVHUW VRLO DQG D VRXUFH RI PLFURELDO LQRFXOXP UHOHDVHG E\ H[IROLDWLRQ 'LVWULEXWLRQ LV SULPDULO\
'' :\QQ:LOOLDPV
ORFDO EXW NDWDEDWLF ZLQGV FDQ GLVWULEXWH UHODWLYHO\ ODUJH SLHFHV RI URFN RYHU PDQ\ NLORPHWUHV ,W LV LPSRUWDQW WR QRWH WKDW WKH SULPDU\ SURGXFWLYLW\ RI WKHVH HXNDU\RWLF DQG SURNDU\RWLF SKRWRWURSKV VXSSRUWV ODUJH DQG GLYHUVH HQGROLWKLF SRSXODWLRQV RI KHWHURWURSKLF EDFWHULD +LUVFKHWDO 6LHEHUW HW DO
9 6WUHVV)DFWRUVLQ$ULG5HJLRQV $ :DWHU'HILFLHQF\ 2I DOO WKH IDFWRUV DVVRFLDWHG ZLWK GHVHUW VXUYLYDO ZDWHUDYDLODELOLW\LVSDUDPRXQW ,QDFRPSUHKHQVLYHO\ UHYLHZ RI SURNDU\RWLF GHVLFFDWLRQ WROHUDQFH 3RWWV HPSKDVLVHG WKH UHPDUNDEOH DELOLWLHV RI F\DQREDFWHULD WR VXUYLYH EH\RQG WKH OLPLWV RI H[WLQFWLRQ RI RWKHU GHVHUW WD[D ,Q D PRUH IRFXVHG VWXG\ RI Chroococcidiopsis DQG Chroococcus, 3RWWV DQG)ULHGPDQQ IRXQGFRQVLVWHQWGLIIHUHQFHVLQ UHVSRQVH WR ZDWHU VWUHVV DQG VXJJHVWHG WKDW WKLV PD\ EH D SK\VLRORJLFDO SURSHUW\ RI F\DQREDFWHULD RI WD[RQRPLF YDOXH 7KH\ FRQFOXGHG WKDW Chroococcidiopsis LV SUREDEO\ WKH PRVW GHVLFFDWLRQ UHVLVWDQWF\DQREDFWHULXPLQH[LVWHQFH +RZHYHU LWLV DEVHQW IURP WKH KLJKDOWLWXGH !P K\SROLWKLF GHVHUW SDYHPHQW KDELWDW RI WKH $WDFDPD GHVHUW DOWKRXJK WKH JHRORJLFDOHGDSKLF FRQGLWLRQV DUH VLPLODUWRRWKHUDUHDVRILWVRFFXUUHQFH 7KLVGHVHUWLV UHQRZQHG IRU LWV YLUWXDO DEVHQFH RI SUHFLSLWDWLRQ PP \ *URVMHDQ HW DO H[FHSW IRU D \HDU F\FOH DVVRFLDWHG ZLWK WKH (O 1Lx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p p
ZKHUH T WHPSHUDWXUH. p YDSRXUSUHVVXUHRIWKHVROXWLRQ po YDSRXUSUHVVXUHRIZDWHUDWWKH VDPHWHPSHUDWXUH 9DOXHV RI DUH H[SUHVVHG DV QHJDWLYH YDOXHV LQ NLOR 3DVFDOV N3D ZKHUH N3D EDU DQG IRU UHIHUHQFH VDWXUDWHG &D&O KDV D 03D Crinalium episammum FDQ VXUYLYH DW D ZDWHU SRWHQWLDO RI 03D DQG Nostoc LV W\SLFDOO\ H[SRVHG WR GHVHUW HQYLURQPHQWV DW 03D GH :LQGHU 3RWWV XQSXEO FLWHG LQ 3RWWV Chroococcus DQG Chroococcidiopsis FDQERWK IL[ &2 DW UHPDUNDEO\ ORZ ZDWHU SRWHQWLDOV EHORZ 03D 3RWWV DQG )ULHGPDQQ 7KH PHDQ ORZHVW ZDWHU SRWHQWLDO UHFRUGHG LQ $QWDUFWLF HQGROLWKLF KDELWDWV LV 03D 1LHQRZ DQG )ULHGPDQQ ZKLOVWWKH HTXLYDOHQW LQ KRW GHVHUW URFNV FRORQLVHG E\ Chroococcidiopsis LV 03D 3DOPHUDQG)ULHGPDQQ 7KHHTXLYDOHQWVWUHVV OLPLWV LQ KRW FUXVWV DUH H[HPSOLILHG E\ 03DIRU SKRWRV\QWKHVLV DQG 03D IRU JURZWK LQ Microcoleus %URFN +RZHYHU DOWKRXJK Chroococcidiopsis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ostoc LQ PRUDLQH SRQGV RQ WKH 0F0XUGR ,FH 6KHOI WKDW DUH VHDVRQDOO\ IUHH]HGULHG UHFRYHU UHVSLUDWRU\ DFWLYLW\ UDSLGO\ RQ UHK\GUDWLRQ DOWKRXJK UHFRYHU\ RI QLWURJHQDVH DFWLYLW\ LV PXFK VORZHU WKDQ WKDW RI SKRWRV\QWKHVLV+DZHVHWDO 3DOPHUDQG)ULHGPDQQ IRXQGWKDWWKH1HJHY 'HVHUW FU\SWRHQGROLWKLF F\DQREDFWHULDO FRPPXQLW\ SKRWRV\QWKHVLVHV RQO\ DW PDWULF ZDWHU SRWHQWLDOV DSSURDFKLQJ WKDW RI OLTXLG ZDWHU ,W LV WKHUHIRUH VLJQLILFDQWWKDWSKRWRV\QWKHVLV E\ Chroococcidiopsis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icrocoleus sociatus IURP KRWGHVHUWFUXVWVEHFDPHLQDFWLYHXQGHUUHODWLYHO\ORZ ZDWHU VWUHVV /DQJH HW DO D VXEVHTXHQWO\ FRQILUPHGWKDWXQOLNHTrebouxia ZKLFKLVDGRPLQDQW DOJDO V\PELRQW RIHQGROLWKLF OLFKHQV M. sociatus GLG QRWVKRZQHWSKRWRV\QWKHVLV XQOHVV OLTXLG ZDWHUIURP GHZZDV DYDLODEOH :DWHUYDSRXUZDV QRWVXIILFLHQW ,Q FRQWUDVW HQGROLWKLF OLFKHQV GRPLQDWHG E\ HXNDU\RWLF DOJDH LQ %HDFRQ VDQGVWRQH DW /LQQDHXV 7HUUDFH LQFRUSRUDWH &2 XVLQJ ZDWHU YDSRXU DORQH DW
D ZDWHU SRWHQWLDO DV ORZ DV 03D 5+ DW & 3DOPHUDQG)ULHGPDQQ
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³FRPSDWLEOH´ VROXWHV RU RVPRSURWHFWDQWV ZKLFKFDQ DFFXPXODWHWR KLJK FRQFHQWUDWLRQV ZLWKRXW D GHWULPHQWDO HIIHFW RQ HQ]\PH HIILFLHQF\ RU PHWDEROLF DFWLYLW\ 7KHLU FROOLJDWLYH SURSHUWLHV SURYLGHSURWHFWLRQDJDLQVWORVVRIWXUJLGLW\ ,QRUJDQLF VROXWHVLQFOXGH. VXFKDV LQKDORSKLOLFAphanothece halophytica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ynechocystis VS IURP VWURPDWROLWHV DW $O.KLUDQ .XZDLW DQG Aphanothece halophytica IURP 6RODU /DNH6LQDL:DUUHWDO ,Q KLV GHWDLOHG DQDO\VLV RI SURNDU\RWLF GHVLFFDWLRQ WROHUDQFH 3RWWV GHVFULEHV D ³SUHIHUHQWLDO H[FOXVLRQ PHFKDQLVP´ ZKHUHE\ FRPSDWLEOH VROXWHV FDQ SURWHFW WKH VKHOO RI ZDWHU URXQG SURWHLQV WR SUHYHQW WKHLU GHQDWXUDWLRQ DQG LQDFWLYDWLRQ XQGHU LQWHUPHGLDWHRVPRWLFVWUHVV +RZHYHU XQGHUH[WUHPH ZDWHUGHILFLW RQO\SRO\K\GUR[\ FRPSRXQGV VXFK DV WKH GLVDFFKDULGHV VXFURVH DQG WUHKDORVH FDQ SUHYHQW GHQDWXUDWLRQ RI SURWHLQV E\ UHSODFLQJ WKH VKHOO RI
ZDWHU DURXQG WKH PDFURPROHFXOHV 7KLV IXQFWLRQ LV GHVFULEHG E\ D ³ZDWHU UHSODFHPHQW K\SRWKHVLV´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hroococcidiopsis VS +HUVKNRYLW] HW DO )ULHGPDQQ DQG 2FDPSR)ULHGPDQQ VXSSRUWV WKHLU UROH LQ GHVLFFDWLRQ WROHUDQFH HVSHFLDOO\ LQ FROG GHVHUWV ZKHQWKH\ DUH DOVR QHHGHG DV FU\RSURWHFWDQWV 0RQWLHODQG&RZDQ $FFXPXODWLRQV RIGLVDFFKDULGH VXJDUV DUH FRPPRQ LQ GHVHUW F\DQREDFWHULD 8QGHU H[SHULPHQWDO ZDWHU SRWHQWLDO VWUHVV RI F 03D GHVLFFDWLRQ UHVLVWDQW VWUDLQV RIPhormidium autumnale IURP WKH 'DQ DUHD VHZDJHSODQW ,VUDHO DQG Chroococcidiopsis VS IURP GHVHUW VDQGVWRQH IURP 0DNKWHVK 5DPRQ ,VUDHO DFFXPXODWHG XS WR J DQG J WUHKDORVH J FKORURSK\OO UHVSHFWLYHO\ +HUVKNRYLW] HW DO 7KH\DOVRDFFXPXODWHGJDQG JVXFURVHJ FKORURSK\OO UHVSHFWLYHO\ $W KLJKHU ZDWHU SRWHQWLDO WKH FRQFHQWUDWLRQ GHFUHDVHG SUREDEO\ EHFDXVH PHWDEROLF DFWLYLW\ ZDV EHLQJ GLVUXSWHG E\ WKH VWUHVV LWVHOI 1RQGURXJKW UHVLVWDQW VWUDLQV (Plectonema boryanum DQG D Synechococcus VWUDLQ GLG QRW DFFXPXODWHHLWKHUVXJDUXQGHUWKHVDPHFRQGLWLRQV
& 7HPSHUDWXUH 7KH JURZWK DQG SURGXFWLYLW\ RIGHVHUW F\DQREDFWHULD GHSHQGVQRWRQO\RQPRLVWXUHEXWDOVRRQWHPSHUDWXUH DQG OLJKW 7KHUROH RIWHPSHUDWXUH LV FRQQHFWHG ZLWK WKDW RI ZDWHU EHFDXVH RI WKH IUHH]LQJ WUDQVLWLRQ +RZHYHU WKH SUHVHQFH RIFRPSDWLEOH VROXWHV H[WHQGV WKHLUPHWDEROLFZLQGRZWR VXE]HURWHPSHUDWXUHV DQG .DSSHQ DQG )ULHGPDQQ IRXQG WKDW PHWDEROLF
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in situ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rebouxia DQG Hemichloris antarctica WKH\ REWDLQHG IURP VRLO DQGURFNVDW/LQQDHXV7HUUDFH2FDPSR)ULHGPDQQHW DO IRXQG WKDW WKH F\DQREDFWHULXP Chroococcidiopsis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icrocoleusGRPLQDWHG VRLO FUXVWIURP WKH VDPH GHVHUWUHJLRQ KDV DPHDQ&9 GDLO\PD[LPXPQHWSKRWRV\QWKHVLV RI PRO &2 P V PJ &2 P K /DQJH HW DO *DUFLD3LFKHO DQG %HOQDS XVHG UDSLGUHVSRQVHILEUHRSWLF SUREHV DQG PLFURHOHFWURGHV WR PRQLWRU PLFURVFDOH HQYLURQPHQWDO JUDGLHQWV DQG DVVRFLDWHG ELRPDVV DQG SULPDU\ SURGXFWLYLW\ RI WZR GHVHUW VRLO FUXVWV IURP 8WDK86$ $ UHG FUXVW GRPLQDWHG E\ Microcoleus vaginatus DQG Nostoc VS VSDUVH ZLWKDGU\ ELRPDVV RI PJ P PJ &KO P KDG D FRPPXQLW\ QHWSURGXFWLYLW\6( RI PPRO2 P K ZKLOVW D GHQVHU EODFN FUXVW ELRPDVV PJ P &KOPJP FRQWDLQLQJ DEXQGDQWM. vaginatus, Nostoc VS DQG Scytonema VS SURGXFHG PPRO 2 P K &RQVLGHULQJ WKH GLIIHUHQW PHWKRGRORJLHV XVHG WKHVH YDOXHV DUH UHPDUNDEO\
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
' /LJKWDQG89UDGLDWLRQ 3KRWRV\QWKHWLFDOO\$FWLYH 5DGLDWLRQ )URP WKH HDUOLHVW HFRSK\VLRORJLFDO VWXGLHV RI GHVHUW HQGROLWKLF FRPPXQLWLHV OLJKW KDV EHHQ FRQVLGHUHG WR EH WKH SULPH IDFWRU IRU VWUDWLILFDWLRQ )ULHGPDQQ )ULHGPDQQ HW DO VKRZHG WKDW OLJKW WHPSHUDWXUH DQG PRLVWXUH FRQGLWLRQV ZHUH IDYRXUDEOH IRUPLFURELDOPHWDEROLVPIRUEHWZHHQ WRK\ DW /LQQDHXV 7HUUDFH )LHOG GDWD DQG VDPSOHV IURP /LQQDHXV 7HUUDFH KDYH EHHQ XVHG WR YDOLGDWH D OLJKW UHJLPH PRGHO 1LHQRZ HW DO E 7KH\ FRQFOXGHG WKDW WZR DVSHFWV RI OLJKW IOX[ UHJXODWHG VWUDWLILFDWLRQ H[FHVVLYH OLJKW UHVXOWV LQ SKRWR R[LGDWLYH EOHDFKLQJ ZKHUHDV DW WKH ORZHU HQG WKHUH LV LQVXIILFLHQW OLJKW HQHUJ\ WR VXVWDLQ SKRWRV\QWKHWLF DFWLYLW\DQGJURZWK ,WLVNQRZQWKDWGDUNSLJPHQWVLQ WKHZDOOVRIKRWGHVHUWIXQJLDQGLQWKHVKHDWKVRIWKHLU
HXNDU\RWLF SK\FRELRQWV SURWHFW DJDLQVW SKRWR EOHDFKLQJ )ULHGPDQQ DQG *DOXQ 7KLV LV DQDORJRXVWRWKHGHFUHDVHLQVKHDWKSLJPHQWDWLRQZLWK GHSWK LQ HQGROLWKLF F\DQREDFWHULD DV VHHQ LQ WKH 1HJHY 'HVHUW )ULHGPDQQ +RZHYHU Chroococcidiopsis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emichloris antarctica ]RQH DQG PRO DW LWV ORZHU ERXQGDU\ ZKHUH F\DQREDFWHULD PLJKW EH H[SHFWHG 0RUH OLJKW SHQHWUDWHV WKH URFN ZKHQ LW LV ZHW EXWHYHQ WKHQ WKH SKRWRQ IOX[ DW WKH ERWWRP RI WKH Hemichloris ]RQH RQO\ UHDFKHV PRO P -2 \ 7KH SKRWRQ IOX[ RI PRO P V GU\ RU HYHQ PRO P V ZKHQ ZHW LV EHORZ WKH FRPSHQVDWLRQ SRLQW RI PRVW DOJDH +RZHYHU F\DQREDFWHULD DUH W\SLFDOO\ FDSDEOH RI VKDGHDGDSWDWLRQ DQG Phormidium frigidurn-Lyngbya martensiana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ostoc VSS IURP GLYHUVH JHRJUDSKLFDO ORFDWLRQV PD\ ZHOO SURYLGHSDUW RIWKLV
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
8OWUD 9LROHW5DGLDWLRQ 0XFK KDV EHHQ SXEOLVKHG RQ WKH HIIHFWV RI89% RQ F\DQREDFWHULD:\QQ:LOOLDPV *DUFLD3LFKHO EXW OLWWOH RQ WKH HIIHFWV RI 89% RQ GHVHUW FRPPXQLWLHV +RZHYHU F\DQREDFWHULD UHODWHG WR GHVHUW VWUDLQV KDYH EHHQ VWXGLHG IRU WKHLU SURGXFWLRQ 7KHVH LQFOXGH RI 89DEVRUELQJ SLJPHQWV VF\WRQHPLQ LQ WKH (36 VKHDWK RI F\DQREDFWHULD VXFK DV Nostoc commune *DUFLD3LFKHO DQG &DVWHQKRO] (KOLQJ6FKXOW] HWDO DQG P\FRVSRULQH OLNH DPLQR DFLGV 0$$V ZLWKLQ WKH FHOOV RI Gloeocapsa VSS 6FKHUHUHW DO *DUFLD3LFKHO HWDO 6F\WRQHPLQKDV DOVR EHHQIRXQG LQ \HOORZLVKEURZQ FRORXUHG Calothrix parietina LQ Microcoleus FUXVWV RQ GXQH VRLOV RI WKH 1L]]DQD GLVWULFW 1HJHY 'HVHUW /DQJH HW DO *DUFLD 3LFKHO DQG %HOQDS FRQFOXGHG WKDW WKH LQYHVWPHQW ZKLFK VXUIDFHERXQG F\DQREDFWHULD PDNH LQ V\QWKHVLVLQJ VF\WRQHPLQ LQGLFDWHV WKH HFRORJLFDO LPSRUWDQFH RI FRORQLVLQJ RI VXFK H[WUHPH HQYLURQPHQWV ,QSRODUGHVHUWVWKHUHZLOOEHPHWDEROLF FRPSHWLWLRQ IRU ELRFKHPLFDO UHVRXUFHV WR GHDO ZLWK HQYLURQPHQWDO VWUHVVHV 'XULQJ WKH ODWH ZLQWHU ³2]RQH +ROH´ VRPH RI WKH FDUERQ DOORFDWHG WR FU\RSURWHFWDQWVPD\QHHGWREHUHWULHYHGIRUV\QWKHVLV LI 89%SURWHFWDQWV VXFK DV VF\WRQHPLQ RU 0$$V 7KLVLVDQDVSHFWZKLFKUHTXLUHVIXUWKHUUHVHDUFK
9, %LR:HDWKHULQJ DQG 1XWULHQW $YDL,DEL,LW\ $IHDWXUHRIURFNVFRORQLVHGE\HQGROLWKLFPLFUREHVLV WKH FKDUDFWHULVWLF PRVDLF HIIHFW UHVXOWLQJ IURP H[IROLDWLRQ )ULHGPDQQ DQG 2FDPSR)ULHGPDQQ 3DUWRIWKHPHFKDQLVPRIWKLVSURFHVVGHSHQGV RQWKHSK\VLFDOHIIHFWVRIKHDWLQJDQG FRROLQJDQGWKH
&KDSWHU &\DQREDFWHULDLQ 'HVHUWV H[SDQVLRQ DQG FRQWUDFWLRQ RI FRQGHQVHG ZDWHU DV LW FU\VWDOOLVHV ZLWKLQ WKH IDEULF RI WKH URFN &DPSEHOO DQG&ODULGJH +RZHYHU WKH VFDUFLW\ RIOLTXLG ZDWHU LQ HQGROLWKLF FRPPXQLWLHV PDNHV IUHH]HWKDZ GLVUXSWLRQ DQ DOPRVW QHJOLJLEOH IDFWRU 7KH DEVHQFH RIFOHDYDJHSODQHVLQVDQGVWRQHPDNHVFUXPEOLQJLQWR LQGLYLGXDO JUDLQV WKHLU PDLQ GLVLQWHJUDWLRQ UHVSRQVH 6DQGVWRQH OD\HUV DV GLVWLQFW IURP FOHDYDJH SODQHV DUH KRZHYHU VSOLW DSDUW E\ WKH IUHH]HWKDZ DFWLRQ RI VQRZ PHOWZDWHU DQG ZLQG HURVLRQ 7KLV LV RQ D GLIIHUHQW VFDOH IURP WKH H[IROLDWLRQ DSSDUHQWO\ DVVRFLDWHG ZLWK HQGROLWKLF FRPPXQLWLHV 7KHUPDO GLVUXSWLRQ LV XQOLNHO\ WR EH D SULPH FDXVH RI HQGROLWKLF H[IROLDWLRQ LQ VDQGVWRQH VLQFH &RRNH DQG 6PDOOH\ FRQFOXGHG WKDW H[SDQVLRQ DQG FRQWUDFWLRQ RI URFN PLQHUDOV DV D UHVXOW RI GDLO\ WHPSHUDWXUH FKDQJHV LQ GHVHUWV DUH E\ WKHPVHOYHV LQDGHTXDWH FDXVHV RI URFN GLVLQWHJUDWLRQ 6DOW GLVLQWHJUDWLRQ LV OLNHZLVH XQOLNHO\ WR EH LQIOXHQWLDO LQ WKH VWUDWLILHG ZHDWKHULQJ REVHUYHG LQ WKHVH HOHYDWHG VXEVWUDWDGLVWDQW IURP VHHSDJH DQG VDOWDFFXPXODWLRQ ]RQHV 2EVHUYDWLRQV RIWKH H[IROLDWLRQ RIHQGROLWKLFV VKRZV WKDW WKH\ WHQG WR VSOLW DORQJ WKH OD\HU FRQWDLQLQJ WKH PLFURELDO FRPPXQLW\ ZKLFK VXJJHVWV WKDW WKLV LV D ]RQH RI LQIOXHQFH DQG VWUXFWXUDO ZHDNQHVV 7RLQYHVWLJDWHWKH VROXEOLVLQJDFWLRQRIPLQHUDOVE\ HQGROLWKV FKHPLFDOZHDWKHULQJ -RKQVWRQ DQG9HVWDO XVHG GHQWDO SLFNV WR VHSDUDWH WKH RI]RQHV RI HQGROLWKLF FRPPXQLWLHV GRPLQDWHG E\ F\DQREDFWHULD IURP %DWWOHVKLS 3URPRQWRU\ %3 DQG OLFKHQV DW /LQQDHXV 7HUUDFH /7 IRU LQRUJDQLF DQDO\VLV 7KH RXWHU FUXVW DQG LQQHU EHGURFN ]RQHV DW %3 FRQWDLQHG FKDUDFWHULVWLFDOO\ PXFK OHVV LURQ WKDQ DW /7 F DQGJ J URFNUHVSHFWLYHO\ 7KH%3PLFURELDO ]RQH FRQWDLQHG PXFK PRUH FDOFLXP F J J DQG PDJQHVLXP F J J WKDQ WKDW RI/7 F DQG J J UHVSHFWLYHO\ %RWK FRQWDLQHG VLPLODUDPRXQWVRIWRWDOSKRVSKRUXVDQGSRWDVVLXPF 7KH %3 DQG J J UHVSHFWLYHO\ F\DQREDFWHULDO ]RQH ZDV PXFK OHVV DFLG S+ WKDQWKDWRI/7S+ ZKLFKKDGDPXFKORZHU)H FRQWHQW F J J WKDQ DQ DFFXPXODWLRQ ]RQH EHQHDWK F JJ 7KH\ VXJJHVWHGWKDWWKH)H KDGEHHQOHDFKHGLQWRWKLV]RQHE\WKHFKHODWLYHHIIHFW RIR[DODWHSURGXFHGE\WKHHQGROLWKLFFRPPXQLW\ 7R LQYHVWLJDWHWKLVIXUWKHU WKH\ DQDO\VHGWKHGLVWULEXWLRQ RI R[DODWH H[WUDFWHG IURP GLIIHUHQW ]RQHV RI WKH HQGROLWKLF FRPPXQLW\ DW /7 -RKQVWRQ DQG 9HVWDO 7KH\ FRQFOXGHG WKDW R[DODWH ZDV GLVVROYLQJ WKH VDQGVWRQH FHPHQW LQ WKH OLFKHQGRPLQDWHG ]RQH
DQG FRPSRXQGLQJ WKH ZHDWKHULQJ HIIHFW E\ FRPSOH[LQJZLWK6L)HDQG$O 5XVVHOO HW DO XVHG QRQGHVWUXFWLYH )7 5DPDQVSHFWURVFRS\(GZDUGVDQG6HDZDUG WR GHPRQVWUDWH WKH VSDWLDO GLVWULEXWLRQ RI R[DODWH LQ GLIIHUHQW K\GUDWLRQ VWDWHV ZLWKLQ WKH ]RQHV RI (DVW %HDFRQHQGROLWKV in situ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³IRVVLO´ FRPPXQLWLHV )ULHGPDQQ DQG :HHG ZKLFK DUHUHYHDOHG E\ WKH OHDFKLQJ SDWWHUQV RI WKHLU DFLGV ZKLFK PRELOLVH LURQ DQG RWKHU PLQHUDOV -RKQVWRQDQG9HVWDO 7R HVWDEOLVK WKH OLPLWV RI VXUYLYDO WKUHVKROG RI H[WLQFWLRQ IRU FU\SWRHQGROLWKLF FRPPXQLWLHV )ULHGPDQQ HW DO HVWDEOLVKHG D WUDQVHFW RI PRQLWRUHG VLWHV FRQWDLQLQJ %HDFRQ VDQGVWRQH SRWHQWLDOO\ FDSDEOH RI VXSSRUWLQJ OLIH LQ WKH 0F0XUGR 'U\ 9DOOH\V GHVHUW UHJLRQ 7KH\IRXQGDVHTXHQWLDOGHFUHDVH LQPHDQDLU DQG URFN WHPSHUDWXUHV DW %DWWOHVKLS 3URPRQWRU\
'':\QQ:LOOLDPV
Table 4. 3K\VLFDODQGVWUXFWXUDOGHVFULSWLRQV RI W\SLFDO HQGROLWKLFKDELWDWVLQ VRXWKHUQ9LFWRULD/DQG
'DWDDUHIURP-RKQVWRQ 9HVWDO DQG 1LHQRZ )ULHGPDQQ
%DVHGRQLPDJHDQDO\VLVRIJHQWO\FUXVKHGURFNDXWKRUXQSXEOLVKHG
.DSSHQHWDO
'DWDIURP*UHHQILHOG
:KHUHXVHG SUHVHQW DEXQGDQW
'DWDIURP-RKQVWRQDQG9HVWDO 'DWDIRUIURP)ULHGPDQQHWDO QD GDWDQRWDYDLODEOHRUQRWDSSOLFDEOH
&KDSWHU &\DQREDFWHULDLQ 'HVHUWV /LQQDHXV 7HUUDFH VHH 7DEOH 7\URO 9DOOH\ 6 ( 0RXQW )OHPLQJ 6 ( DQG +RUVHVKRH 0RXQWDLQ 6 ( ZKLFK FRUUHODWHG ZLWK D JUDGXDO H[WLQFWLRQ RI HQGROLWKLF OLIH $OO FRPPXQLWLHV ZHUH DOLYH DW %DWWOHVKLS 3URPRQWRU\ DQG DOO ZHUH IRVVLOLVHG DW +RUVHVKRH 0RXQWDLQ 7KH WKUHVKROG RI VXUYLYDO H[WLQFWLRQDSSHDUHGWREHDW0RXQW)OHPLQJZKHUHWKH DQQXDO PHDQ URFN WHPSHUDWXUH ZDV & PD[ & PLQ & DW D PHDQ DQQXDO 5+ RI PD[ PLQ 'DWD IRU WKH F\DQREDFWHULD GRPLQDWHG FRPPXQLW\ DW %3 ZHUH LQFRPSOHWH IRU WHFKQLFDO UHDVRQV +RZHYHU FRPSDULVRQV RI PLG VXPPHU GDWD -DQXDU\ VKRZHG %DWWOHVKLS 3URPRQWRU\ PHDQ URFN WHPSHUDWXUH WR EH & PD[&PLQ & DWPHDQ5+PD[ PLQ &RQFXUUHQW FRQGLWLRQV DW 0RXQW )OHPLQJZHUHPHDQWHPS &PD[&PLQ & DWPHDQ5+PD[PLQ DQG DW +RUVHVKRH 0RXQWDLQ ZHUH PHDQ WHPS & PD[ & PLQ & DW PHDQ 5+ 55% PD[ 85%, PLQ $OWKRXJK OLIH SURFHVVHV WDNHSODFH
DW WHPSHUDWXUHV DERYH & )ULHGPDQQ HW DO WKH QXPEHU RI KRXUV SHU \HDU WKDW WKH URFN WHPSHUDWXUHZDV DERYH& ZDVHVSHFLDOO\ LQIOXHQWLDO LQGHILQLQJWKHYLDELOLW\WKUHVKROGRQWKHWUDQVHFW %
([RELRORJ\
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
Fig. 2. 'LDJUDPRIFROGGHVHUWELRPHVDQGPRUHEHQLJQKDELWDWVUHVXOWLQJIURPWKHLQWHUDFWLRQEHWZHHQPHDQDQQXDO WHPSHUDWXUHDQGPHDQ DQQXDOSUHFLSLWDWLRQLQ$QWDUFWLFDDQGSURMHFWHGIRU0DUV >IURP:\QQ:LOOLDPV DIWHU&DPHURQHWDO ZLWKSHUPLVVLRQ@
'' :\QQ:LOOLDPV
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hroococcidiopsis ZRXOG PDNH D VXLWDEOH SLRQHHU PLFURRUJDQLVP IRU WHUUDIRUPLQJ 0DUV
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³SULPRUGLDO VRXS´ DUH QRZ EHLQJ VWXGLHG IRU WKHLU VXUYLYDO SRWHQWLDO LQ WKH XOWLPDWH GHVHUW 0DUV
$FNQRZOHGJHPHQWV , WKDQN WKH %ULWLVK $QWDUFWLF 6XUYH\ %$6 WKH 8QLWHG 6WDWHV $QWDUFWLF 3URJUDPPH HVSHFLDOO\ 3URI 'LDQD )UHFNPDQ 86 1DWLRQDO 6FLHQFH )RXQGDWLRQ HVSHFLDOO\ 'U 3ROO\ 3HQKDOH DQG WKH ,WDOLDQ $QWDUFWLF 3URJUDPPH (1($ HVSHFLDOO\ ,QJ 0DULR =XFFHOOL IRU RSSRUWXQLWLHV WR ZRUN LQ GHVHUWV RI $QWDUFWLFD , HVSHFLDOO\ WKDQN 'U 'DYLG :DOWRQ %$6 IRU KLV HQFRXUDJHPHQW DQG VXSSRUW LQ WKLV OLQH RI UHVHDUFK , DP JUDWHIXO WR 'U 3HGUR 0RQWLHO DQG WKH HGLWRUV IRU YDOXDEOH FRPPHQWV RQ WKH SDSHU
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hroococcidiopsis DQG LWV DSSOLFDWLRQ WR LGHQWLILFDWLRQ RI ftsZ. $SSO (QYLURQ 0LFURELRO
&KDSWHU &\DQREDFWHULDLQ'HVHUWV %RQDQL *)ULHGPDQQ (, 2FDPSR)ULHGPDQQ 5 0F.D\ &3 DQG :RHOIOL : 3UHOLPLQDU\ UHSRUW RI UDGLRFDUERQ GDWLQJ RI FU\SWRHQGROLWKLF PLFURRUJDQLVPV 3RODUIRUVFK %RUFKHLP .< %UDWEDN* DQG +HOGDO 0 (QXPHUDWLRQ DQG ELRPDVV HVWLPDWLRQ RI SODQNWRQLF EDFWHULD DQG YLUXVHV E\ WUDQVPLVVLRQ HOHFWURQ PLFURVFRS\ $SSO (QYLURQ 0LFURELRO %UDFN $ )RUWHUUH 33LOOLQJHU&7 +RUQHFN * 6FKLGORZVNL0DQG :lQNH + (6$([RELRORJ\ 7HDP6WXG\RQWKH 6HDUFKIRU /LIH LQ WKH 6RODU 6\VWHP )LQDO 5HSRUW ,QWHUQDO 5HSRUW (XURSHDQ 6SDFH&HQWUH6DVVHQKHLP SS %UDXQH :DQG6DQNH+ ,QWHUIHURPHWULVFKH 8QWHUVXFKXQJHQ ]XU '\QDPLN YRQ +\GUDWXU XQG 7URFNHQPDVVHJHKDOW ZDKUHQG GHU /LFKWDQKlQJLJHQ .HLPXQJ GHU $NLQHWHQ YRQ Anabaena variabilis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ostoc commune. -%DFWHULRO (QJHOVNM¡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
FRYHULQJ VDQG GXQHV LQ WKHZHVWHUQ1HJHY'HVHUW)XQFWLRQDO(FRORJ\ /DQJH 2/ 0H\HU $ DQG %LLGHO % D 1HW SKRWRV\QWKHVLV DFWLYDWLRQ RID GHVLFFDWHG F\DQREDFWHULXP ZLWKRXW OLTXLG ZDWHU LQ KLJK DLU KXPLGLW\ DORQH ([SHULPHQWV ZLWK Microcoleus sociatus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phanothece halophytica. $UFK0LNURE 0LRWNH)' 'LH'QHQLP9LFWRULD9DOOH\9LFWRULD/DQG $QWDUNWLV HLQ %HLWUDJ ]XU DROLVFKHQ )RUPXQJ LP H[WUHP NDOWHQ .OLPD3RODUIRUVFK 0RQWLHO 32 DQG &RZDQ '$ 7KH SRVVLEOH UROH RI VROXEOH FDUERK\GUDWHVDQGSRO\ROVDVFU\RSURWHFWDQWVLQ$QWDUFWLFSODQWV ,Q +H\ZRRG5% HG 8QLYHUVLW\5HVHDUFKLQ$QWDUFWLFD 3URFHHGLQJV RI WKH %ULWLVK $QWDUFWLF 6XUYH\ $QWDUFWLF 6SHFLDO 7RSLF $ZDUG 6FKHPH 5RXQG 6\PSRVLXP 6HSWHPEHU 2FWREHU SS %ULWLVK $QWDUFWLF 6XUYH\&DPEULGJH 1LHQRZ-$DQG)ULHGPDQQ(, 7HUUHVWULDOOLWKRSK\WLFURFN FRPPXQLWLHV ,Q)ULHGPDQQ(,HG $QWDUFWLF0LFURELRORJ\ SS :LOH\/LVV1HZ
1LHQRZ -$ 0F.D\ &3 DQG )ULHGPDQQ (, E &U\SWRHQGROLWKLF PLFURELDO HQYLURQPHQW LQ WKH 5RVV 'HVHUW RI $QWDUFWLFD OLJKW LQ SKRWRV\QWKHWLFDOO\ DFWLYH UHJLRQ 0LFURE (FRO 2FDPSR)ULHGPDQQ 5 0H\HU 0$ &KHQ 0 DQG )ULHGPDQQ (, 7HPSHUDWXUH UHVSRQVH RI $QWDUFWLF FU\SWRHQGROLWKLF SKRWRV\QWKHWLFPLFURRUJDQLVPV3RODUIRUVFK 2OLH-- DQG 3RWWV 0 3XULILFDWLRQ DQG ELRFKHPLFDO DQDO\VLV RI WKH F\WRSODVPLF PHPEUDQH IURP WKH GHVLFFDWLRQWROHUDQW F\DQREDFWHULXP Nostoc commune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ostoc commune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ostoc commune. 3ODQW 3K\VLRO 6FKHUHU 6 DQG =KRQJ =3 'HVLFFDWLRQ LQGHSHQGHQFH RI WHUUHVWULDO Nostoc commune HFRW\SHV &\DQREDFWHULD 0LFURE (FRO 6FKRSI-: 0LFURIRVVLOV RIWKH HDUO\ $UFKHDQ $SH[ FKHUW QHZHYLGHQFHRIWKHDQWLTXLW\RIOLIH 6FLHQFH 6FKRSI-: DQG :DOWHU 05 2ULJLQ DQG HDUO\ HYROXWLRQ RI F\DQREDFWHULD WKH JHRORJLFDO HYLGHQFH ,Q &DUU 1* DQG
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
Summary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
B A Whitton and M Potts (eds). The Ecology of Cyanobacteria, Kluwer Academic Publishers Printed LQ the Netherlands
1+0DQQ
E\ SKRVSKRU\ODWLRQ LV UHFRJQLVHG 7KH SRWHQWLDO WR XQGHUVWDQG F\DQREDFWHULDO VLJQDOOLQJ PHFKDQLVPV KDV EHHQ VLJQLILFDQWO\HQKDQFHGE\JHQRPLFVHTXHQFHLQIRUPDWLRQDQGZKHUHSRVVLEOHWKLVKDVEHHQXWLOLVHGWRFRQILUPRU H[WHQGWKHSUHVHQWHPSLULFDONQRZOHGJH
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bbreviations: &$03 F\FOLF ¶¶DGHQRVLQH PRQRSKRVSKDWH &L LQRUJDQLF FDUERQ *3'+ JOXFRVHSKRVSKDWH GHK\GURJHQDVH +3. KLVWLGLQH SURWHLQ NLQDVH 3L LQRUJDQLF SKRVSKDWH SS*SS JXDQRVLQH¶GLSKRVSKDWH¶GLSKRVSKDWH 34 SODVWRTXLQRQH 34+SODVWRTXLQRO36,SKRWRV\VWHP,36,,SKRWRV\VWHP,,55 UHVSRQVHUHJXODWRU
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lac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ynechocystis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ynechocystis 3&& JHQRPH.DQHNR HW DO WKRXJK TXRUXP VHQVLQJ W\SH PROHFXOHV 1DF\ODWHGKRPRVHULQH ODFWRQHV KDYHEHHQ GHWHFWHG LQ F\DQREDFWHULDO EORRPV %DFKRIHQ DQG 6FKHQN
,, 3URWHLQ 3KRVSKRU\ODWLRQ
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
histidine protein kinase
1+0DQQ
response regulator
Figure 1. $GLDJUDPLOOXVWUDWLQJ WKHGRPDLQRUJDQL]DWLRQRIWKHW\SLFDO KLVWLGLQHSURWHLQNLQDVHV DQGUHVSRQVHUHJXODWRUV RIWZRFRPSRQHQW VHQVRU\V\VWHPVDQGSDWWHUQRISKRVSKRWUDQVIHU 7KH+1')DQG*ER[HVRIWKHKLVWLGLQHSURWHLQNLQDVHVDUHLQGLFDWHGWRJHWKHUZLWKWKH FRQVHUYHG DPLQR DFLG VHTXHQFH IHDWXUHV ' DVSDUWDWH )SKHQ\ODODQLQH * JO\FLQH + KLVWLGLQH . O\VLQH [ DQ\ DPLQR DFLG IRU ERWK WKH KLVWLGLQHSURWHLQNLQDVHV DQGWKHUHVSRQVHUHJXODWRUV 7KHQXPEHULQJRIWKHFRQVHUYHGUHVLGXHV LQWKHUHVSRQVHUHJXODWRULV WKDWDSSO\LQJWR WKH Escherichia coli &KH<SURWHLQ
UHFHLYHUGRPDLQRI55VLVXVXDOO\DSSUR[LPDWHO\ DPLQR DFLGV LQ VL]H $Q\ WZR UHFHLYHU GRPDLQV XVXDOO\H[KLELW LGHQWLW\DQGWKHUHVLGXHVWKDW FRUUHVSRQG WR $VS $VS DQG /\V LQ WKH ZHOO VWXGLHGDQG DUFKHW\SDOE. coli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ynechocystis 3&& .DQHNR HWDO LQGLFDWHV DW OHDVW +3.V DQG
55V &\DQREDFWHULDO SURWHLQV FRQWDLQLQJ HLWKHU +3. RU 55 PRGXOHVKDYH EHHQ LPSOLFDWHG LQ DZLGH UDQJH RI DGDSWLYHUHVSRQVHV 7DEOH 7KH 55 1E5 IURP Synechococcus 3&& LV LQYROYHG LQ WKH UHJXODWLRQ RI SK\FRELOLVRPH GHJUDGDWLRQ LQ UHVSRQVH WR QXWULHQW OLPLWDWLRQ EXW DOVR PRGXODWHV DGGLWLRQDO IXQFWLRQV FULWLFDO IRU FHOO VXUYLYDO GXULQJ QXWULHQW OLPLWHG DQG KLJK OLJKW FRQGLWLRQV 6FKZDU] DQG *URVVPDQ $ JHQH patA LQYROYHG LQ WKH FRQWURO RI KHWHURF\VW SDWWHUQ IRUPDWLRQHQFRGHVD DPLQR DFLG SURWHLQ FRQVLVWLQJ RI GRPDLQV RI ZKLFK WKH &WHUPLQDO GRPDLQ H[KLELWHG WKH VHTXHQFH IHDWXUHV FKDUDFWHULVWLF RI D 55 /LDQJ HW DO 7UDQVSRVRQ PXWDJHQHVLV RI D V\PELRWLF QLWURJHQ IL[LQJ Nostoc VWUDLQOHGWRWKHLGHQWLILFDWLRQ RI D JHQH devR, HQFRGLQJ DSURWHLQ ZLWK WKH FRQVHUYHGIHDWXUHV RI D 55 VLPLODU WR WKH E. coli &KH< DQG Bacillus subtilis 6SR2) ZKLFK ZDV UHTXLUHG IRU QLWURJHQ IL[DWLRQ XQGHU DHURELF FRQGLWLRQV EXW GLG QRW EORFN KHWHURF\VW GHYHORSPHQW &DPSEHOO HW DO 6LPLODUO\ WKH rcaC JHQH ZKLFK FRPSOHPHQWV D PXWDWLRQ FDXVLQJ UHGOLJKW LQGLIIHUHQFH LQ WKH
&KDSWHU 'HWHFWLQJWKH(QYLURQPHQW
7DEOH Cyanobacterial adaptive responses for which there is evidence implicating proteins with features characteristic of the modules of two component sensory system histidine protein kinases or response regulators.
3K\VLRORJLFDOIXQFWLRQ
*HQHVRU25)V
2UJDQLVP
5HIHUHQFHV
3K\FRELOLVRPHGHUDGDWLRQDQG VXUYLYDO XQGHU QXWULHQWOLPLWHGKLJK OLJKWFRQGLWLRQV
nblR
Synechococcus 3&&
6FKZDU] DQG *URVVPDQ
&RPSOHPHQWDU\ FKURPDWLFDGDSWDWLRQ
rcaC rcaE rcaF
Fremyella diplosiphon Frernyella diplosiphon Fremyella diplosiphon
&KLDQJ HW D .HKRH DQG *URVVPDQ .HKRH DQG *URVVPDQ
+HWHURF\VWSDWWHUQIRUPDWLRQ
patA
Anabaena 3&&
/LDQJ HW DO
1LWURJHQ IL[DWLRQ
devR
Nostoc $7&&
&DPSEHOOHWDO
&2 DYDLODELOLW\
hatR
Synechocystis 3&&
%pGX HW DO
Synechocystis 3&&
.DQHNRHWDO
phy plpA
Synechocystis 3&& Synechocystis 3&&
+XJKHV HW DO :LOGH HW DO
+HUELFLGH UHVLVWDQFHFKHPLFDO VLJQDOOLQJ
dspA
Synechocystis 3&&
%DUWVHYLFKDQG6KHVWDNRY
3KRVSKDWHDYDLODELOLW\
sphR/sphS phoB/phoR
Synechococcus 3&& Synechococcus :+
$LED HW DO :DWVRQHWDO
3K\WRFKURPH DQG HWK\OHQHUHVSRQVH VOU VOO VOUVOU OLNHVLJQDOOLQJ
FRPSOHPHQWDU\ FKURPDWLF DGDSWLQJ Fremyella diplosiphon, HQFRGHVDDPLQRDFLGSURWHLQWKH1 WHUPLQXVRIZKLFKFRQWDLQVWZR55UHFHLYHUGRPDLQV RQHDWWKH1WHUPLQXVDQGWKHRWKHUDWWKH&WHUPLQXV &KLDQJ HW DO 0RUH UHFHQWO\ .HKRH DQG *URVVPDQ KDYHVKRZQWKDWWKHrcaE DQG rcaF JHQHVRIF. diplosiphon, ZKLFKPD\EHLQYROYHG LQ WKH VHQVLQJ RIOLJKW TXDOLW\ HQFRGH SURWHLQV ZLWK WKH IHDWXUHV RI DQ +3. DQG 55 UHVSHFWLYHO\ 0XWDWLRQV LQ Synechocystis 3&& LQ D JHQH GHVLJQDWHG dspA, ZKLFKOHDGWRFURVVUHVLVWDQFHWRWKH KHUELFLGHV GLIXQRQ DQG GLXURQ DV ZHOO DV WKH FDOPRGXOLQ DQWDJRQLVWV FKORUSURPD]LQH DQG WULIOXRSHUD]LQH GHILQH DJHQHHQFRGLQJ DQ +3.ZLWK WZRW\SLFDOWUDQVPHPEUDQHVHJPHQWV%DUWVHYLFKDQG 6KHVWDNRY $LED HW DO FKDUDFWHUL]HG WZR FRQWLJXRXV JHQHV sphS DQG sphR IURP WKH IUHVKZDWHU VWUDLQ Synechococcus 3&& ZKLFK HQFRGH DQ +3. DQG 55 UHVSHFWLYHO\ 0XWDWLRQV LQ WKHVH JHQHV SUHYHQW WKH LQGXFWLRQ RI DONDOLQH SKRVSKDWDVH DQG RWKHU SRO\SHSWLGHV DVVRFLDWHG ZLWK 3L VWDUYDWLRQ ,PSRUWDQWO\ 1DJD\DHWDO KDYH VKRZQ WKDW 6SK5 LV SKRVSKRU\ODWHG E\
SKRVSKRWUDQVIHUIURP 6SK6 DQG ELQGV WR WZR GLVWLQFW VLWHV XSVWUHDP IURP WKH DONDOLQH SKRVSKDWDVH (phoA) SURPRWHU ,Q WKH FDVH RI WKH RFHDQLF VWUDLQ Synechococcus :+ WZR FRQWLJXRXV JHQHV GHVLJQDWHG phoB DQG phoR HQFRGH D 55 DQG +3. UHVSHFWLYHO\ DQG WUDQVFULSWLRQ RI WKHVH WZR JHQHV LV LQGXFHG E\ 3L VWDUYDWLRQ :DWVRQ HW DO 6HYHUDO RWKHU +3. DQG 55 JHQHV RI XQFOHDU SK\VLRORJLFDO VLJQLILFDQFH KDYH EHHQ LGHQWLILHG LQ F\DQREDFWHULD0DQQ VXFK DV WKRVH FORQHG E\ 1DJD\DHWDO IURPSynechococcus 3&& RQ WKH EDVLV RI WKHLU DELOLW\ WR FRPSOHPHQW +3. PXWDQWV RI E. coli, DQG DQ LQFRPSOHWH 25) IRXQG GRZQVWUHDP IURP WKH pet H 1$'3IHUUHGR[LQ R[LGRUHGXFWDVH JHQH RI Synechococcus 3&& 6FKOXFKWHU DQG %U\DQW $Q DGMDFHQW SDLU RI +3. DQG 55 JHQHV KDYH EHHQ FKDUDFWHUL]HG IURP Synechococcus 3&& EXW LQVHUWLRQDO PXWDJHQHVLV RI WKH 55 JHQH \LHOGHG QR REYLRXV SKHQRW\SH 6FDQODQ DQG 0DQQ DQG VLPLODU REVHUYDWLRQV ZHUH PDGH IROORZLQJ WKH GLVUXSWLRQ RI WZR SXWDWLYH 55 JHQHV GHVLJQDWHG srrA DQG srrB,
IURP Synechococcus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ynechocystis 3&& .DQHNR HW DO KDVERWKWUDQVPLWWHUDQG UHFHLYHU GRPDLQV DV GR VHYHUDO RWKHU 25)V ,Q WKH FDVH RI VRPH RI WKHVH PRUH FRPSOH[ VLJQDOOLQJ SURWHLQV LW LV WKRXJKW WKDWWKH\PD\SOD\ DSDUWLQ DIRXUVWHSSKRVSKRUHOD\ LQ ZKLFK WKHUH LV D +LV$VS+LV$VS VHTXHQWLDO SKRVSKRWUDQVIHU $SSOHE\ HW DO 6XFK D V\VWHP LV W\SLILHG E\ WKH SDWKZD\ JRYHUQLQJ WKH LQLWLDWLRQRIVSRUXODWLRQLQ B. subtilis DQG DOVRRFFXUV LQWKH%YJ6%YJ$WZRFRPSRQHQWV\VWHPLQYROYHGLQ WKH UHJXODWLRQ RI YLUXOHQFH IDFWRUV LQ Bordetella pertussis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ynechocystis 3&& LQ UHODWLRQ WR RWKHUJHQHVRINQRZQSK\VLRORJLFDOIXQFWLRQPD\JLYH VRPHFOXHVDVWRWKHLUOLNHO\SK\VLRORJLFDOUROH
1+0DQQ % 3K\WRFKURPH DQGHWK\OHQH UHVSRQVH UHODWHG SURWHLQV *LYHQ WKH LPSRUWDQFH RI SK\WRFKURPHV LQ HQDEOLQJ SODQWV WR UHVSRQG WR SDUWLFXODU FKDQJHV LQ WKHLU OLJKW HQYLURQPHQW WKH TXHVWLRQ RIZKHWKHU VXFK PROHFXOHV SOD\ D VLPLODU UROH LQ F\DQREDFWHULD KDV EHHQ UDLVHG IUHTXHQWO\ 7KH SK\WRFKURPHV FRQVWLWXWH D IDPLO\ RI SODQW VHQVRUV WKDW FDQ H[LVW LQ WZR SKRWR LQWHUFRQYHUWLEOH LVRPHULF IRUPV DQG HQDEOH SODQWV WR UHVSRQG WR SDUWLFXODU FKDQJHV LQ WKHLU OLJKW HQYLURQPHQW E\ D ZLGH YDULHW\ RI PRUSKRJHQHWLF UHVSRQVHV 7KH FKURPRSKRUH LV D OLQHDU WHWUDS\UUROH WKDW EHFRPHV DWWDFKHG WR WKH DSRSURWHLQ YLD D WKLRHWKHU ERQG IRUPHG E\ DQ DXWRFDWDO\WLF VHOI DVVHPEO\ SURFHVV /DJDULDV DQG /DJDULDV ,W ZLOOEHXVHIXOWRORRNEULHIO\DWWKHVWDWHRINQRZOHGJH LQ Arabidopsis thaliana. )LYH GLVWLQFW SK\WRFKURPH JHQHVKDYHEHHQGHWHFWHG6KDUURFNDQG4XDLO 3K\WRFKURPH $ SUREDEO\ PHGLDWHV WKH IDUUHG KLJK LUUDGLDQFH UHVSRQVH ZKHUHDV SK\WRFKURPH % LV UHVSRQVLEOH IRUWKHUHGIDUUHGUHVSRQVH DQG WKH RWKHU SK\WRFKURPHV SOD\ PLQRU UROHV 6PLWK 'HOHWLRQ DQDO\VLV KDV UHYHDOHG WKDW WKHUH DUH VHSDUDWH GRPDLQV ZLWKLQ WKH a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
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW 7KH HWK\OHQH KRUPRQHUHVSRQVH SDWKZD\ LQ SODQWV DIIHFWV PDQ\ DVSHFWV RI SODQW JURZWK DQG GHYHORSPHQW 6WXGLHV ZLWK Arubidopsis PXWDQWV GLVSOD\LQJ GRPLQDQW HWK\OHQH LQVHQVLWLYLW\ IDFLOLWDWHG WKH FKDUDFWHUL]DWLRQ RI WKH etrl JHQH &KDQJ HW DO $OWKRXJK WKH VHTXHQFH RIWKH DPLQRWHUPLQDO KDOI RI WKH GHGXFHG (75O SURWHLQ DSSHDUV WR EH QRYHO WKH FDUER[\OWHUPLQDO KDOI FRQWDLQV ERWK WKH WUDQVPLWWHU DQG UHFHLYHU GRPDLQV RI +3.V DQG 55V UHVSHFWLYHO\ &KDQJ HW DO &KDQJ DQG 0H\HURZLW] $ VHFRQG JHQH UHODWHG WR etrl, KDV EHHQFKDUDFWHUL]HG DQG WKH GHGXFHG(56 SURWHLQ ZKLFK H[KLELWV LGHQWLW\ ZLWK (75 KDV D WUDQVPLWWHU GRPDLQ EXW QR UHFHLYHU GRPDLQ +XD HW DO ,W LV VXJJHVWHG WKDW ERWK SURWHLQV DUH HWK\OHQH VHQVRUV DQG WKXV WKHUH LV UHGXQGDQF\ LQ WKH VLJQDOOLQJ SDWKZD\ 7KH 25) VOO LQ WKH JHQRPH RI Synechocystis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rcaE JHQH RIFremyella diplosiphon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ynechocystis 3&& .DQHNR HW DO DOWKRXJK WKLV RUJDQLVP GRHV QRW XQGHUJR FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ 7KH IRXU PRVW VLPLODU25)V VOO VOU VOO DQG VOU DOOH[KLELWWKHFRQVHUYHG7/DQG5/PRWLIV DQGDOO EXW RQH VOO DOVR KDYH &WHUPLQDO +3.
WUDQVPLWWHU GRPDLQV 25)V VOO DQG VOO H[KLELW VRPH VLPLODULW\ WR WKH SK\WRFKURPH FKURPRSKRUH DWWDFKPHQW UHJLRQ 7KH JHQH plpA VOO ZDV GLVUXSWHG LQSynechocystis 3&& DQG WKH PXWDQW ZDV XQDEOH WR JURZ LQ EOXH OLJKW :LOGHHWDO ,Q FRQWUDVW WR WKH RWKHU 25)V HQFRGHG LQ WKH JHQRPH RI Synechocystis 3&& ZKLFK H[KLELW OLPLWHGUHJLRQRIVLPLODULW\WRWKHSODQWSK\WRFKURPHV 25) VOU GLVSOD\V PDUNHG H[WHQGHG VLPLODULWLHV .DQHNR HW DO 7KH VHTXHQFH VLPLODULWLHV DUH QRW FRQILQHG WR WKH UHJLRQ DOLJQLQJ ZLWK +3. EXW H[WHQG LQWR WKH UHJLRQV LQYROYHG LQ FKURPRSKRUH DWWDFKPHQWDQGVSHFWUDOLQWHJULW\,QGHHGWKHF\VWHLQH LQYROYHG LQ FKURPRSKRUH DWWDFKPHQWLV SUHVHQWLQ WKH F\DQREDFWHULDO KRPRORJXH 7KLV SXWDWLYH F\DQR EDFWHULDO SK\WRFKURPH JHQH (phy) LV VLWXDWHG LPPHGLDWHO\XSVWUHDPIURP DQ 25)WKDWGLVSOD\V WKH UHFHLYHUGRPDLQ RID55 ([SUHVVLRQ RIWKHphy JHQH LQ E. FROL UHYHDOHG WKDW WKH SURWHLQ ZDV DEOH WR DXWRFDWDO\WLFDOO\ UHDFW ZLWK SK\FRF\DQRELOLQ WR SURGXFH D SKRWRFKURPLF KRORSURWHLQ ZLWK DEVRUSWLRQ PD[LPD DW DQG QP DIWHU UHG DQG IDUUHG LUUDGLDWLRQ +XJKHV HW DO /DPSDUWHU HW DO
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
H[FOXGLQJ WKH FDVH RI LVRFLWUDWH GHK\GURJHQDVH SKRVSKRU\ODWLRQLQE. coli, WKHUHZDVOLPLWHGSURJUHVV LQWKLVDUHDXQWLOUHFHQWO\+RZHYHUWKHUHLVH[WHQVLYH XQDPELJXRXV HYLGHQFH IRU WKH PRQRHVWHU SKRVSKRU\ODWLRQ RI SURWHLQV LQ D UDQJH RI F\DQREDFWHULDO VWUDLQV XVLQJERWK in vivo DQG in vitro ODEHOOLQJ WHFKQLTXHV DQG FKDQJHV LQ WKH SDWWHUQ RI SURWHLQ SKRVSKRU\ODWLRQ KDYH EHHQ FRUUHODWHG ZLWK D YDULHW\RISK\VLRORJLFDOFKDQJHV0DQQ (XNDU\RWLF VHULQHOWKUHRQLQH DQG W\URVLQH SURWHLQ NLQDVHV IDOO LQWR D VLQJOH VXSHUIDPLO\ +DQNV HW DO 7KH LQLWLDO GLVFRYHU\ RI VXFK ³HXNDU\RWLF´ SURWHLQ NLQDVHV LQ SURNDU\RWHV ZDV EDVHG RQ WKH GHVLJQ RI 3&5 SULPHUV DJDLQVW FRQVHUYHG UHJLRQV LQ WKH FDWDO\WLF GRPDLQ RI VHULQHWKUHRQLQH SURWHLQ NLQDVHV DQG OHG WR WKH FKDUDFWHUL]DWLRQ RI D VHULQHOWKUHRQLQH SURWHLQ NLQDVH JHQH (pknl) IURP Myxococcus xanthus 0XLLR]'RUDGR HW DO DQG HVWDEOLVKPHQW WKDW WKLV NLQDVH ZDV UHTXLUHG IRU QRUPDO GLIIHUHQWLDWLRQ 6XEVHTXHQW VWXGLHV LQGLFDWHG WKDWWKHUHZDV DIDPLO\ RIVXFKNLQDVHV DQG WKDWWKH\ RFFXUUHG LQ D UDQJH RI RUJDQLVPV =KDQJ .HQQHOO\ DQG 3RWWV 7KH ILUVW UHSRUW RI DQ ³HXNDU\RWLF´ NLQDVH LQ D F\DQREDFWHULXPHPSOR\HG D VLPLODU 3&5 DSSURDFK WR WKDW XVHG ZLWK M. xanthus =KDQJ 7KH pknA JHQH RI Anabaena 3&& HQFRGHV D SURWHLQ WKH 1WHUPLQDO UHJLRQ RI ZKLFK H[KLELWV VLJQLILFDQW VLPLODULW\ WR WKH FDWDO\WLF GRPDLQV RI VHULQHOWKUHRQLQH NLQDVHV ,QDFWLYDWLRQ RI pknA UHGXFHG WKH IUHTXHQF\ RI KHWHURF\VWV LQ WKH ILODPHQWV DQG FRORQLHV RI WKH PXWDQW DSSHDUHG OLJKW JUHHQDQGURXJKLQWKHDEVHQFHRIFRPELQHGQLWURJHQ 6RXWKHUQ K\EULGL]DWLRQ H[SHULPHQWV LQGLFDWHG WKDW WKHUH ZDV DIDPLO\ RIVXFK JHQHV LQAnabaena 3&& =KDQJ $Q25)H[KLELWLQJVLPLODULW\WR UDW P\RVLQ OLJKW FKDLQ VHULQHOWKUHRQLQH NLQDVH KDV EHHQ UHSRUWHG IRU Fremyella diplosiphon *URVVPDQ HW DO $ WRWDO RI VL[ VHULQHWKUHRQLQH SURWHLQ NLQDVH JHQHV KDYH QRZ EHHQ FKDUDFWHUL]HG IURP Anabaena 3&& DQG DOWKRXJK WKHLU DQDO\VLV LV QRWFRPSOHWHWKHUHLVHYLGHQFHWKDWVRPHRIWKHPPD\ EH UHTXLUHG IRU JURZWK LQ WKH DEVHQFH RI FRPELQHG QLWURJHQ=KDQJ =KDQJDQG/LEV =KDQJ HW DO D 3K\ORJHQHWLF DQDO\VLV EDVHG RQ WKH FDWDO\WLF GRPDLQV RI VHULQHOWKUHRQLQH SURWHLQ NLQDVHV IURPSURNDU\RWHV VKRZWKDWILYH3NQ$3NQ%3NQ& 3NQ' DQG 3NQ( RIWKH VL[ NLQDVHV IURP Anabaena 3&&FOXVWHUWRJHWKHU ZKLOHRQH +VW. LVPRUH FORVHO\ UHODWHG WR 67( IURP Saccharomyces cerevisiae =KDQJ 7KH JHQRPLF VHTXHQFH RI Synechocystis 3&& UHYHDOV VHYHQ JHQHV DSSDUHQWO\ HQFRGLQJ VHULQHOWKUHRQLQH SURWHLQ NLQDVHV
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³HXNDU\RWLF´W\SH SURWHLQ NLQDVHV KDYH EHHQ GLVFRYHUHG LQ F\DQREDFWHULD DQG RWKHUSURNDU\RWHVLWPD\ZHOOEHWKHFDVHWKDWPXFKRI SKRVSKRU\ODWLRQ RISURWHLQV DV PRQRHVWHUV LV FDUULHG E\NLQDVHVWKDWH[KLELWQRKRPRORJ\WRWKHHXNDU\RWLF NLQDVHV 7KLV LV WKH FDVH IRU WKH LVRFLWUDWH GHK\GURJHQDVH NLQDVHOSKRVSKDWDVH RIE. coli &RUWD\ HW DO WKXV D QHZ FODVV RU FODVVHV RI SURNDU\RWLF NLQDVHV PD\ DZDLW GLVFRYHU\ 6HFWLRQ ,,' ,Q HXNDU\RWHV WKH QXPEHU DQG YDULHW\ RI SURWHLQ NLQDVHV DUH SDUDOOHOHG E\ SURWHLQ SKRVSKDWDVHV UHVSRQVLEOH IRU GHSKRVSKRU\ODWLQJ VHULQHOWKUHRQLQH DQG W\URVLQH UHVLGXHV 7KLV DEXQGDQFH RI SKRVSKDWDVHV KDV QRW VR IDU EHHQ UHIOHFWHG LQ SURNDU\RWHV H[FHSW LQ WKH FDVH RI DQDO\VLV RI HXNDU\RWLFOLNH VLJQDOOLQJ SURWHLQV HQFRGHG E\ WKH Synechocystis 3&& JHQRPH =KDQJ HW DO E *HQHVHQFRGLQJHXNDU\RWLFOLNHSKRVSKDWDVHV KDYH DOVR EHHQ FKDUDFWHUL]HG LQAnabaena 3&& =KDQJ HW DO DQG Microcystis aeruginosa VS 87(; 6KL DQG &DUPLFKDHO 7KH PRVW GHWDLOHG FKDUDFWHUL]DWLRQ RI D F\DQREDFWHULDO SURWHLQ SKRVSKDWDVH LQYROYHV WKH PRQRHVWHUDVH ,SK3 IURP Nostoc commune 87(; 7KH JHQH (iph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
&KDSWHU 'HWHFWLQJWKH(QYLURQPHQW
7DEOH Cyanobacterial adaptive responses in which the monoester phosphorylation of proteins LV implicated.
(QYLURQPHQWDO 6LJQDO
$GDSWLYH UHVSRQVH
2UJDQLVP
5HIHUHQFHV
/LJKW VSHFWUDO TXDOLW\
FRPSOHPHQWDU\
Fremyella diplosiphon
&KLDQJ HW DO
FKURPDWLF DGDSWDWLRQ
Calothrix 3&& Fremyella diplosiphon
6REF]\N HW DO .HKRH DQG *URVVPDQ
/LJKW VSHFWUDO TXDOLW\ /LJKW LQWHQVLW\
VWDWH WUDQVLWLRQ
Synechococcus 3&&
"
Synechococcus 3&& Anabaena 3&&
$OOHQ :DUQHU DQG %XOOHUMDKQ 0F&DUWQH\ HW DO
$PPRQLXP LRQV&2 IL[DWLRQ
3,, SKRVSKRU\ODWLRQ
Synechococcus 3&&
"
3SK\FRF\DQLQ SKRVSKRU\ODWLRQ
Synechococcus 3&&
6DOLQLW\ &2 DYDLODELOLW\ $YDLODELOL\ RI IL[HG QLWURJHQ 1 IL[DWLRQ
"
Synechocystis 3&&
&L WUDQVSRUW +HWHURF\VW IRUPDWLRQ
Synechocystis 3&& Anabaena 3&&
+DJHPDQQ HW DO %OR\H HW DO =KDQJ
$PPRQLXP LRQV
1LWUDWH WUDQVSRUW
Synechococcus 3&&
5RGULJXH] HW DO
SURWHLQ W\URVLQH SKRVSKRU\ODWLRQ
,QWHUHVWLQJO\ ,KS3 H[KLELWHG 0$3 NLQDVH SKRVSKDWDVH DFWLYLW\ ,QWURGXFWLRQ RI WKH iphP JHQH RQD PXOWLFRS\ SODVPLG LQWR Anabaena 3&& OHDGV WR FKDQJHV LQ WKH SDWWHUQ RI SURWHLQ W\URVLQH SKRVSKRU\ODWLRQ 6 6PLWK DQG 0 3RWWV SHUV FRPP +RZHYHULWLVFOHDUIURPDYDULHW\RIin vivo DQG in vitro VWXGLHVWKDWWXUQRYHURISURWHLQPRQRHVWHU SKRVSKDWHV GRHV RFFXU LQ F\DQREDFWHULD DQG WKDW WKH DFWLYLW\ RIWKHVH SKRVSKDWDVHV PD\ EH PRGXODWHG E\ UHGR[ FRQGLWLRQV DV ZHOO DV WKH FRQFHQWUDWLRQV RI PHWDEROLWHVVXFKDVULEXORVHSKRVSKDWHDQG JOXFRVH SKRVSKDWH0DQQHWDO $OOHQ
' 2WKHU0RGHV RI3KRVSKRU\ODWLRQ 3URWHLQ SKRVSKRU\ODWLRQ LQYROYLQJ PRQRHVWHUV KDV EHHQ LPSOLFDWHG LQ D ZLGH UDQJH RI SK\VLRORJLFDO UHVSRQVHV LQ F\DQREDFWHULD 7DEOH KRZHYHU IHZ RI WKH SURWHLQV LQYROYHG KDYH EHHQ LGHQWLILHG DQG NLQDVHV RWKHU WKDQ WKRVH RI WKH HXNDU\RWLFW\SH PD\ EH LQYROYHG 3,, UHSUHVHQWV WKH VLQJOH H[DPSOH LQ F\DQREDFWHULD ZKHUH WKH PRQRHVWHU SKRVSKRU\ODWLRQ RID VSHFLILF LGHQWLILHG SURWHLQ DQG WKH SK\VLRORJLFDO VLJQLILFDQFH RI WKH SURFHVV KDYH EHHQ ZHOO FKDFWHUL]HG,QE. coli WKH3,, *OQ% SURWHLQLVVXEMHFW
+DUULVRQ HW DO 7VLQRUHPDV HW DO )RUFKKDPPHU DQG 7DQGHDX GH 0DUVDF DE +DUULVRQ
WR XULG\O\ODWLRQ DV D UHVSRQVH WR D GHFUHDVLQJ JOXWDPLQH FRQFHQWUDWLRQ UHODWLYH WR R[RJOXWDUDWH DQG WKH PRGLILHG IURP RI 3,, DFWLYDWHV DQ DGHQ\OWUDQVIHUDVH ZKLFK GHDGHQ\ODWHV JOXWDPLQH V\QWKHWDVH FDXVLQJLWV DFWLYDWLRQ 7KH XULG\O\ODWLRQ RI 3,, DOVR FDXVHV DQ DFWLYDWLRQ RI WKH GHSHQGHQW WUDQVFULSWLRQ RI WKH glnALG RSHURQ DQG LQFUHDVHG V\QWKHVLV RI JOXWDPLQH V\QWKHWDVH YLD D SKRVSKRU\ODWLRQ FDVFDGH 1LQID HW DO $ 3,, *OQ% KRPRORJXH ZDV LGHQWLILHG LQ Synechococcus 3&& DQGZDVIRXQG WREH VXEMHFWWRVRPHIRUP RI SRVWWUDQVODWLRQDO PRGLILFDWLRQ ZKLFK ZDV PRGXODWHG E\ WKH DPPRQLXP LRQ FRQFHQWUDWLRQ DQG SRVVLEO\ E\ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW 7VLQRUHPDVHWDO )RUFKKDPPHUDQG7DQGHDX GH0DUVDF VKRZHGWKDW LQIDFW WKH 3,, SURWHLQ RI Synechococcus 3&& ZDV VXEMHFW WR SKRVSKRU\ODWLRQ RI D VHULQH UHVLGXH UDWKHU WKDQ XULG\O\ODWLRQ DQG LWZDV IXUWKHUGHPRQVWUDWHG WKDWWKH SKRVSKRU\ODWLRQ RI 3,, ZDV SULPDULO\ GHWHUPLQHG E\ WKH 1VWDWXV RI WKH FHOO EXW DOVR E\ &2 IL[DWLRQ )RUFKKDPPHU DQG 7DQGHDX GH0DUVDF E 7KH SKRVSKRU\ODWLRQ VLWH RI 3,, ZDV FKDUDFWHUL]HG DQG IRXQG WR KDYH DQ 5;6 PRWLI DOVR IRXQG IRU HXNDU\RWLF F$03GHSHQGHQW SURWHLQ NLQDVHV DQG WKH
3,, NLQDVH DFWLYLW\ ZDV GHWHUPLQHG E\ WKH FRQFHQWUDWLRQRIR[RJOXWDUDWHDQG QRWE\ JOXWDPLQH )RUFKKDPPHU DQG 7DQGHDX GH 0DUVDF D ,Q IDFW 3,, ELQGV R[RJOXWDUDWH DQG $73 LQ D PXWXDOO\ GHSHQGHQW PDQQHU DQG LW KDV EHHQ SURSRVHG WKDW WKH SULPDU\IXQFWLRQ RIWKH Synechococcus 3,, LV WR VHQVH R[RJOXWDUDWH WKH FDUERQ VNHOHWRQ UHTXLUHG IRU QLWURJHQ DVVLPLODWLRQ )RUFKKDPPHU DQG +HGOHU +RZHYHUWKHUHDOVRDSSHDUVWREHDUROHIRU3,, LQ WKH FRQWURO RI QLWUDWHQLWULWH WUDQVSRUW /HH HW DO 7KH 3,, NLQDVH DQG SKRVSKDWDVH DFWLYLWLHV DUH ELRFKHPLFDOO\VHSDUDEOHDQGWKHODWWHUZDV FDSDEOH RI GHSKRVSKRU\ODWLQJ SKRVSKRVHULQHWKUHRQLQH UHVLGXHV RQ VHYHUDO SURWHLQV WKRXJK RQO\ 3,, GH SKRVSKRU\ODWLRQ ZDV UHJXODWHG E\ R[RJOXWDUDWH DQG $73,UPOHUHWDO
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
1+0DQQ WKH ODWWHU DV WKH PRVW LPSRUWDQW URXWH RI R[LGDWLYH JOXFRVHGLVVLPLODWLRQ &DUU 6PLWK 7KH 233 LV WKRXJKW DOVR WR EH ODUJHO\ UHVSRQVLEOH IRU WKH VXSSO\ RI UHGXFWDQW WR QLWURJHQDVH LQ WKH KHWHURF\VW $SWHHWDO 6XPPHUVHWDO E ,QKLJKHU SODQWV LW LV ZHOO HVWDEOLVKHG WKDW VHYHUDO HQ]\PHV RI SKRWRV\QWKHWLF DQG UHVSLUDWRU\ PHWDEROLVP DUH DFWLYDWHGE\DSURFHVVLQYROYLQJWKLRUHGR[LQDFWLQJDV D SURWHLQ GLVXOSKLGH UHGXFWDVH %XFKDQDQ 7KH VZLWFK EHWZHHQ ³OLJKW´ DQG ³GDUN´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ynechocystis 3&& .DQHNR HW DO KDV UHYHDOHG IRXU WKLRUHGR[LQ JHQHV RQHRIZKLFKVOU LVFOHDUO\VLPLODUWRWKH DW\SLFDO Anabaena WKLRUHGR[LQRQHRIWKHPLVPW\SH VOU DQG WZR H[KLELW OHVV FHUWDLQ DIILQLWLHV VOVOU $WWHPSWVKDYHEHHQPDGHWRVWXG\ WKH OLJKWGDUN UHJXODWLRQ RI SKRWRV\QWKHWLF HQ]\PHV ZLWK LQWDFW FHOOV RIWKH F\DQREDFWHULXPNostoc 0$& $XVWLQ HW DO DQG HYLGHQFH ZDV REWDLQHG IRU WKH UHYHUVLEOH OLJKWDFWLYDWLRQ RI IUXFWRVH ELVSKRVSKDWDVH VHGRKHSWXORVHELVSKRVSKDWDVH ULEXORVHSKRVSKDWH NLQDVH DQG 1$'3OLQNHG JO\FHUDOGHK\GHSKRVSKDWH GHK\GURJHQDVH 7KH OLJKWGHSHQGHQW FRQWURO RI DFWLYLW\ RI IHUUHGR[LQ1$'3 UHGXFWDVH SRVVLEO\ YLD D WKLROGLVXOSKLGH H[FKDQJH KDV EHHQ UHSRUWHG IRU Anabaena variabilis )LOODW HW DO 7KH PRVW FOHDUFXWHYLGHQFHIURP LQ YLWUR VWXGLHV UHODWHV WR WKH R[LGDWLYH SHQWRVH SKRVSKDWH SDWKZD\ HQ]\PH *3'+ ZKHUH WKLRUHGR[LQ FRQWURO YLD D WKLROGLVXOSKLGH H[FKDQJH PHFKDQLVP KDV DOVR EHHQ
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW SURSRVHG ERWK E\ &RVVDU HW DO DQG 8GYDUG\ HW DO ,QKLELWLRQ RI *3'+ IURP Anabaena 3&& ZDV IDU PRUH VHQVLWLYH WR WKLRUHGR[LQ WKDQWRWKLRUHGR[LQ*OHDVRQ 7KLVSURSRVHG PHFKDQLVPKDVUHFHLYHGVXSSRUWIURPWKH REVHUYDWLRQ WKDWWZRF\VWHLQHUHVLGXHVDUHDEVROXWHO\FRQVHUYHGLQ DOOWKHF\DQREDFWHULDO]ZIJHQHVVRIDUVHTXHQFHG DQG DUH DEVHQW IURP WKH SURWHLQ IURP RWKHU SURNDU\RWLF VRXUFHV1HZPDQHWDO $GGLWLRQDOO\ WKHUHLV DOVR LQ YLWUR HYLGHQFH IRUWKH WKLRUHGR[LQPHGLDWHG FRQWURO RI IUXFWRVH ELVSKRVSKDWDVH DFWLYLW\ IURP Anacystis nidulans Synechococcus 3&& Anabaena cylindrica, Synechocystis 3&& DQG Anabaena variabilis 8GYDUG\ HW DO ,S HW DO 3DFROG HW DO ,Q FRQWUDVW %LVKRS UHSRUWHG QR WKLRO DFWLYDWLRQIRUWKHHQ]\PHIURPAnacystis nidulans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hlorogloeopsis fritschii E\ GLWKLRQLWH DQG JOXWDWKLRQH 0DUVWHQ DQG &RGG DQG WKHUH DUH WZR F\VWHLQH UHVLGXHV LQ WKH HQ]\PH IURPSynechocystis 3&& ZKLFK DUH LQDVLPLODUSRVLWLRQWRWKRVHLQYROYHGLQWKLRUHGR[LQ PHGLDWHG FRQWURO RI WKH SODQW HQ]\PH 6X DQG %RJRUDG 7KH S\ULGLQH QXFOHRWLGHGHSHQGHQW 'JOXFRVH GHK\GURJHQDVH RI Nostoc 0$& ZDV UHSRUWHGO\ GHDFWLYDWHG E\ WKLRUHGR[LQ -XKiV] HW DO )LQDOO\ JO\FHUDOGHK\GHSKRVSKDWH GHK\GURJHQDVH IURP Anabaena variabilis DQG Synechocystis 3&& ZKLFKERWK ODFNRQH RIWKH FRQVHUYHGF\VWHLQHV LQYROYHG LQUHJXODWLRQRIDFWLYLW\ RIWKH SODQW HQ]\PH LV LQVHQVLWLYH WR WUHDWPHQW ZLWK GLWKLRQLWH VWURQJO\ VXJJHVWLQJ DQ DEVHQFH RI
WKLRUHGR[LQ FRQWURO 7KXV WKHUH LV FRQIOLFWLQJ HYLGHQFH IRU WKH UROH RIWKLRUHGR[LQ LQ WKH OLJKWGDUN FRQWURO RI HQ]\PH DFWLYLW\ ZLWK WKH VWURQJHVW HYLGHQFHVRIDUEHLQJREWDLQHGIRUWKHOLJKWGHSHQGHQW DFWLYDWLRQ RI IUXFWRVH ELVSKRVSKDWDVH DQG LQDFWLYDWLRQ RI*3'+ +RZHYHU WKH LPSRUWDQFH RI WKH PW\SH WKLRUHGR[LQ LQ F\DQREDFWHULD LV LQGLFDWHG E\ WKH IDFW WKDW LQ FRQWUDVW WR WKH VLWXDWLRQ ZLWK E. coli, DWWHPSWVDWLQVHUWLRQDOPXWDJHQHVLVRIWKHJHQHLQ Synechococcus 3&& \LHOGHG QR YLDEOH VHJUHJDQWV ODFNLQJ WKLRUHGR[LQ 0XOOHU DQG %XFKDQDQ 2QHODVWSRLQWLVZRUWKQRWLQJ 7KH JHQH HQFRGLQJ D WKLRUHGR[LQOLNH SURWHLQ KDV EHHQ FKDUDFWHUL]HG IURP Synechococcus 3&& DQG PXWDWLRQVLQWKHWKLRUHGR[LQOLNHGRPDLQ OHDGWRFHOOV ZKLFKFDQVFDUFHO\JURZZKHUHDVPXWDWLRQVLQWKH& WHUPLQXVFRXOGOHDGWRDQLQFUHDVHGFRQWHQWRIQRUPDO SK\FRELOLVRPHV&ROOLHUDQG*URVVPDQ
% 3ODVWRTXLQRQH DQG WKH &\WRFKURPH EI &RPSOH[ *LYHQ WKH FHQWUDO SRVLWLRQ RI WKH LQWHUSKRWRV\VWHP HOHFWURQ WUDQVSRUW FKDLQ LQ WKH HQHUJ\ JHQHUDWLQJ SURFHVVHV RIF\DQREDFWHULD DQG WKH VHQVLWLYLW\ RILWV UHGR[ VWDWH WR WKH LQWHQVLW\ DQG TXDOLW\ RIOLJKW LW LV QRWVXUSULVLQJ WKDWWKHUH DUH PDQ\ UHSRUWV FRQFHUQLQJ WKH VLJQDOOLQJ UROH RI LWV FRPSRQHQWV SDUWLFXODUO\ SODVWRTXLQRQH DQG WKH F\WRFKURPH b6f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
,9 /RZ PROHFXODU :HLJKW 6LJQDOOLQJ 0ROHFXOHV $ &$03 F$03 LV DQ LPSRUWDQW VLJQDOOLQJ PROHFXOH LQ ERWK SURNDU\RWHVDQGHXNDU\RWHV,QHQWHULFEDFWHULD &$03 YLDLWVLQWHUDFWLRQZLWKWKH&53SURWHLQH[HUWVFRQWURO RYHUWKHH[SUHVVLRQRIODUJHQXPEHURIRSHURQVZKLFK DUHVHQVLWLYHWRFDWDEROLWHUHSUHVVLRQ&53FRPSOH[HG ZLWK F$03 DFWV DV D WUDQVFULSWLRQDO DFWLYDWRU YLD LQWHUDFWLRQ ZLWK WKH &WHUPLQDO GRPDLQ RI WKH VXEXQLWRIWKH IRUPRI51$SRO\PHUDVH (EULJKW DQG %XVE\ ,Q DQLPDO FHOOV F$03 DFWV DV D VHFRQGPHVVHQJHULQVLJQDOWUDQVGXFWLRQSULPDULO\YLD WKH DFWLYDWLRQ RI SURWHLQ NLQDVH $ $ YDULHW\ RI HYLGHQFH VXJJHVWV WKDWF$03 PD\ KDYH D VLJQLILFDQW SK\VLRORJLFDO UROH LQ VLJQDO WUDQVGXFWLRQ LQ F\DQREDFWHULD F$03 LQ F\DQREDFWHULD ZDV ILUVW GHWHFWHG LQ Anabaena variabilis DQG D FRUUHODWLRQ EHWZHHQ &$03 FRQWHQW DQG QLWURJHQ VWDUYDWLRQ ZDV DSSDUHQW +RRG HW DO )XUWKHU HYLGHQFH OLQNHGF$03 FRQFHQWUDWLRQV WRQLWURJHQ VWDUYDWLRQLQ A. flos-aquae )UDQFNR DQG :HW]HO DQG KHWHURF\VW SDWWHUQ IRUPDWLRQ LQ A. variabilis 6PLWK DQG 2ZQE\ 0RUH UHFHQWO\ F$03 FRQFHQWUDWLRQV KDYH EHHQ VKRZQ WR FKDQJH LQ UHVSRQVH WR OLJKWGDUN WUDQVLWLRQV DQG S+ VKLIWV LQ A. cylindrica 2KPRUL HW DO 2KPRUL DQG H[RJHQRXVO\ DGGHG F$03 DIIHFWHG ERWK UHVSLUDWRU\ DQG JOLGLQJ DFWLYLWLHV RISpirulina platensis 2KPRUL HW DO 0RVW UHFHQWO\ LW KDV EHHQ VKRZQ WKDW LQWUDFHOOXODU F$03 FRQFHQWUDWLRQV LQ Anabaena cylindrica FKDQJH UDSLGO\ LQ UHVSRQVH WR UHGIDU UHG OLJKW ZLWKUHG OLJKWFDXVLQJ DGHFUHDVHLQ F$03DQG IDU UHG FDXVLQJ DQ LQFUHDVH 2KPRUL HW DO %LDQFKLQL HW DO SDUWLDOO\ FKDUDFWHUL]HG DQ DGHQ\ODWH F\FODVH DFWLYLW\ IURP Anabaena $7&& 7KHDFWLYLW\ZKLFKZDVLQWKHVROXEOHIUDFWLRQ RIWKH FHOO DQG KDG DQ DSSDUHQW PROHFXODU ZHLJKW RI DERXWZDVDFWLYDWHGE\&D DQGERYLQHEUDLQ RU VSLQDFK FDOPRGXOLQ DQG LQKLELWHG E\ (*7$ 7KH JHQH (cya) HQFRGLQJ WKH HQ]\PH DGHQ\ODWH F\FODVH UHVSRQVLEOH IRU WKH V\QWKHVLV RI F$03 ZDV ILUVW FKDUDFWHUL]HG IURP WKH F\DQREDFWHULXP Anabaena cylindrica .DWD\DPD HW DO 7KH GHGXFHG SULPDU\ VWUXFWXUH RI WKH SURWHLQ UHYHDOHG LW WR EH D PHPEHU RIWKH XQLYHUVDO FODVV RIDGHQ\ODWH F\FODVHV 'DQFKLQ ZKLFK LQFOXGHV PHPEHUV IURP HXNDU\RWHV DQG SURNDU\RWHV DQG LW H[KLELWHG PDUNHG VLPLODULWLHV WR WKH FRQVHUYHG UHJLRQV RI DGHQ\ODWH
1+0DQQ F\FODVH IURP YDULRXV HXNDU\RWHV ,Q FRQWUDVW WKHUH ZDV QR GHWHFWDEOH VLPLODULW\ ZLWK WKH DGHQ\ODWH F\FODVHVIURPRWKHUSURNDU\RWLFFODVVHVLQFOXGLQJWKDW RI E. coli. 6HTXHQFH IHDWXUHV LQ WKH A. cylindrica HQ]\PH VXJJHVW WKDW LW KDV D VLJQDO VHTXHQFH IRU SURWHLQ H[SRUW DQG RQH WUDQVPHPEUDQH GRPDLQ 6XEFHOOXODU IUDFWLRQDWLRQ ORFDOL]HG WKH HQ]\PH WR WKH WK\ODNRLG PHPEUDQH )RXU DGHQ\ODWH F\FODVH JHQHV KDYH EHHQ FKDUDFWHUL]HG IURP Anabaena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cyaC ZDV IRXQG WR EH WKH PRVW DEXQGDQW DQG LQVHUWLRQDO PXWDJHQHVLV RI cyaC FDXVHG DPDUNHGUHGXFWLRQ LQ FHOOXODU &$03 FRQFHQWUDWLRQ $VLPLODUO\ FRPSOH[ DGHQ\ODWH F\FODVH DSSHDUVWR EHHQFRGHG E\WKH cyaC JHQH RISpirulina platensis .DVDKDUDHW DO 7KH SUHGLFWHG JHQH SURGXFWFRQWDLQV IURPWKH1WHUPLQXV D55UHFHLYHU GRPDLQ D GRPDLQ VLPLODU WR SDUW RI (75O RI Arabidopsis thaliana, DQ +3. WUDQVPLWWHU GRPDLQ D VHFRQG55UHFHLYHUGRPDLQDQGDFDWDO\WLFGRPDLQ 7KH JHQRPH RI Synechocystis 3&& HQFRGHV WZR SURWHLQV ZKLFK H[KLELW VLPLODULW\ WR DGHQ\ODWH F\FODVHV DQG DOVR D SURWHLQ GLVSOD\LQJ D PDUNHG GHJUHH RI VLPLODULW\ WR E. coli &53 ERWK LQ WKH & WHUPLQDO '1$ELQGLQJ GRPDLQ DQG WKH 1WHUPLQDO QXFOHRWLGHELQGLQJ GRPDLQ .DQHNR HW DO Synechocystis 3&& ZDV IRXQG WR H[KLELW DQ LQFUHDVHLQWKH LQWUDFHOOXODUFRQFHQWUDWLRQRIF$03LQ UHVSRQVH WR D GDUNOLJKW WUDQVLWLRQ DQG WKDW EOXH OLJKW QP ZDVUHVSRQVLEOHIRUWKHHIIHFW7HUDXFKLDQG 2KPRUL 0XWDJHQHVLV RI WKH WZR cya JHQHV UHYHDOHG WKDW WKDW cyal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
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW RU LQGHHG RYHUODS ZLWK WKH PXOWLVWHS SKRVSKRUHOD\ SDWKZD\V6HFWLRQ,,$
% &D LRQV /LNH F$03 &D LRQV DUH LPSRUWDQW DV D VHFRQG PHVVHQJHU LQ VLJQDO WUDQVGXFWLRQ LQ HXNDU\RWHV +RZHYHU WKHLU UROH LQ VLJQDO WUDQVGXFWLRQ LQ SURNDU\RWHV LV IDU OHVV FOHDUO\ HVWDEOLVKHG 7KHUH LV HYLGHQFHIRU DQ LQYROYHPHQWRI&D LRQ VLJQDOOLQJ LQ DUDQJHRISURFHVVHVLQEDFWHULD LQFOXGLQJFKHPRWD[LV LQ Bacillus subtilis DQG E. coli, DQGJOLGLQJPRWLOLW\LQ Myxococcus xanthus DQG Stigmatella aurantiaca 1RUULVHWDO ,Q F\DQREDFWHULD &D KDV EHHQ LPSOLFDWHG LQ D YDULHW\ RIFHOOXODU SURFHVVHV 6PLWK 2QHNDQG 6PLWK LQFOXGLQJ WKHPRWLOLW\ UHVSRQVHVRISpirulina subsalsa $EHOLRYLFKDQG*DQ DQG Phormidium uncinatum +DGHU DV ZHOO DV KHWHURF\VW GLIIHUHQWLDWLRQ LQ Nostoc VWUDLQ 6PLWK HW DO $Q DEXQGDQW FHOO VXUIDFH SURWHLQ LQYROYHG LQ WKH VZLPPLQJ PRWLOLW\ RIPDULQH Synechococcus VWUDLQV KDV VHYHUDO &DELQGLQJ VLWHV %UDKDPVKD 6HFWLRQ 9% DV GRHV WKH RVFLOOLQ SURWHLQ LQYROYHG LQ JOLGLQJ PRWLOLW\ +RLF]\N DQG %DXPHLVWHU (VWLPDWHV RIWKH IUHH LQWUDFHOOXODU &D FRQFHQWUDWLRQLQE. coli DUHLQWKHUDQJH WR 0 *DQJROD DQG 5RVHQ ZKLFK DUH VHYHUDO RUGHUV RI PDJQLWXGH OHVV WKDQ WKRVH RIWHQ IRXQG RXWVLGH WKH FHOO 6LPLODUO\ LQ Anabaena 3&& WKH LQWUDFHOOXODU FRQFHQWUDWLRQ RI&D LV LQ WKH UDQJH Q0 DQG PD\ DOWHU LQ UHVSRQVH WR HQYLURQPHQWDO VLJQDOV VXFK DV FROG VKRFN 7RUUHFLOOD HW DO ,Q HXNDU\RWHV WKH LQWUDFHOOXODU FRQFHQWUDWLRQ RI&D LV PDLQWDLQHG E\ WKH DFWLRQ RI &D$73DVHV KRZHYHU PDQ\ VSHFLHV RI EDFWHULD SXPS &D RXW RI WKH F\WRSODVP E\ VHFRQGDU\ WUDQVSRUW LQYROYLQJ + RU 1D DQWLSRUWV 5RVHQ ,Q IDFW &D$73DVHV GR RFFXU LQ VRPH VSHFLHV RI EDFWHULD LQFOXGLQJ F\DQREDFWHULD ,W ZDV IRXQGWKDWSynechococcus 3&& FRQWDLQV DWOHDVW WZR GLVWLQFW 3W\SH $73DVHV RQH EHORQJV WR WKH IDPLO\ RIW\SLFDOSURNDU\RWLF 3W\SH$73DVHV DQG WKH RWKHUPDUNHGO\UHVHPEOHV HXNDU\RWLF3W\SH$73DVHV .DQDPDUXHWDO ,QVHUWLRQPXWDQWVRIHLWKHURI WKHVH WZR $73DVHJHQHV ZHUH K\SHUVHQVLWLYH WR RVPRWLF VWUHVV ,QDFWLYDWLRQ RI WKH JHQH pacL HQFRGLQJ WKH HXNDU\RWLFW\SH &D$73DVH FDXVHG D FRPSOHWH ORVV RI $73GHSHQGHQW &D WUDQVSRUW DFWLYLW\ DVVRFLDWHG ZLWK WKH SODVPD PHPEUDQH EXW VRPHZKDWVXUSULVLQJO\ VXFK DPXWDQWGLG QRWH[KLELW D &DVHQVLWLYH SKHQRW\SH %HUNHOPDQ HW DO ,W PD\ EH WKDW WKH &DVHQVLWLYH SKHQRW\SH ZDV
VXSSUHVVHG E\ WKH DFWLYLW\ RI RWKHU &D$73DVHV VLQFH WKHUH DSSHDU WR EH DW OHDVW WKUHH VXFK HQ]\PHV HQFRGHG E\ WKH JHQRPH RI Synechocystis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scillatoria limnetica; WKH HYLGHQFH ZDV EDVHG RQ WKH HIIHFWV RI IOXSKHQD]LQH D &DFDOPRGXOLQLQKLELWRU RQ SKRVSKDWH XSWDNH DQG DOVR RQ WKH UDGLRLPPXQRDVVD\ TXDQWLWDWLRQRIDQDQWLJHQZKLFKFURVVUHDFWHGZLWKDQ DQWLFDOPRGXOLQ DQWLERG\ .HUVRQ HW DO 6XEVHTXHQWO\ 3HWWHUVVRQ DQG %HUJPDQ GHVFULEHG DKHDWVWDEOHIDFWRULQ H[WUDFWV RIAnabaena VWUDLQV WKDW H[KLELWHG WKH FDOPRGXOLQOLNH SURSHUW\ RI VWLPXODWLQJ FXFXPEHU 1$' NLQDVH LQ D &D GHSHQGHQW UHDFWLRQ 0RUHRYHU WKH DXWKRUV ZHUH DEOH WR GHWHFW D N'D SURWHLQ E\ ZHVWHUQ EORWWLQJ WKDW FURVVUHDFWHG ZLWK D SRO\FORQDO DQWLERG\ DJDLQVW VSLQDFK FDOPRGXOLQ 7KH DQWLJHQ ZDV GHWHFWDEOH LQ ERWKYHJHWDWLYHFHOOVDQG KHWHURF\VWV %LDQFKLQL HWDO UHSRUWHG WKH SDUWLDO SXULILFDWLRQ RI WKUHH KHDW VWDEOH SRO\SHSWLGHV IURP Anabaena $7&& FDSDEOH RI WKH &DGHSHQGHQW DFWLYDWLRQ RI ERYLQH EUDLQ F$03 SKRVSKRGLHVWHUDVH DQG WKH\ DOVR IRXQG WKDWWKHDGHQ\ODWHF\FODVHDFWLYLW\IURPWKLVVWUDLQZDV DFWLYDWHG E\ &D DQG ERYLQH EUDLQ RU VSLQDFK FDOPRGXOLQ 7KH UHTXLUHPHQW RI F\DQREDFWHULDO SKRWRV\VWHP ,, IRU &D ZDV ILUVW GHPRQVWUDWHG E\ (QJODQG DQG (YDQV DQG WKH VHQVLWLYLW\ RIWKLV &DGHSHQGHQW DFWLYLW\ WR DQWLFDOPRGXOLQ DQWLERG\ ZDV UHSRUWHG E\ 7UDPRQWLQL :HVWHUQ EORW DQDO\VLV RISKRWRV\VWHP,,SUHSDUDWLRQV XVLQJ DQ DQWL FDOPRGXOLQ DQWLERG\ GHWHFWHG D EDQG RI N'D DQG &D RYHUOD\RI6'63$*(JHOV RISKRWRV\VWHP ,, SUHSDUDWLRQV UHYHDOHG EDQGV RI DSSUR[LPDWHO\ DQG N'D0F&ROO DQG (YDQV ,W VKRXOG EH ERUQH LQ PLQG WKDW SK\FRELOLSURWHLQ VXEXQLWV ZKLFK DUH RID VLPLODU VL]H WR WKH N'D SRO\SHSWLGH DOVR
1+0DQQ
ELQG&D LQWKH RYHUOD\SURWRFRO 2QHNHWDO 0RUH UHFHQWO\ D N'D SRO\SHSWLGH ZDV LVRODWHG IURP Nostoc 3&&ZKLFKH[KLELWHG PDQ\RIWKH SURSHUWLHV DVVRFLDWHG ZLWK FDOPRGXOLQ 2QHN HW DO ,W DFWLYDWHG SHD 1$' NLQDVH LQ D &D GHSHQGHQW IDVKLRQ UHDFWHG ZLWK DQWLERGLHV UDLVHG DJDLQVW VSLQDFK FDOPRGXOLQ DQG H[KLELWHG D &D GHSHQGHQW VKLIW LQ PLJUDWLRQ RQ 6'63$*( 8QIRUWXQDWHO\ QR DPLQR DFLG VHTXHQFH LQIRUPDWLRQ ZDV REWDLQHG IRU WKH SURWHLQ DQG WKXV WKH WUXH QDWXUH RI WKLV SXWDWLYH F\DQREDFWHULDO FDOPRGXOLQ ZDV QRW HVWDEOLVKHG +RZHYHU ZKHQ DOO WKH 25)V LQ WKH JHQRPH RI Synechocystis 3&& DUH FRPSDUHG ZLWK FDOPRGXOLQ VHTXHQFHV IURP DYDULHW\ RIVRXUFHV QRVLJQLILFDQWVLPLODULWLHVDUHGHWHFWHG7KXVWKHRQO\ VDIHFRQFOXVLRQIURPDOOWKLVHYLGHQFHLV WKDW LIWKHUH LV D F\DQREDFWHULDO FDOPRGXOLQ LW LV OLNHO\ WR EH D IXQFWLRQDO DQDORJXH UDWKHU WKDQ D WUXH KRPRORJXH 7KH UHDO SK\VLRORJLFDO VLJQLILFDQFH RI &D LRQ VLJQDOOLQJLQF\DQREDFWHULDUHPDLQVWREHHVWDEOLVKHG
& *XDQRVLQHWHWUDSKRVSKDWH 7KH V\QWKHVLV RI PDFURPROHFXODU FRPSRQHQWV PXVW REYLRXVO\EHWLJKWO\FRRUGLQDWHGZLWKWKHJURZWKUDWH RI FHOO DQG JLYHQ WKH 2)7(1 UDSLGO\ IOXFWXDWLQJ HQYLURQPHQWV SUHFLVH PHFKDQLVPV KDYH HYROYHG WR FRQWUROWKHV\QWKHVLVRIU51$DQGWKHUHE\FRQWUROWKH SURWHLQ V\QWKHWLF FDSDFLW\ RIWKH FHOO &RQGRQ HW DO ,Q E. coli WKHUH DSSHDU WR EH DW OHDVW IRXU PHFKDQLVPV E\ ZKLFK U51$ DFFXPXODWLRQ LV GHWHUPLQHGDQGLQWZRRIWKHVHWKHVWULQJHQWUHVSRQVH DQG JURZWK UDWH GHSHQGHQW FRQWURO WKH NH\ LQWUDFHOOXODUVLJQDOLVWKHFRQFHQWUDWLRQRIWKHXQXVXDO QXFOHRWLGH JXDQRVLQH GLSKRVSKDWH GLSKRVSKDWH SS*SS $IXUWKHUUROHIRUSS*SSLVVXJJHVWHGE\WKH REVHUYDWLRQ WKDW LW DFWV DV D SRVLWLYH VLJQDO IRU WUDQVFULSWLRQ RI WKH rpoS JHQH HQFRGLQJ WKH IDFWRU LQYROYHG LQ WKH H[SUHVVLRQ RIPDQ\ VWDWLRQDU\ SKDVH JHQHV *HQWU\ HW DO SS*SS ZDV ILUVW GHWHFWHG LQ WKH XQLFHOOXODU F\DQREDFWHULXP Synechococcus 3&& LQ UHVSRQVH WR D VKLIW GRZQ LQ JURZWKUDWH EURXJKW DERXWE\ DUHGXFWLRQ LQ WKH LQFLGHQW OLJKW LQWHQVLW\ 0DQQ HW DO +RZHYHU WKH FRQFHQWUDWLRQ RISS*SS IHOO EHIRUH WKH DFFXPXODWLRQ RIU51$UHVWDUWHG VXJJHVWLQJWKDWWKHUH ZHUHDGGLWLRQDOPHFKDQLVPVLQYROYHG6PLWKDQG&DUU DOVR GHPRQVWUDWHG SS*SS V\QWKHVLV LQ Synechococcus 3&& XQGHUFRQGLWLRQV RIDPLQR DFLG VWDUYDWLRQ 9DULDWLRQV LQ WKH UHVSRQVHV RI GLIIHUHQW F\DQREDFWHULDO VWUDLQV ZHUH GHWHFWHG E\ $GDPVHWDO ZKRFRQILUPHGWKHSURGXFWLRQRI
SS*SSLQSynechococcus 3&& ZKHQWUDQVIHUUHG IURP WKH OLJKW WR WKH GDUN 6LPLODU HIIHFWV ZHUH REVHUYHG LQ DNostoc VWUDLQ DQGAphanocapsa EXW QRW LQ Anabaena cylindrica DQG Anabaena catenula. ,Q FRQWUDVW WKHUH DUH VHYHUDO UHSRUWV RI SS*SS DFFXPXODWLRQ LQ UHVSRQVH WR QLWURJHQ VWDUYDWLRQ LQ Anabaena cylindrica $NLQ\DQMX DQG 6PLWK DQGDOVRAnacystis nidulans )ULJD HW DO 7KH SURGXFWLRQ RI SS*SS E\ Synechococcus 3&& KDV DOVR EHHQ FRQILUPHG E\ %RUEpO\ HW DO ZKR DOVR HVWDEOLVKHG WKDW EDFWHULRSKDJH LQIHFWLRQ LQWHUIHUHG ZLWK SS*SS DFFXPXODWLRQ LQGXFHG E\ HLWKHU HQHUJ\ RU QLWURJHQ VWDUYDWLRQ 6XUiQ\L HW DO DOVR VKRZHG WKDW WKHUH ZDV DQ LQYHUVH FRUUHODWLRQ EHWZHHQ WKH UDWH RI WRWDO 51$ V\QWKHVLV DQG WKH SRRO VL]H RI SS*SS LQ Synechococcus 3&& H[FHSW LPPHGLDWHO\ DIWHU D WHPSHUDWXUH VKLIW ZKHQ SDUDOOHO LQFUHDVHV ZHUH IRXQG 7KH URXWHV RI SS*SS V\QWKHVLV LQ E. coli DUH UHDVRQDEO\ ZHOO FKDUDFWHULVHG 'XULQJ WKH VWULQJHQW UHVSRQVH WR DPLQR DFLG VWDUYDWLRQ WKH relA JHQH SURGXFW S SS*SS V\QWKHWDVH , FDWDO\VHV WKH V\QWKHVLV RISS*SSDQGWKHSHQWDSKRVSKDWHGHULYDWLYH SSS*SS SSS*SS PD\ EH FRQYHUWHG WR SS*SS E\ WKH DFWLRQRIWKHgpp JHQHSURGXFW&RQGRQHWDO 2YHUH[SUHVVLRQRI5HO$ FDXVHV DPDUNHGUHGXFWLRQLQ WKH JURZWK UDWH SUHVXPDEO\ E\ LQKLELWLQJ U51$ DFFXPXODWLRQ 7KHUH LV D VHFRQG SDWKZD\ IRU WKH V\QWKHVLV RI SS*SS HPSOR\LQJ WKH SURGXFW RI WKH spoT JHQH ZKLFK HQFRGHV D ELIXQFWLRQDO HQ]\PH S SS*SS V\QWKHWDVH ,, ZKLFK FDQ FDWDO\VH ERWK SS*SSV\QWKHVLVDQGK\GURO\VLV7KLVVHFRQGSDWKZD\ VHHPV WR EH DVVRFLDWHG ZLWK SS*SS V\QWKHVLV LQ UHVSRQVH WR IOXFWXDWLRQ LQ LQWUDFHOOXODU HQHUJ\ VRXUFH SRROV 7KH JHQRPH RI Synechocystis 3&& HQFRGHV D SRO\SHSWLGH VOU ZLWK PDUNHG VLPLODULW\ WR WKH S SS*SS S\URSKRVSKRK\GURODVH HQFRGHG E\ spoT. +RZHYHU QR KRPRORJXH RI relA FDQ EH GHWHFWHG 7KXV LW VHHPV SUREDEOH WKDW SS*SS V\QWKHVLV DQG GHJUDGDWLRQ LQ Synechocystis 3&& LV FDWDO\VHG RQO\ E\ WKH 6SR7 KRPRORJXH 7KH WUXH SK\VLRORJLFDO VLJQLILFDQFH RI SS*SS LQ F\DQREDFWHULD ZLOO RQO\ EH UHDOL]HG ZKHQ JHQH GLVUXSWLRQ H[SHULPHQWV KDYH EHHQ FDUULHG RXW LQ Synechocystis 3&& DQG SUHIHUDEO\ DOVR D KHWHURF\VWRXVVWUDLQ
' 0HWDEROLWHSRROV 7KH FRQFHQWUDWLRQV RI FHUWDLQ NH\ PHWDEROLWHV DUH OLNHO\WRDFWDVVLJQDOVUHIOHFWLQJWKHPHWDEROLFVWDWHRI
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ynechocystis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ynechococcus DQG Oscillatoria JURZQ LQ DOLJKWOGDUNF\FOH6WDODQG.UXPEHLQ 0LWVXLHW DO *UREEHODDU HW DO PRUH UHFHQWO\
WKH\ KDYH EHHQ VKRZQ LQ WKH PDULQH Trichodesmium ,06 &KHQ HW DO &KHQ HW DO ,PSRUWDQWO\ WKHVH UK\WKPV SHUVLVW LQ FRQVWDQW OLJKW IROORZLQJHQWUDLQPHQWGXULQJ JURZWK XQGHUOLJKWOGDUN F\FOLQJ WKXV IXOILOOLQJ DQRWKHU UHTXLUHPHQW RID WUXH FLUFDGLDQ UK\WKP ,QGHHG LQ Synechocystis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ynechococcus :+ 6ZHHQH\ DQG %RUJHVH DQG SKRWRV\QWKHWLF 2 SURGXFWLRQ QLWURJHQDVH DFWLYLW\ DQGFHOOGLYLVLRQLQSynechococcus 0LDPL%* 0LWVXLHWDO 6LPLODUREVHUYDWLRQVZHUHPDGH ZLWK UHJDUG WR GLQLWURJHQ IL[DWLRQ DQG DPLQR DFLG WUDQVSRUW IRU Synechococcus VWUDLQ 5) +XDQJ HW DO &KHQHWDO 7KHDSSOLFDWLRQRIJHQHWLF DQG PROHFXODUELRORJLFDO WRROV LVEHJLQQLQJ WR SURYLGH DQ XQGHUVWDQGLQJ RIWKH FORFN PHFKDQLVPV DQG RWKHU IHDWXUHV RI FLUFDGLDQ UK\WKPV LQ F\DQREDFWHULD +XDQJ HW DO HVWDEOLVKHG IRU Synechococcus 5) WKDW QLWURJHQDVH P51$ DEXQGDQFH ZDV UK\WKPLF XQGHU FRQGLWLRQV RI FRQVWDQW OLJKW LPSO\LQJ WKDW WUDQVFULSWLRQ LWVHOI RU 51$ GHJUDGDWLRQ H[KLELWHG SHULRGLFLW\ $Q LPSRUWDQW GHYHORSPHQW ZDV WKH GLVFRYHU\ WKDW WKH JHQHWLFDOO\ DPHQDEOH Synechococcus 3&& H[KLELWHG FLUFDGLDQ UK\WKPV 7KH lux JHQHV IURP Vibrio harveyii ZHUH LQWURGXFHG LQWRWKLV VWUDLQ XQGHU WKH FRQWURO RI WKH SURPRWHU RI WKH psbAI JHQH HQFRGLQJ D FRPSRQHQW RI 36,, DQG WKH FRQVHTXHQW OXPLQHVFHQFH H[KLELWHG D PDUNHG FLUFDGLDQ UK\WKP .RQGRHWDO )XUWKHUVWXGLHVZLWKWKLVV\VWHP FRQILUPHG HQWUDLQPHQW RI WKH OXPLQHVFHQFH UK\WKP GXULQJ D OLJKW GDUN F\FOH VXEVHTXHQW SHUVLVWHQFH RI WKH UK\WKP XQGHU FRQWLQXRXV OLJKW DQG WHPSHUDWXUH FRPSHQVDWLRQ .RQGR HW DO .RQGR DQG ,VKLXUD 6XFK VWXGLHV HVWDEOLVKHG WKDW WKHVH IHDWXUHVUHIOHFWHGWUDQVFULSWLRQ RIWKHpsbAI JHQH /LX HW DO D 0XWDQWV KDYHEHHQ LVRODWHG ZKLFK DUH DSSDUHQWO\DIIHFWHG LQWKHLUFLUFDGLDQUK\WKPV +XDQJ HWDO GHVFULEHGPXWDQWV RISynechococcus 5) LQ ZKLFK HLWKHURUERWK WKH GLQLWURJHQ IL[DWLRQ DQG DPLQR DFLG XSWDNH UK\WKPV ZHUH DIIHFWHG $ ODUJH
1+0DQQ
QXPEHU RI PXWDQWV RI 6\QHFKRFRFFXV 3&& DIIHFWHGLQWKHLUFLUFDGLDQUK\WKPVKDYHEHHQLVRODWHG H[KLELWLQJ D YDULHW\ RI FORFNGLVUXSWHG SKHQRW\SHV VXFK DV DOWHUHG SHULRGLFLW\ DQG DUUK\WKPLD DQG WKH PDMRULW\ RI WKHVH PXWDWLRQV FRXOG EH FRPSOHPHQWHG E\ DQ NE IUDJPHQW RIZLOG W\SH '1$ -RKQVRQ HW DO 5DQGRP VFUHHQLQJ RI IUDJPHQWV RI 6\QHFKRFRFFXV 3&& '1$ IRU FLUFDGLDQ SURPRWHU DFWLYLW\ ZLWK WKH luxAB JHQHV /LX HW DO E UHYHDOHG WKDW WKH PDMRULW\ RI'1$IUDJPHQWV ZKLFKH[KLELWHGSURPRWHU DFWLYLW\GLVSOD\HGUK\WKPLF EHKDYLRXU DQG D YDULHW\ RI GLIIHUHQW SKDVH UHODWLRQVKLSV ZHUH DSSDUHQW 7KXV WKH PHWDEROLVP RI WKH RUJDQLVP LV OLNHO\ WR EH GRPLQDWHG E\ UK\WKPLF EHKDYLRXU 7KH cis- DQG WUDQVDFWLQJFRPSRQHQWV WKDW HIIHFW WKH FLUFDGLDQ FRQWURO RI JHQH H[SUHVVLRQ KDYH VWLOO ODUJHO\WR EHGHILQHG +RZHYHU RQH JHQH IURP 6\QHFKRFRFFXV 3&& ZKLFK DIIHFWV WKH DPSOLWXGH RI WKH UK\WKP DVVRFLDWHG ZLWK VRPH SURPRWHUV KDV EHHQ VKRZQ WR HQFRGH D OLNH WUDQVFULSWLRQ IDFWRU 7KLV JHQH LV D PHPEHU RI WKH IDPLO\ RI IRXU SULQFLSDO IDFWRU JHQHV LQ WKLV RUJDQLVP 7VLQRUHPDV HW DO 6HFWLRQ ,9$ 7KHUHODWLRQVKLSRIFLUFDGLDQUK\WKPVWRWKHFHOOF\FOH KDV EHHQ LQYHVWLJDWHG DQG LW LV DSSDUHQW WKDW WKH FLUFDGLDQ FORFN LV FRPSOHWHO\ LQGHSHQGHQW RIWKH FHOO F\FOH EXW WKH WLPLQJ RI '1$ UHSOLFDWLRQ DQG FHOO GLYLVLRQ DUH QRW LQGHSHQGHQW RI WKH FLUFDGLDQ FORFN -RKQVRQHWDO .RQGRHWDO DQGWKLVLV LQNHHSLQJZLWKWKHFLUFDGLDQUK\WKPRIH[SUHVVLRQ RI dnaN DQ HVVHQWLDO '1$ UHSOLFDWLRQ JHQH /LX DQG 7VLQRUHPDV &RQWURO RIWKH FLUFDGLDQ UK\WKP LV DVVRFLDWHG ZLWK WKH kaiABC JHQH FOXVWHU DQG WKHUH LV HYLGHQFH WR VXJJHVW WKDW D QHJDWLYH IHHGEDFN RI kaiC H[SUHVVLRQ E\ .DL& JHQHUDWHV WKH FLUFDGLDQ RVFLOODWLRQZKLFKLV VXVWDLQHGE\.DL$,VKLXUDHWDO 7KH pex JHQH SURGXFW DSSHDUV WR H[WHQG WKH SHULRG RIWKH FLUFDGLDQ FORFN SRVVLEO\ E\ LQWHUDFWLQJ ZLWK.DL&.XWVXQDHWDO ,WVHHPVOLNHO\WKDW F\DQREDFWHULDO SK\VLRORJ\ LQ QDWXUH LV GRPLQDWHG E\ UK\WKPLF F\FOHV DQG WKLV FRPSOLFDWHV DWWHPSWV WR XQGHUVWDQG WKHLU HFRORJ\ EDVHG RQ LQIRUPDWLRQ IURP FXOWXUHV JURZQ XQGHU FRQVWDQW LOOXPLQDWLRQ 7KH DGDSWLYH VLJQLILFDQFH RID UHVRQDWLQJ FLUFDGLDQ FORFN KDV QRZ EHHQ GHPRQVWUDWHG E\ WKH IDFW WKDW WKH UHODWLYH ILWQHVV RI YDULRXV VWUDLQV RI F\DQREDFWHULD H[SUHVVLQJ GLIIHUHQW FLUFDGLDQ UK\WKPV ZDV RSWLPDO ZKHQ WKH FLUFDGLDQ SHULRG ZDV VLPLODU WR WKH OLJKWGDUNF\FOH
% 0RWLOLW\ 0RGHVRI0RWLOLW\DQG7D[HV &\DQREDFWHULD H[KLELW WZR IRUPV RI PRYHPHQW JOLGLQJDQGVZLPPLQJZKLFKHQDEOHWKHPWRUHVSRQG WR FKDQJHV LQ WKHLU HQYLURQPHQW )XUWKHUPRUH VRPH SODQNWRQLF VSHFLHV DUH DEOH WR SURGXFH JDV YHVLFOHV ZKLFK DGMXVW WKHLU EXR\DQF\ DQG FRQVHTXHQWO\ WKHLU SRVLWLRQ LQ WKH ZDWHU FROXPQ :DOVE\ *OLGLQJ PRWLOLW\ KDV ORQJ EHHQ UHFRJQLVHG DV D SURSHUW\ RI PDQ\ JURXSV RI F\DQREDFWHULD DQG ZDV RQFH WKRXJKW WR EH WKH RQO\ DFWLYH IRUP RI PRWLOLW\ H[KLELWHG E\ WKHVH RUJDQLVPV *OLGLQJ PRWLOLW\ LQYROYHV VHOISURSXOVLRQ DFURVV D VROLG RU VHPLVROLG VXUIDFH ZLWKRXWWKH LQYROYHPHQW RIDQ RUJDQHOOH HJ D IODJHOOXP RU D FKDQJH RI VKDSH RI WKH RUJDQLVP 3UHYLRXVO\ WKHUH KDYH H[WHQVLYH VWXGLHV RQ WKH JOLGLQJ PRWLOLW\ EHKDYLRXU RI F\DQREDFWHULD DQG WKH HFRORJLFDO VLJQLILFDQFH RI WKHVH PRYHPHQWV &DVWHQKRO] 0XFKRIWKLVHDUOLHUZRUNIRFXVHG RQ WKH PRWLOLW\ UHVSRQVHV WR OLJKW DOWKRXJK FKHPRWDFWLF DQG DJJUHJDWLYH PRYHPHQWV ZHUH REVHUYHG &DVWHQKRO] GHVFULEHV WKUHH SKRWLF PRWLOLW\ UHVSRQVHV DOO RI ZKLFK KDYH EHHQ REVHUYHG ZLWK YDULRXV VWUDLQV RIF\DQREDFWHULD 3KRWRNLQHVLV LV ZKHUH WKHUDWH RIPRYHPHQW LV HQKDQFHG RU LQKLELWHG E\ FKDQJHV LQ WKH OLJKW LQWHQVLW\ ,Q D SKRWRSKRELF UHVSRQVHDFKDQJHLQOLJKWLQWHQVLW\HOLFLWVDFKDQJHLQ PRYHPHQW XVXDOO\ D VWRS IROORZHG E\ UHYHUVDO RI GLUHFWLRQ 3KRWRWD[LV LV D SRVLWLYH RU QHJDWLYH PRYHPHQWRULHQWHGZLWKUHVSHFWWRWKHGLUHFWLRQRIWKH OLJKW )HZ VWXGLHV KDYH DGGUHVVHG WKH QDWXUH RI WKH FKHPRWDFWLFUHVSRQVHDVVRFLDWHGZLWKJOLGLQJPRWLOLW\ 1HJDWLYH FKHPRWDFWLF UHVSRQVHV WR DFLGV ZHUH REVHUYHG ZLWKVRPH 2VFLOODWRULDFHDH )HFKQHU FLWHG E\ 5LFKDUGVRQ DQG &DVWHQKRO] DQG VXOSKLGH ZLWK O. WHUHEULIRUPLV 5LFKDUGVRQ DQG &DVWHQKRO] 3RVLWLYHOLJKWGHSHQGHQWUHVSRQVHV WR &2 +&2 DQG 2 ZHUH REVHUYHG ZLWK DQRWKHU 2VFLOODWRULD 0DOLQ DQG :DOVE\ 5LFKDUGVRQ DQG&DVWHQKRO] GHVFULEHGDWUDQVLHQWLQKLELWLRQ RI JOLGLQJ PRELOLW\ LQ O. WHUHEULIRUPLV E\ IUXFWRVH 7KH GXUDWLRQ RI LQKLELWLRQ ZDV SURSRUWLRQDO WR WKH FRQFHQWUDWLRQ RIIUXFWRVH ZKLOH WKHUDWH RIPRYHPHQW IROORZLQJ UHVWRUDWLRQ RI PRWLOLW\ ZDV LQYHUVHO\ SURSRUWLRQDO WR WKH FRQFHQWUDWLRQ 6XOILGH H[HUWHG D VLPLODUHIIHFWEXWQRWJOXFRVHDFHWDWHRUODFWDWH /LWWOH LV NQRZQ DERXW WKH PHFKDQLVP RI JOLGLQJ PRWLOLW\H[FHSWWKDWLWLVDSSDUHQWO\GHSHQGHQWRQERWK WKH S+ DQG FRPSRQHQWV RI WKH SURWRQ PRWLYH
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ynechocystis 3&& /DEEH HW DO DQG Anabaena VSS *XHUUHUR%DUUHUDHWDO 5HFHQWO\WKHVXUIDFH ILEULOV RIPhormidium uncinatum ZHUH VKRZQ WR EH FRPSRVHG RI D VLQJOH URGVKDSHG N'D SURWHLQ WHUPHG RVFLOOLQ ZKLFK KDV D 1WHUPLQDO GRPDLQ FRQWDLQLQJ UHSHDWV RI D &DELQGLQJ PRWLI +RLF]\NDQG%DXPHLVWHU 7KH ILUVW UHSRUW RI D F\DQREDFWHULXP H[KLELWLQJ VZLPPLQJ PRWLOLW\ ZDV IRU PDULQH SK\FRHU\WKULQ FRQWDLQLQJ Synechococcus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
,QVHUWLRQDO LQDFWLYDWLRQ RI WKH JHQH swmA HQFRGLQJ WKLV SURWHLQ UHVXOWV LQ ORVV RI DELOLW\ WR WUDQVORFDWH EXW WKH FHOOV VWLOO JHQHUDWH WRUTXH 7KH 6ZP$ SURWHLQ LV OLNHO\ WR EH JO\FRV\ODWHG DQG FRQWDLQV D QXPEHU RI&DELQGLQJ PRWLIV ZKLFK PD\ EH LQYROYHG LQ LWV DWWDFKPHQW WR WKH FHOO VXUIDFH %UDKDPVKD 7KLV LV FRQVLVWHQW ZLWK WKH REVHUYDWLRQ WKDW FDOFLXP LV UHTXLUHG IRU VZLPPLQJ PRWLOLW\ E\ Synechococcus :+ 3LWWD HW DO 7KH WKLUG IRUP RI PRWLOLW\ H[KLELWHG E\ VRPH VSHFLHVRIF\DQREDFWHULDLVWKHFRQWURORIEXR\DQF\WR GHWHUPLQH SRVLWLRQ ZLWKLQ WKH ZDWHU FROXPQ YLD WKH IRUPDWLRQRIJDV YHVLFOHV :DOVE\ &KDSWHU 7KHH[WHUQDOVWLPXOXVWULJJHULQJJDVYHVLFOHIRUPDWLRQ DSSHDUVWREHORZLQFLGHQWOLJKWLQWHQVLW\ EXWQRWKLQJ LV \HW NQRZQ DERXW WKH QDWXUH RI WKH LQWHUQDO VLJQDO RWKHUQWKDQWKDWLWUHVXOWVLQDWUDQVFULSWLRQDOUHVSRQVH
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yxococcus
1+0DQQ
xanthus H[KLELWVJOLGLQJPRWLOLW\DQGVRFLDOEHKDYLRXU DQG WKH frz JHQHV LQYROYHG LQ WKLV PRWLOLW\ DUH KRPRORJRXV WR WKH HQWHULF EDFWHULDO che JHQHV LQYROYHGLQVZLPPLQJPRWLOLW\DQGDWOHDVWRQHRIWKH 0&3HQFRGLQJ JHQHV 6KL DQG =XVPDQ 7KXV WKHUHDUHPDUNHGVLPLODULWLHVLQWKHVLJQDOWUDQVGXFWLRQ SDWKZD\V LQYROYHG LQ WKH VZLPPLQJ DQG JOLGLQJ PRWLOLW\ RI WZR GLYHUVH JURXSV RI EDFWHULD 3URWHLQV DQWLJHQLFDOO\UHODWHGWRWKH0&3VZHUHGHWHFWHGLQWKH PHPEUDQH IUDFWLRQ RISynechocystis 3&& EXW ZHUH QRW GHWHFWHG LQ Oscillatoria tenuis 0RUJDQ HW DO $QDO\VLV RI WKH JHQRPLF VHTXHQFH RI Synechocystis 3&& UHYHDOV VHYHUDO 25)V ZLWK PDUNHG VLPLODULW\ WR FKHPRWD[LV SURWHLQV IURP RWKHU RUJDQLVPV .DQHNRHWDO 7KHUHDUHWKUHHZHOO VHSDUDWHG GLVFUHWH JURXSV RI DSSDUHQWO\ FKHPRWD[LV UHODWHG 25)V LQ ZKLFK WKHLQGLYLGXDO PHPEHU25)V DUHFRQWLJXRXVVXJJHVWLQJWKDWWKH\FRQVWLWXWHSXWDWLYH RSHURQV 7DEOH $ QRQH[KDXVWLYH DQDO\VLV RIWKH SRO\SHSWLGHV HQFRGHG E\ WKHVH SXWDWLYH RSHURQV UHYHDOV VLPLODULWLHV WR WKHFKHPRWD[LVUHODWHGSURWHLQV RI Myxococcus xanthus 6KL DQG =XVPDQ Pseudomonas aeruginosa 'DU]LQV Bacillus subtilis +DQORQ HW DO DQG Desulfovibrio vulgaris 'HFNHUVDQG9RRUGRXZ ZKLFKDUHLQ WXUQ UHODWHG WR WKH FKHPRWD[LV SURWHLQV RI E. coli $PVOHU DQG 0DWVXPDUD 2YHUDOO WKH PRVW IUHTXHQW VLPLODULWLHV ZHUH WR WKH pil JHQHV RI Ps. aeruginosa DVVRFLDWHG ZLWK SLOXV V\QWKHVLV DQG WZLWFKLQJPRWLOLW\'DU]LQV DQGLWLVVRPHZKDW VXUSULVLQJ WKDW HDFK RI WKH SXWDWLYH RSHURQV EHJLQV ZLWK WZR 25)V HDFK H[KLELWLQJ VLPLODULW\ WR WKH 3LO*+ SURWHLQV DV GRHV WKH pil RSHURQ LWVHOI &RQVHTXHQWO\ LW VHHPV YHU\ OLNHO\ WKDW FKHPRWDFWLF UHVSRQVHV LQ F\DQREDFWHULD LQYROYLQJ ERWK JOLGLQJ DQG VZLPPLQJPRWLOLW\ PD\ LQYROYHSURWHLQV VLPLODU WRWKRVHLQYROYHGLQPRWLOLW\LQRWKHUEDFWHULD
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
$
IDFWRUV
6HTXHQFHLQIRUPDWLRQIRUDODUJHQXPEHURI IDFWRUV KDV UHYHDOHG WKDWWKH\ IDOO LQWR WZR FODVVHV EDVHG RQ WKH E. coli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nabaena 3&& %UDKDPVKD DQG +DVHONRUQ IROORZLQJ WKH ELRFKHPLFDO FKDUDFWHUL]DWLRQ RIWKH PDLQ IRUP RIWKH KRORHQ]\PH 6FKQHLGHU HW DO ,W LV QRZ FOHDU WKDW WKHUH DUH PXOWLSOH JHQHV HQFRGLQJ OLNH VXEXQLWVLQVHYHUDOF\DQREDFWHULDOVWUDLQV7ZRIXUWKHU RIDFWRUJHQHVsigB DQG sigC KDYHEHHQFKDUDFWHUL]HG IURP Anabaena 3&& %UDKDPVKD DQG +DVHONRUQ DQG IRXU RIDFWRU JHQHV IURP Synechococcus 3&& 7DQDND HW DO D 7DQDNDHWDO E 6LPLODUO\ WKHUHDUHIRXUJHQHV OLNH VXEXQLWV LQ WKH JHQRPH RI HQFRGLQJ Synechocystis 3&&.DQHNRHWDO 'HYHORSPHQWDO SURFHVVHV LQ SURNDU\RWHV VXFK DV VSRUXODWLRQ LQ Bacillus subtilis, FRPPRQO\ LQYROYH VHTXHQWLDO JHQH H[SUHVVLRQ ZLWK DOWHUQDWLYH RIDFWRUV +DOGHQZDQJ 7KH SDWWHUQ RI WUDQVFULSWLRQ RI sigA LQ Anabaena 3&& LV FRPSOH[ ZLWK PXOWLSOHWUDQVFULSWVVRPHRIZKLFKYDU\LQ DEXQGDQFH XQGHU QLWURJHQIL[LQJ JURZWK FRQGLWLRQV %UDKDPVKD DQG +DVHONRUQ 7UDQVFULSWLRQ IURP sigB DQG sigC RFFXUV WUDQVLHQWO\ DIWHU QLWURJHQ VWDUYDWLRQ EXW QHLWKHUJHQH DORQHLVUHTXLUHGIRUQLWURJHQ IL[DWLRQRU KHWHURF\VW GHYHORSPHQW %UDKDPVKD DQG +DVHONRUQ 7KXV WKH UROH RIDOWHUQDWLYH VLJPD IDFWRUV LQ WKH GLIIHUHQWLDWLRQ DQG SK\VLRORJ\ RI KHWHURF\VWV UHPDLQV XQFOHDU 2QH FOHDUO\ HVWDEOLVKHG UROH IRU RQH
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW
Table 3. 7KH25)V FRQVWLWXWLQJWKUHHSXWDWLYHFKHPRWD[LV RSHURQV HQFRGHG LQ WKHJHQRPH RISynechocystis VS 3&& .DQHNR HWDO 7KHKRPRORJLHVZHUHGHWHFWHGXVLQJWKH%/$67DOJRULWKPDQGWKH6:,663527GDWDEDVHXQSXEOLVKHGUHVXOWV WKLVODERUDWRU\
2SHURQ
25)
+RPRORJXH
2UJDQLVP
VOO VO, VOO VOO 5 VOO VOO
3LO*+ 3LO*+ )U]$ 3LO'FU+ )U](
Pseudomonas aeruginosa Pseudomonas aeruginosa Myxococcus xanthus Pseudomonas aeruginosa Desulfovibrio vulgaris Myxococcus xanthus
VOO VOO VOO VOO 5 VOO
3LO*+ 3LO*+ 3LOO 3LO)U](
Pseudomonas aeruginosa Pseudomonas aeruginosa Pseudomonas aeruginosa Pseudomonas aeruginosa Myxococcus xanthus
VOU VOU VOU VOU
3LO*+ 3LO*+ &KH: 3LO-
Pseudomonas aeruginosa Pseudomas aeruginosa Bacillus subtilis Pseudomonas aeruginosa
RIWKHDGGLWLRQDO VLJPDIDFWRUV LV IRUWKDWHQFRGHG E\ WKH rpoD2 JHQH RISynechococcus 3&& ZKLFK LV LQYROYHG LQ WKH FRQWURO RI FLUFDGLDQ UK\WKPV 7VLQRUHPDV HW DO 6HFWLRQ 9$ DQG ZRXOG FRQVHTXHQWO\UHSUHVHQWD*URXS IDFWRU 7KHJURXS IDFWRU 6LJ( RI Synechococcus 3&& LV DVVRFLDWHG ZLWK WKH WUDQVFULSWLRQ RI SRVWH[SRQHQWLDO SKDVH JHQHV *UXEHU DQG %U\DQW Nostoc punctiforme $7&& FDQ HVWDEOLVK D V\PELRWLF UHODWLRQVKLS ZLWK VHYHUDO SODQWV LQFOXGLQJAnthoceros punctatus YLD LQIHFWLRQ E\ PRWLOH KRUPRJRQLXP ILODPHQWVDQGWUDQVFULSWLRQRIDJURXS IDFWRUJHQH LV WUDQVFULEHG RQO\ LQ WKH SUHVHQFH RI SODQW H[XGDWH &DPSEHOO HW DO 7KH JHQRPLF VHTXHQFH RI Synechocystis 3&& UHYHDOV WZR IDFWRUV ZKLFK SUREDEO\FDQEHFODVVLILHGDV*URXS VLQFH DOWKRXJK WKH\ VKRZ VRPH UHODWHGQHVV WR WKH\ DUH PRUH FORVHO\ UHODWHG WR WKH DQG VSRUXODWLRQVSHFLILF RIDFWRUV RI Bacillus subtilis .DQHNR HW DO $OWKRXJK JHQHV DUH ZLGHO\ GLVWULEXWHG DPRQJVW SURNDU\RWHV WKHUH LV QR GLUHFW HYLGHQFH IRU WKHLU RFFXUUHQFHLQF\DQREDFWHULD ,QGHHG WKHUHLV QR25) HQFRGHG E\ WKH JHQRPH RI Synechocystis 3&& WKDWH[KLELWVH[WHQGHG VLPLODULW\ZLWKDQ\PHPEHUVRI WKH IDPLO\+RZHYHUWKHUHLVHYLGHQFHRISURWHLQV ZLWKPDUNHG VLPLODULWLHVWRWKHHQKDQFHUOLNHSURWHLQV DVVRFLDWHG ZLWKWKH DFWLYDWLRQ RIWUDQVFULSWLRQ RI
(
)
GHSHQGHQW JHQHV $QDO\VLV RI D OLJKWUHSUHVVHG WUDQVFULSW IRXQG LQ Synechococcus 3&& UHYHDOHG DQ 25) HQFRGLQJ D SRO\SHSWLGH H[KLELWLQJ VLPLODULW\ WR WUDQVFULSWLRQDO PRGXODWRUV IURP Klebsiella pneumoniae DQG Azotobacter vinelandii 7DQHWDO 7KXV PXFKUHPDLQV WREHGRQHWR FODULI\ WKH UROH RI DOWHUQDWLYH IDFWRUV LQ F\DQREDFWHULD SDUWLFXODUO\ ZLWK UHJDUG WR WKH SK\VLRORJLFDOIXQFWLRQVRI DQG KRPRORJXHV
(
)
% 5HJXODWRU\3URWHLQV 6HYHUDO IHDWXUHV RI WKH UROH RI WUDQVFULSWLRQDO UHJXODWRU SURWHLQV DUH GHVFULEHG HOVHZKHUH LQ WKLV YROXPH &RQWURO RI H[SUHVVLRQ RIWKH psbA JHQHV LQ UHVSRQVH WR OLJKW LQWHQVLW\ DQG SURPRWHU ELQGLQJ IDFWRUV LQYROYHG LQ FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ DUH ERWK GLVFXVVHG LQ &KDSWHU DQG WKH UROHRIWKH 6PW% SURWHLQ LQFRQWURO RIPHWDOORWKLRQHLQ H[SUHVVLRQ LV FRYHUHG LQ &KDSWHU $QDO\VLV RIWKH JHQRPH RI Synechocystis 3&& KDV LGHQWLILHG VHYHUDOSXWDWLYH WUDQVFULSWLRQDO UHJXODWRU\ SURWHLQV RQ WKH EDVLV RI VHTXHQFH KRPRORJLHV .DQHNR HW DO WKRXJK UHODWLYHO\ IHZ H[DPSOHV ZLWK JRRG HYLGHQFH DERXW WKH VLJQDO WKDW WKHVH SURWHLQV DUH LQYROYHGLQWUDQVGXFLQJ
1+0DQQ
,URQLVHVVHQWLDOIRUF\DQREDFWHULD6WUDXV DV LURQ LV LQYROYHG LQ D YDULHW\ RI FHOOXODU SURFHVVHV LQFOXGLQJ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW ,URQ VWUHVV UHVXOWV LQ PDMRU SK\VLRORJLFDO DQG XOWUDVWUXFWXUDO FKDQJHV DQG LQGXFHV WKH V\QWKHVLV RI VHYHUDO SURWHLQV LQFOXGLQJ D QRYHO FKORURSK\OO ELQGLQJ SURWHLQ DQG IODYRGR[LQ ZKLFK DFWV DV D UHSODFHPHQWIRU WKH LURQFRQWDLQLQJ IHUUHGR[LQ ,Q E. coli, H[SUHVVLRQRIJHQHVLQUHVSRQVHWRLURQOLPLWDWLRQ LQYROYHV WKHfur JHQH ZKLFK HQFRGHV D '1$ELQGLQJ UHSUHVVRUSURWHLQZKLFKLV DFWLYDWHGE\WKHELQGLQJRI LURQ $ KRPRORJXH RIWKH JHQHfur LQYROYHG LQ IHUULF XSWDNH UHJXODWLRQ LQ E. coli ZDV FKDUDFWHUL]HG IURP Synechococcus 3&& DQG WKH SRO\SHSWLGH SURGXFWH[KLELWV VLPLODULW\ LGHQWLW\ WRWKH E. coli )XU SURWHLQ DQG LQFOXGHV WKH SXWDWLYH )H ELQGLQJ VLWH *KDVVHPLDQ DQG 6WUDXV ,QVHUWLRQDOPXWDJHQHVLV RIWKHF\DQREDFWHULDOfur JHQH FDXVHG FRQVWLWXWLYH V\QWKHVLV RI IODYRGR[LQ DQG K\GUR[DPDWHVLGHURSKRUHV 7ZR GLVWLQFW W\SHV RIUHJXODWRU\ SURWHLQ 1WF$ DQG 1WF% DUH WUDQVFULSWLRQDO DFWLYDWRUV WKDW KDYH EHHQ LPSOLFDWHG LQ QLWURJHQ PHWDEROLVP 7KH ntcB JHQH LV DSSDUHQWO\ LQYROYHG LQ WKH SDUWLWLRQLQJ RI IL[HG 1 &KDSWHU DQG HQFRGHV D PHPEHU RI WKH /\V5 IDPLO\ RI WUDQVFULSWLRQDO DFWLYDWRUV 6X]XNL HW DO +HQLNRIIHW DO 7KH lrrA JHQH SURGXFW RI Synechococcus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ynechocystis 3&& DQG WKHUH DUH WZR DGGLWLRQDO 25)V VOO VOU ZKLFK DSSHDUWREHORQJ WR&53)15IDPLO\ .DQHNR HWDO 7KH VWUXFWXUH RI WKH GLPHULF E. coli &53 PROHFXOH LV NQRZQ DW c UHVROXWLRQ :HEHU DQG 6WHLW] 7KH PRQRPHU VXEXQLWV FRQVLVW RI DQ DPLQRWHUPLQDO GRPDLQ FDUU\LQJ WKH &$03ELQGLQJ VLWH DQG WKH FDUER[\WHUPLQDO GRPDLQ FRQWDLQLQJ WKH KHOL[WXUQKHOL['1$ELQGLQJGRPDLQ7KH&53RIE. coli DQGSUHVXPDEO\WKHRWKHUPHPEHUVRIWKLVIDPLO\ DUH WKRXJKW WR DFW DV WUDQVFULSWLRQDO DFWLYDWRUV YLD LQWHUDFWLRQV ZLWK WKH &WHUPLQDO GRPDLQ RI WKH VXEXQLW RI 51$ SRO\PHUDVH (EULJKW DQG %XVE\
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nabaena 3&& DQG KDYHEHHQ LPSOLFDWHG LQ WKH UHVSRQVHV WR FROG VKRFN DQG UHGR[ VWDWH UHVSHFWLYHO\&KDPRWHWDO
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opcA, LPPHGLDWHO\ GRZQVWUHDP IURP WKHzwf JHQH HQFRGLQJ *3'+ LV UHTXLUHG IRU FHOOV WR H[SUHVV VLJQLILFDQW *3'+ DFWLYLW\ 0XWDWLRQV LQ WKH opcA JHQH DSSHDU WR DOWHU WKH ROLJRPHULF VWDWH RI*3'+ ZLWKRXW DOWHULQJ WKH WRWDO DPRXQW RI *3'+ SRO\SHSWLGH EHLQJ V\QWKHVLVHG 6XQGDUDP et al., 7KXV 2SF$ PD\ UHSUHVHQW DQRWKHU FRPSRQHQW RI WKH FRQWURO V\VWHP UHJXODWLQJ *3'+ DFWLYLW\ YLD DVVHPEO\ LQWR WKH IXQFWLRQDO ROLJRPHULFHQ]\PH
% 3RVWWUDQVODWLRQDO FRYDOHQWPRGLILFDWLRQV ,Q DGGLWLRQ WR SKRVSKRU\ODWLRQ SURWHLQV DUH SRWHQWLDOO\VXEMHFWWRDYDULHW\RIRWKHUPRGLILFDWLRQV LQFOXGLQJ SURWHRO\WLF FOHDYDJH $'3ULERV\ODWLRQ
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW XULG\O\ODWLRQ PHWK\ODWLRQ OLSR\ODWLRQ HWF 7KHUH LV VRPH OLPLWHG RFFXUUHQFH RI IRUPV RI SURWHLQ PRGLILFDWLRQ RWKHU WKDQ SKRVSKRU\ODWLRQ LQ F\DQREDFWHULD =HKU HW DO UHSRUWHG D GLHO PRGLILFDWLRQ UHVHPEOLQJ $'3ULERV\ODWLRQ RI WKH )HSURWHLQ RI QLWURJHQDVH LQ QDWXUDO SRSXODWLRQV RI Trichodesmium thiebautii. 6LPLODUO\ SHULRGLF PRGLILFDWLRQ RI WKH )HSURWHLQ RI QLWURJHQDVH LQ Gloeothece $7&& JURZQ XQGHU D OLJKWGDUN F\FOHZDVGHWHFWHG 'X DQG*DOORQ 6LOPDQHW DO UHSRUWHG WKH $'3ULERV\ODWLRQGHSHQGHQW LQDFWLYDWLRQ RIJOXWDPLQH V\QWKHWDVH LQ Synechocystis 3&& WKRXJKWKLVREVHUYDWLRQLVLQFRQIOLFWZLWK RWKHU UHSRUWV FRQFHUQLQJ WKH UHJXODWLRQ RI JOXWDPLQH V\QWKHWDVH DFWLYLW\ &KDSWHU +RZHYHU -RVHSK DQG 0HHNV KDYH DOVR UHSRUWHG DQ DSSDUHQW SRVWWUDQVODWLRQDO PRGLILFDWLRQ RI JOXWDPLQH V\QWKHWDVHIURPDV\PELRWLFNostoc VWUDLQ ,Q DOO RUJDQLVPV HQHUJ\GHSHQGHQW SURWHDVHV LQ DGGLWLRQ WR WKHLU FHOOXODU KRXVHNHHSLQJ UROH PD\ PHGLDWHWKHWXUQRYHURIVSHFLILFVKRUWOLYHGUHJXODWRU\ SURWHLQV ZKLFK JLYHV WKHP D FHQWUDO SRVLWLRQ LQ UHJXODWLQJWKHUHVSRQVHWRPDQ\VRUWVRISK\VLRORJLFDO VWUHVV 7KH&OS$73DVHVZHUHRULJLQDOO\LGHQWLILHGDV UHJXODWRU\ FRPSRQHQWV RIEDFWHULDO SURWHDVHV VXFK DV &OS3 RI E. coli :DZU]\QRZ HW DO DQG DUH PHPEHUV RIWKH XELTXLWRXV +63 6FKLUPHU HW DO ,WZDV VKRZQ WKDW Synechococcus 3&& FRQWDLQHG D KRPRORJXH RI WKH FKORURSODVW &OS& SURWHLQ &ODUNH DQG (ULNVVRQ DQG IURP DQDO\VLV RI WKH Synechocystis 3&& JHQRPH LW DSSHDUV WKDW WKH &OS3 SURWHDVH LWVHOILV HQFRGHG E\ D PXOWLJHQHIDPLO\.DQHNRHWDO LQFRQWUDVWWR RWKHU HXEDFWHULD ,Q Synechococcus 3&& PXWDWLRQ RIWKH clpP1 JHQH HQFRGLQJ RQH RIWKH IRXU &OS3 SURWHDVH LVR]\PHV OHG WR UHVWULFWHG JURZWK DQG OLJKW DFFOLPDWLRQ &ODUNH HW DO DV ZHOO DV IDLOXUH WR DGDSW WR 89% DQG ORZ WHPSHUDWXUHV 3RUDQNLHZLF] HW DO 0XWDWLRQ RI WKH JHQH HQFRGLQJ RQH RI WKH &OS $73DVHV &OS% DOVR VLJQLILFDQWO\ DIIHFWHG ERWK WKH GHYHORSPHQW RI WKHUPRWROHUDQFH (ULNVVRQ DQG &ODUNH DQG FROG DFFOLPDWLRQ 3RUDQNLZLF] DQG &ODUNH E\ Synechococcus 3&& 7KXV HQHUJ\GHSHQGHQW SURWHDVHV SOD\ DQ LPSRUWDQW UROH LQ UHJXODWLQJ LPSRUWDQWF\DQREDFWHULDODGDSWLYHUHVSRQVHV
9,, &URVVWDONDQG ,QWHJUDWLRQRI 6LJQDOOLQJ .H\ DUHDV RI WKH LQWHJUDWLRQ RI PHWDEROLVP LQ F\DQREDFWHULD DUH SKRWRV\QWKHVLV DQG UHVSLUDWLRQ
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ynechocystis 3&& H[KLELWHG D UDSLG UHGXFWLRQ LQ ELFDUERQDWH XSWDNH UDWH DQG WKLV ZDV DFFRPSDQLHG E\ WKH SKRVSKRU\ODWLRQ RID QXPEHU RISRO\SHSWLGHV UDQJLQJ LQVL]HIURP N'D%OR\HHWDO 6LPLODU FKDQJHVLQWKHELFDUERQDWHXSWDNHUDWHDQGWKHSURILOH RI SKRVSKRU\ODWHG SRO\SHSWLGHV ZHUH DOVR REVHUYHG ZKHQ ELFDUERQDWHOLPLWHG FXOWXUHV ZHUH VKLIWHG LQWR
1+0DQQ
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b6f FRPSOH[ 5H\HV HW DO DQG WKHUH LV HYLGHQFHIRUWKH EXONRIWKH HQ]\PH EHLQJ DVVRFLDWHG ZLWK WKH WK\ODNRLG PHPEUDQHV 6DOODO 8QGHU FRQGLWLRQVRIQLWURJHQOLPLWDWLRQSynechococcus 3&& H[KLELWHG D PDUNHG UHGXFWLRQ LQ WKH QXPEHU RI DFWLYH36,,UHDFWLRQFHQWUHV %HUJHVHWDO DQG FKDQJHV RIQLWURJHQVRXUFHHIIHFWHG DVWDWHWUDQVLWLRQ OLNH PRGLILFDWLRQ LQ WKH GLVWULEXWLRQ RI H[FLWDWLRQ HQHUJ\ LQPhormidium laminosum 2FKRDGH $OGDHW DO 6XFK HIIHFWV PD\ QRW EH FRQILQHG WR QLWURJHQ DVVLPLODWLRQ DV LQFUHDVHV LQ 36,GULYHQ WUDQVSRUW UHODWLYH WR OLQHDU HOHFWURQ WUDQVSRUW KDYH EHHQREVHUYHGIROORZLQJ3RU6GHSULYDWLRQ)RUNDQG +HUEHUW 7KH LQIOXHQFH RIWKH UHGXFWLRQ VWDWH RI34H[WHQGVWRUHJXODWLRQRIFHOOXODUGLIIHUHQWLDWLRQ ERWK ZLWK UHJDUG WR KRUPRJRQLXP IRUPDWLRQ DQG KHWHURF\VWSURGXFWLRQ LQ Calothrix VSS &DPSEHOO HW DO 7KXV DOWKRXJK LQWHUDFWLRQV EHWZHHQ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW DQG QXWULHQW DFTXLVLWLRQ V\VWHPV DUH FHQWUDO WR LQWHJUDWLQJ WKH PHWDEROLVP RI WKH FHOO RWKHU DVSHFWV RI FHOO JURZWK DQG GLIIHUHQWLDWLRQ PD\ EH GHWHUPLQHG E\ WKH UHGR[ SRLVHRIWKHWZRSKRWRV\VWHPV
9OOO &RQFOXVLRQV 7KHUH LV QRZ D FOHDU XQGHUVWDQGLQJ RI WKH W\SHV RI VLJQDOOLQJ SURFHVVHV WKDW HQDEOH F\DQREDFWHULD WR GHWHFW DQG UHVSRQG WR FKDQJHV LQ WKHLU HQYLURQPHQW ,W LV DOVR DSSDUHQW WKDW WKHVH VLJQDOOLQJ SDWKZD\V HPSOR\HOHPHQWV IURPHYHU\ OHYHO LQ WKH JHQRW\SHWR SKHQRW\SH SDWKZD\ 7KXV IRU H[DPSOH WKRXJK
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
$FNQRZOHGJHPHQWV , WKDQN 'DYH 6FDQODQ 1RHO &DUU DQG -RKQ 2UPHURG IRU WKHLU FULWLFDO UHDGLQJ RI WKH PDQXVFULSW DQG FRQVWUXFWLYHFRPPHQWV
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¶GLSKRVSKDWH ¶GLSKRVSKDWH V\QWKHVLV LQ F\DQREDFWHULD-%DFWHULRO $NLQ\DQMX - DQG 6PLWK 5- 7KH DFFXPXODWLRQ RI SKRVSKRU\ODWHG JXDQLQH QXFOHRWLGHV LQ $QDEDHQD F\OLQGULFD 1HZ3K\WRO $ODP - &XUWLV 6 *OHDVRQ ). *HUDPL1HMDG 0 DQG )XFKV -$ ,VRODWLRQ VHTXHQFHDQGH[SUHVVLRQ LQ(VFKHULFKLD FROL RI
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW DQXQXVXDOWKLRUHGR[LQJHQHIURPWKHF\DQREDFWHULXPAnabaena 3&&-%DFWHULRO $OOHQ -) 3URWHLQ SKRVSKRU\ODWLRQ LQ UHJXODWLRQ RI SKRWRV\QWKHVLV%LRFKLP%LRSK\V$FWD $OOHQ -) 5HGR[ FRQWURO RIWUDQVFULSWLRQ VHQVRUV UHVSRQVH UHJXODWRUVDFWLYDWRUVDQGUHSUHVVRUV)(%6/HWW $OOHQ -) $OH[FLHY . DQG +nNDQVVRQ * 5HJXODWLRQ E\ UHGR[VLJQDOOLQJ&XUU%LRO $PVOHU &' DQG 0DWVXPDUD 3 &KHPRWDFWLF VLJQDO WUDQVGXFWLRQ LQ Escherichia coli DQG Salmonella typhimurium. ,Q +RFK -$ DQG 6LOKDY\ 7- HGV 7ZR&RPSRQHQW 6LJQDO 7UDQVGXFWLRQSS $603UHVV:DVKLQJWRQ'& $QDQGDQ 6 1DOW\ 06 &RJGHOO '( DQG *ROGHQ 66 ,GHQWLILFDWLRQRIWZRFODVVHVRIWUDQVFULSWLRQDOUHJXODWRUJHQHV LQ WKH F\DQREDFWHULXPSynechococcus 3&& $UFK 0LFURELRO $RNL 6.RQGR7:DGD+,VKLXUD0 &LUFDGLDQUK\WKPRI WKH F\DQREDFWHULXP Synechocystis 3&& LQ WKH GDUN - %DFWHULRO $SSOHE\ -/ 3DUNLQVRQ -6 DQG %RXUUHW 5% 6LJQDO WUDQVGXFWLRQ YLD WKH PXOWLVWHS SKRVSKRUHOD\ QRW QHFHVVDULO\ D URDGOHVVWUDYHOOHG&HOO $SWH6. 5RZHOO3DQG 6WHZDUW :'3 (OHFWURQ GRQDWLRQ WR IHUUHGR[LQ LQ KHWHURF\VWV RI WKH 1IL[LQJ DOJD Anabaena cylindrica. 3URF5R\6RF% $XVWLQ 3$ 5RVV ,6 DQG 0LOOV-' /LJKWGDUN UHJXODWLRQ RI SKRWRV\QWKHWLF HQ]\PHV ZLWKLQ LQWDFW FHOOV RI WKH F\DQREDFWHULXP Nostoc VS 0$& %LRFKLP%LRSK\V $FWD %DFKRIHQ 5 DQG 6FKHQN $ 4XRUXP VHQVLQJ DXWRLQGXFHUV 'RWKH\SOD\DUROHLQQDWXUDOPLFURELDOKDELWDWV"0LFURELRO5HV %DGJHU 05 DQG 6FKUHLEHU 8 (IIHFWV RI LQRUJDQLF FDUERQ DFFXPXODWLRQ RQ SKRWRV\QWKHWLF R[\JHQ UHGXFWLRQ DQG F\FOLF HOHFWURQ IORZ LQ WKH F\DQREDFWHULXP Synechococcus 3&& 3KRWRV\QWK5HV %DUWVHYLFK99 DQG 6KHVWDNRY 69 7KH dspA JHQH SURGXFW RI WKH F\DQREDFWHULXP Synechocystis 3&& LQIOXHQFHV VHQVLWLYLW\ WR FKHPLFDOO\ GLIIHUHQW JURZWK LQKLELWRUV DQG KDV DPLQRDFLG VLPLODULW\WRKLVWLGLQHSURWHLQNLQDVHV 0LFURELRORJ\ %HUJHV -$ &KDUOHERLV '2 0DX]HUDOO '& DQG )DONRZVNL 3* 'LIIHUHQWLDO HIIHFWV RI QLWURJHQ OLPLWDWLRQ RQ SKRWRV\QWKHWLFHIILFLHQF\RISKRWRV\VWHPV, DQG,,LQPLFURDOJDH 3ODQW3K\VLRO %HUNHOPDQ 7 *DUUHWHQJHOH 3 DQG +RIIPDQ 1( 7KH pacL JHQHVRISynechococcus 3&&HQFRGHVD&DWUDQVSRUWLQJ $73DVH-%DFWHULRO %LDQFKLQL *0 3DVWLQL $& 0XVFKLHWWL -3 7pOOH],xRQ 07 0DUWLQHWWR +7 7RUUHV +1 DQG )ODZLD 00 $GHQ\ODWH F\FODVHDFWLYLW\LQF\DQREDFWHULD DFWLYDWLRQE\&DFDOPRGXOLQ DQG FDOPRGXOLQOLNH DFWLYLW\ %LRFKLP %LRSK\V $FWD %LVKRS 5+ 5HJXODWRU\ FKDUDFWHULVWLFV RI D IUXFWRVH ELVSKRVSKDWDVH IURP WKH EOXHJUHHQ EDFWHULXP Anacystis nidulans. $UFK%LRFKHP%LRSK\V %OR\H6$ 6LOPDQ1-0DQQ1+DQG&DUU1* %LFDUERQDWH FRQFHQWUDWLRQE\Synechocystis 3&& PRGXODWLRQRISURWHLQ SKRVSKRU\ODWLRQ DQG LQRUJDQLF FDUERQ WUDQVSRUW E\ JOXFRVH 3ODQW3K\VLRO %RUEpO\*.DUL&*XO\iV$DQG)DUNDV*/ %DFWHULRSKDJH LQIHFWLRQ LQWHUIHUHV ZLWK JXDQRVLQH¶GLSKRVSKDWH¶ GLSKRVSKDWH DFFXPXODWLRQ LQGXFHG E\ HQHUJ\ DQG QLWURJHQ
VWDUYDWLRQLQWKHF\DQREDFWHULXPAnucystis nidulans. -%DFWHULRO %UDKDPVKD % $Q DEXQGDQW FHOOVXUIDFH SRO\SHSWLGH LV UHTXLUHG IRU VZLPPLQJ E\ WKH QRQIODJHOODWHG PDULQH F\DQREDFWHULXP Synechococcus. 3URF 1DWO $FDG 6FL 86$ %UDKDPVKD % DQG +DVHONRUQ 5 ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RIWKH JHQH HQFRGLQJ WKH SULQFLSDO VLJPDIDFWRU RIWKHYHJHWDWLYHFHOO51$SRO\PHUDVHIURPWKHF\DQREDFWHULXP Anabaena 3&&-%DFWHULRO %UDKDPVKD % DQG +DVHONRUQ 5 ,GHQWLILFDWLRQ RI PXOWLSOH 51$ SRO\PHUDVH VLJPD IDFWRUKRPRORJV LQ WKH F\DQREDFWHULXP Anabaena 3&& - %DFWHULRO %XFKDQDQ %% 5HJXODWLRQ RI&2 PHWDEROLVP LQ R[\JHQLF SKRWRV\QWKHVLV WKH IHUUHGR[LQWKLRUHGR[LQ V\VWHP $UFK %LRFKHP%LRSK\V %XVWRV 6$ 6FKDHIHU 05 DQG *ROGHQ 66 'LIIHUHQW DQG UDSLG UHVSRQVHV RI F\DQREDFWHULDOpsbA WUDQVFULSWV WR FKDQJHV LQOLJKWLQWHQVLW\-%DFWHULRO &DPSEHOO '$ +RXPDUG - DQG 7DQGHDX GH 0DUVDF 1 (OHFWURQ WUDQVSRUW UHJXODWHV FHOOXODU GLIIHUHQWLDWLRQ LQ WKH ILODPHQWRXVF\DQREDFWHULXPCalothrix. 3ODQW&HOO &DPSEHOO (/ %UDKDPVKD % DQG 0HHNV -& ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WKH Nostoc punctiforme sigH DQG ctpH JHQHV$EVWUDFWVRIWKH9,&\DQREDFWHULDO:RUNVKRS% &DPSEHOO(/+DJHQ.'&RKHQ0)6XPPHUV0/DQG0HHNV-& 7KHdevR JHQHSURGXFWLVFKDUDFWHULVWLFRIUHFHLYHUVRI FRPSRQHQW UHJXODWRU\ V\VWHPV DQG LV HVVHQWLDO IRU KHWHURF\VW GHYHORSPHQW LQ WKH ILODPHQWRXV F\DQREDFWHULXP Nostoc $7&& -%DFWHULRO &DUU1* 0HWDEROLFFRQWURODQGDXWRWURSKLFSK\VLRORJ\,Q &DUU1*DQG :KLWWRQ %$HGV 7KH%LRORJ\RIWKH %OXH*UHHQ $OJDHSS%ODFNZHOO6FLHQWLILF3XEOLFDWLRQV2[IRUG &DVWHQKRO] 5: 0RWLOLW\ DQG WD[HV ,Q &DUU 1* DQG :KLWWRQ %$ HGV 7KH %LRORJ\ RI &\DQREDFWHULD SS %ODFNZHOO6FLHQWLILF3XEOLFDWLRQV2[IRUG &KDPRW'.XMDW6<X(DQG2ZWWULP*: &ROGDQGUHGR[ LQGXFHG F\DQREDFWHULDO 51$ KHOLFDVHV $EVWUDFWV RI WKH 9, &\DQREDFWHULDO:RUNVKRS$ &KDQJ & .ZRN 6) %OHHFNHU $% DQG 0H\HURZLW] (0 Arabidopsis HWK\OHQHUHVSRQVH JHQH (75O VLPLODULW\ WR FRPSRQHQWUHJXODWRUV 6FLHQFH &KDQJ & DQG 0H\HURZLW] (0 7KH HWK\OHQH UHVSRQVH LQ Arabidopsis DHXNDU\RWLF FRPSRQHQWVLJQDOOLQJV\VWHP 3URF 1DWO $FDG6FL 86$ &KHQ7+ &KHQ7/+XQJ/0DQG+XDQJ7& FLUFDGLDQ UK\WKP LQ DPLQR DFLG XSWDNH E\ Synechococcus 5) 3ODQW 3K\VLRO &KHQ<% 'RPLQLF% 0HOORQ07DQG=HKU-3 &LUFDGLDQ UK\WKP RI QLWURJHQDVH JHQH H[SUHVVLRQ LQ WKH GLD]RWURSKLF ILODPHQWRXV QRQKHWHURF\VWRXV F\DQREDFWHULXP Trichodesrniurn ,06-%DFWHULRO &KHQ <% =HKU -3 DQG 0HOORQ 0 *URZWK DQG QLWURJHQ IL[DWLRQ RI WKH GLD]RWURSKLF ILODPHQWRXV QRQKHWHURF\VWRXV F\DQREDFWHULXP Trichodesrniurn VS ,06 LQGHILQHGPHGLD HYLGHQFHIRUDFLUFDGLDQUK\WKP -3K\FRO &KHUU\ -5 +RQGUHG ' :DONHU -0 .HOOHU -0 +HUVKH\ +3 9LHUVWUD 5' &DUER[\WHUPLQDO GHOHWLRQ DQDO\VLV RI RDW SK\WRFKURPH $ UHYHDOV WKH SUHVHQFH RI VHSDUDWH GRPDLQV UHTXLUHGIRU VWUXFWXUHDQGELRORJLFDO DFWLYLW\ 3ODQW &HOO &KLDQJ *& 6FKDHIHU 05 DQG *URVVPDQ $5 &RPSOHPHQWDWLRQ RI D UHG OLJKWLQGLIIHUHQW F\DQREDFWHULDO PXWDQW3URF1DWO$FDG6FL86$
&ODQFH\&-DQG*LOEHUW+) 7KLROGLVXOILGHH[FKDQJHLQ WKH WKLRUHGR[LQFDWDO\VHGUHGXFWLYHDFWLYDWLRQRIVSLQDFK FKORURSODVW IUXFWRVHELVSKRVSKDWDVH-%LRO&KHP &ODUNH $. DQG (ULNVVRQ 0- 7KH F\DQREDFWHULXP Synechococcus VS 3&& SRVVHVVHV D FORVH KRPRORJXH WR WKHFKORURSODVW&OS&SURWHLQRIKLJKHUSODQWV3ODQW0RO%LRO &ROOLHU -/ DQG *URVVPDQ $5 'LVUXSWLRQ RI D JHQH HQFRGLQJ D QRYHO WKLRUHGR[LQOLNH SURWHLQ DOWHUV WKH F\DQREDFWHULDOSKRWRV\QWKHWLF DSSDUDWXV - %DFWHULRO &RQGRQ & 6TXLUHV & DQG 6TXLUHV &/ &RQWURO RI U51$ WUDQVFULSWLRQLQEscherichia coli. 0LFURELRO5HY &RUWD\ -& %OHLFKHU ) 5LHXO & 5HHYHV +& DQG &R]]RQH $- 1XFOHRWLGH VHTXHQFH DQG H[SUHVVLRQ RI WKH aceK JHQH FRGLQJ IRU LVRFLWUDWH GHK\GURJHQDVH NLQDVHSKRVSKDWDVH LQ Escherichia coli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seudomonas aeruginosa JHQH FOXVWHU LQYROYHG LQ SLOXV ELRV\QWKHVLV DQG WZLWFKLQJ PRWLOLW\ VHTXHQFH VLPLODULW\ WR WKH FKHPRWD[LV SURWHLQV RI HQWHULFV DQG WKH JOLGLQJ EDFWHULXP Myxococcus xanthus. 0ROHF 0LFURELRO 'HFNHUV +0 DQG 9RRUGRXZ * ,GHQWLILFDWLRQ RI D ODUJH IDPLO\ RI JHQHV IRU SXWDWLYH FKHPRUHFHSWRU SURWHLQV LQ DQ RUGHUHG OLEUDU\ RI WKH Desulfovibrio vulgaris +LOGHQERURXJK JHQRPH-%DFWHULRO 'X & DQG*DOORQ-5 0RGLILFDWLRQ RIWKH)HSURWHLQ RIWKH QLWURJHQDVH RI Gloeothece 1DJHOL VS $7&& GXULQJ JURZWK XQGHU DOWHUQDWLQJ OLJKW DQG GDUNQHVV 1HZ 3K\WRORJLVW (EULJKW 5+ DQG %XVE\ 6 7KH Escherichia coli 51$ SRO\PHUDVH a VXEXQLW VWUXFWXUH DQG IXQFWLRQ &XUU 2SLQLRQ *HQHW'HYHO (KOHUV.06DPXHO$'7%HUJ+&DQG0RQWJRPHU\5 'R F\DQREDFWHULD VZLP XVLQJ WUDYHOOLQJ VXUIDFH ZDYHV" 3URF 1DWO $FDG6FL86$ (QJODQG 55 DQG (YDQV + $ UHTXLUHPHQW IRU &D LQ WKH H[WUDFWLRQ RI2HYROYLQJ SKRWRV\VWHPV SUHSDUDWLRQV IURP WKH F\DQREDFWHULXP Anacystis nidulans. %LRFKHP- (VFK -- 5RGULJXH] ), %LQGHU %0 +HW]HO &( DQG %OHHFNHU $% 6WUXFWXUH IXQFWLRQDQGHYROXWLRQDU\RULJLQRIWKHFRSSHU EDVHGHWK\OHQHELQGLQJ GRPDLQ RIWKH (75O UHFHSWRU $EVWUDFWV RIWKH,QWHUQDWLRQDO&RQIHUHQFHRQArabidopsis 5HVHDUFK )HFKQHU 5 'LH FKHPRWD[LV GHU 2VFLOODWRULHQ XQG LKUH %HZHJXQJVHUVFKHLQXQJHQXEHUKDXSW=%RW )LOODW 0) (GPRQGVRQ '( DQG *RPH]0RUHQR & /LJKW GHSHQGHQW GHDFWLYDWLRQKHDFWLYDWLRQ RI Anabaena variabilis IHUUHGR[LQ1$'3 UHGXFWDVH%LRFKHP- )RUFKKDPPHU . DQG +HGOHU $ 3KRVSKRSURWHLQ 3,, IURP F\DQREDFWHULD $QDO\VLV RIIXQFWLRQDO FRQVHUYDWLRQ ZLWK WKH 3 ,, VLJQDOWUDQVGXFWLRQ SURWHLQ IURP Escherichia coli. (XU - %LRFKHP
1+0DQQ )RUFKKDPPHU.DQG7DQGHDXGH0DUVDF 1 7KH3,,SURWHLQ LQWKHF\DQREDFWHULXPSynechococcus 3&&LVPRGLILHGE\ VHULQH SKRVSKRU\ODWLRQ DQG VLJQDOV WKH FHOOXODU 1VWDWXV - %DFWHULRO )RUFKKDPPHU . DQG 7DQGHDX GH 0DUVDF 1 D )XQFWLRQDO DQDO\VLV RI WKH SKRVSKRSURWHLQ 3,, (glnB JHQH SURGXFW LQ WKH F\DQREDFWHULXP Synechococcus 3&& - %DFWHULRO )RUFKKDPPHU . DQG 7DQGHDX GH 0DUVDF 1 E 3KRVSKRU\ODWLRQ RI WKH 3,, SURWHLQ (glnB JHQH SURGXFW LQ WKH F\DQREDFWHULXP Synechococcus 3&& DQDO\VLV RIin vitro NLQDVHDFWLYLW\-%DFWHULRO )RUN '& DQG +HUEHUW 6. (OHFWURQ WUDQVSRUW DQG SKRWRSKRVSKRU\ODWLRQ E\ SKRWRV\VWHP , LQ YLYR LQ SODQWV DQG F\DQREDFWHULD3KRWRV\Q5HV )ULJD * %RUEpO\ * DQG )DUNDV */ $FFXPXODWLRQ RI JXDQRVLQH WHWUDSKRVSKDWH SS*SS XQGHU QLWURJHQ VWDUYDWLRQ LQ Anacystis nidulans, D F\DQREDFWHULXP $UFK 0LFURELRO )UDQFNR '$ DQG :HW]HO 5* '\QDPLFV RI FHOOXODU DQG H[WUDFHOOXODU F$03 LQ Anabaena flos-aquae &\DQRSK\WD LQWULQVLF FXOWXUH YDULDELOLW\ DQG FRUUHODWLRQ ZLWK PHWDEROLF YDULDEOHV-3K\FRO )XTXD:&:LQDQV6&DQG*UHHQEHUJ(3 4XRUXPVHQVLQJ LQ EDFWHULD WKH /X[5/X[, IDPLO\ RI FHOO GHQVLW\UHVSRQVLYH WUDQVFULSWLRQDOUHJXODWRUV-%DFWHULRO *DQJROD 3 DQG 5RVHQ %3 0DLQWHQDQFH RI LQWUDFHOOXODU FDOFLXPLQEscherichia coli. -%LRO&KHP *HQWU\'U+HUQDQGH]9- 1JX\HQ /+ -HQVHQ '% DQG&DVKHO 0 6 LV 6\QWKHVLV RI WKH VWDWLRQDU\SKDVH VLJPD IDFWRU SRVLWLYHO\UHJXODWHGE\SS*SS-%DFWHULRO *KDVVHPLDQ0DQG 6WUDXV1$ )XUUHJXODWHVWKHH[SUHVVLRQ RILURQVWUHVV JHQHV LQWKH F\DQREDFWHULXP Synechococcus 3&& 0LFURELRORJ\ *OHDVRQ ). 7KLRUHGR[LQV LQ F\DQREDFWHULD VWUXFWXUH DQG UHGR[ UHJXODWLRQ RI HQ]\PH DFWLYLW\ ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU DFDGHPLF3XEOLVKHUV'RUGUHFKW 7KH1HWKHUODQGV *OHDVRQ ). *OXFRVHSKRVSKDWH GHK\GURJHQDVH IURP WKH F\DQREDFWHULXP Anabaena VS 3&& SXULILFDWLRQ DQG NLQHWLFV RIUHGR[PRGXODWLRQ $UFK%LRFKHP%LRSK\V *ROGHQ 66 ,VKLXUD 0 -RKQVRQ &+ DQG .RQGR 7 &\DQREDFWHULDOFLUFDGLDQUK\WKPV$QQX5HY3ODQW3K\VLRO3ODQW 0RO %LRO *URKEHODDU 1 +XDQJ 7& /LQ +< DQG &KRZ 7- 'LQLWURJHQIL[LQJ HQGRJHQRXV UK\WKP LQ Synechococcus 5) )(060LFURELRO/HWW *URVVPDQ $5 DQG .HKRH '0 3KRVSKRUHOD\ FRQWURO RI SK\FRELOLVRPH ELRJHQHVLV GXULQJ FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ3KRWRV\QWK5HV *URVVPDQ$DQG0F*RZDQ5( 5HJXODWLRQRIWKHJOXFRVH SKRVSKDWH GHK\GURJHQDVH LQ EOXHJUHHQ DOJDH 3ODQW 3K\VLRO *URVVPDQ $5 6FKDHIHU 05 &KLDQJ ** DQG &ROOLHU -/ 7KH SK\FRELOLVRPH D OLJKWKDUYHVWLQJ FRPSOH[ UHVSRQVLYH WR HQYLURQPHQWDOFRQGLWLRQV0LFURELRO5HY *UXEHU 70 DQG %U\DQW '$ &KDUDFWHUL]DWLRQ RI WKH DOWHUQDWLYHVLJPDIDFWRUV 6LJ'DQG6LJ(LQSynechococcus 3&& 6LJ( LV LPSOLFDWHG LQ WUDQVFULSWLRQ RI SRVWH[SRQHQWLDO SKDVHVSHFLILFJHQHV$UFK0LFURELRO *XHUUHUR%DUUHUD $/ *DUFLD&XpOODU &0 9LOODOED -' 6HJXUD 1HULWR 0 *yPH]/RMHUR & 5H\HV 0( +HUQiQGH] -0 *DUFLD 50 DQG GH OD *DU]D 0 $FWLQUHODWHG SURWHLQV LQ
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW Anabaena VSS DQG Escherichia FROL 0LFURELRORJ\ +DGHU '3 *DWHG LRQ IOX[HV LQYROYHG LQ SKRWRSKRELF UHVSRQVHV RI WKH EOXHJUHHQ DOJD Phonidium uncinatum. $UFK 0LFURELRO +DJHPDQQ0*ROOGDFN' %LJJLQV-DQG(UGPDQQ1 6DOW GHSHQGHQW SURWHLQ SKRVSKRU\ODWLRQ LQ WKH F\DQREDFWHULXP Synechocystis VS 3&& )(06 0LFURELRO /HWW +DOGHQZDQJ :* 7KH VLJPD IDFWRUV RI Bacillus subtilis. 0LFURELRO5HY +DQNV .+ 4XLQQ $0 DQG +XQWHU 7 7KH SURWHLQ NLQDVH IDPLO\ FRQVHUYHG IHDWXUHV DQG GHGXFHG SK\ORJHQ\ RI WKH FDWDO\WLFGRPDLQV6FLHQFH +DQORQ':0DUTXH]0DJDQD/0&DUSHQWHU3%&KDPEHUOLQ0- DQG2UGDO*: 6HTXHQFHDQGFKDUDFWHUL]DWLRQRIBacillus subtilis &KH:-%LRO&KHP +DUULVRQ 0$ 0ROHFXODU PHFKDQLVPV RI DGDSWDWLRQ LQ WKH SKRWRV\QWKHWLFDSSDUDWXV3K'WKHVLV8QLYHUVLW\RI/HHGV +DUULVRQ 0$ .HHQ -1 )LQGOD\ -%& DQG $OOHQ -) 0RGLILFDWLRQ RI D *OQ%OLNH JHQHSURGXFW E\ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW LQ WKH F\DQREDFWHULXP Synechococcus )(%6/HWW +DVHONRUQ 5 'HYHORSPHQWDOO\ UHJXODWHG JHQH UHDUUDQJHPHQWVLQSURNDU\RWHV$QQ5HY*HQHW +HQLNRII 6 +DXJKQ *: &DOYR -0 DQG :DOODFH -& $ ODUJH IDPLO\ RIEDFWHULDO DFWLYDWRUSURWHLQV 3URF 1DWO $FDG 6FL 86$ +HUEHUW 6. 0DUWLQ 5( DQG )RUN '& /LJKW DGDSWDWLRQ RI F\FOLF HOHFWURQ WUDQVSRUW WKURXJK SKRWRV\VWHP , LQ WKH F\DQREDFWHULXP Synechococcus VS 3&& 3KRWRV\QWK 5HV +RFK-$DQG6LOKDY\7- 7ZR&RPSRQHQW6LJQDO 6\VWHPV $603UHVV:DVKLQJWRQ'& +RLF]\N ( DQG %DXPHLVWHU : (QYHORSH VWUXFWXUH RIIRXU JOLGLQJILODPHQWRXVF\DQREDFWHULD-%DFWHULRO +RLF]\N ( DQG %DXPHLVWHU : 2VFLOOLQ DQ H[WUDFHOOXODU &DELQGLQJ JO\FRSURWHLQ HVVHQWLDO IRU WKH JOLGLQJ PRWLOLW\ RI F\DQREDFWHULD 0ROHF 0LFURELRO +RRG ( $UPRXU 6 2ZQE\ -' +DQGD $. %UHVVDQ 5$ (IIHFW RI QLWURJHQ VWDUYDWLRQ RQ WKH OHYHO RI DGHQRVLQH ¶¶ PRQRSKRVSKDWH LQAnabaena variabilis. %LRFKLP %LRSK\V $FWD +RZHOO /' *ULIILWKV & 6ODGH /: 3RWWV 0 DQG .HQQHOO\ 3- 6XEVWUDWH VSHFLILFLW\ RI ,SK3 D F\DQREDFWHULDO GXDO VSHFLILFLW\ SURWHLQ SKRVSKDWDVH ZLWK 0$3 NLQDVH SKRVSKDWDVH DFWLYLW\%LRFKHPLVWU\ +XD - &KDQJ & 4XQ 4 DQG 0H\HURZLW] (0 (WK\OHQH LQVHQVLWLYLW\ FRQIHUUHG E\Arabidopsis (56 JHQH 6FLHQFH +XDQJ 7& DQG *UREEHODDU 1 7KH FLUFDGLDQ FORFN LQ WKH SURNDU\RWH Synechococcus 5) 0LFURELRORJ\ +XDQJ 7& 7X - &KRZ 7- DQG &KHQ 7+ &LUFDGLDQ UK\WKPRIWKHSURNDU\RWHSynechococcus VS 5) 3ODQW3K\VLRO +XDQJ7&:DQJ67DQG*UREEHODDU1 &LUFDGLDQUK\WKP PXWDQWVRIWKHSURNDU\RWLF Synechococcus 5) &XUU0LFURELRO +XJKHV - /DPSDUWHU 7 0LWWPDQQ ) +DUWPDQ ( *lUWQHU : :LOGH $ DQG %RPHU 7 $ SURNDU\RWLF SK\WRFKURPH 1DWXUH ,S605RZHOO3$LWNHQ$DQG6WHZDUW:'3 3XULILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WKLRUHGR[LQ IURP WKH 1IL[LQJ
F\DQREDFWHULXP Anabaena cylindrica. (XU- %LRFKHP ,UPOHU $ 6DQQHU 6 'LHUNV + DQG )RUFKKDPPHU . 'HSKRVSKRU\ODWLRQ RIWKHSKRVSKRSURWHLQ3,LQSynechococcus 3&& LGHQWLILFDWLRQ RI DQ $73 DQG R[RJOXWDUDWH UHJXODWHGSKRVSKDWDVH0ROHF0LFURELRO ,VKLXUD0.XWVXQD6$RNL6 ,ZDVDNL+$QGHUVVRQ&57DQDEH $*ROGHQ66-RKQVRQ&+DQG.RQGR7 ([SUHVVLRQRID JHQH FOXVWHU kaiABC DV D FLUFDGLDQ IHHGEDFN SURFHVV LQ F\DQREDFWHULD6FLHQFH -LDQJ ) 0DQQHUYLN % %HUJPDQ % (YLGHQFH IRU UHGR[ UHJXODWLRQ RI WKH WUDQVFULSWLRQ IDFWRU 1WF$ DFWLQJ ERWK DV DQ DFWLYDWRUDQG DUHSUHVVRU LQWKHF\DQREDFWHULXP Anabaena 3&& %LRFKHP- -RKQVRQ&+*ROGHQ66,VKLXUD0DQG.RQGR7 &LUFDGLDQ FORFNVLQSURNDU\RWHV 0ROHF0LFURELRO -RVHSK &0 DQG 0HHNV -& 5HJXODWLRQ RI H[SUHVVLRQ RI JOXWDPLQH V\QWKHWDVH LQ D V\PELRWLF Nostoc VWUDLQ DVVRFLDWHG ZLWK Anthoceros punctatus. -%DFWHULRO -XKDV] $ &VL]PDGLD 9 %RUEpO\ * 8GYDUG\ - DQG )DUNDV */ 7KH S\ULGLQH QXFOHRWLGHGHSHQGHQW 'JOXFRVH GHK\GURJHQDVHRINostoc 0$& DF\DQREDFWHULXP LV VXEMHFWWR WKLRUHGR[LQPRGXODWLRQ)(%6/HWW .DQDPDUX . .DVKLZDJL 6 DQG 0L]XQR 7 7KH F\DQREDFWHULXP Synechococcus VS 3&& SRVVHVVHV WZR GLVWLQFW JHQHV HQFRGLQJ FDWLRQWUDQVSRUWLQJ 3W\SH $73DVHV )(%6/HWW .DQHNR76DWR6 .RWDQL+7DQDND$$VDPL]X(1DNDPXUD< 0L\DMLPD1 +LURVDZD 0 6XJLXUD 0 6DVDPRWR 6 .LPXUD7 +RVRXFKL 7 0DWVXQR $ 0XUDNL $ 1DND]DNL 1 1DUXR . 2NXPXUD 6 6KLPSR 6 7DNHXFKL & :DGD 7 :DWDQDEH $
.RQGR70RUL7/HEHGHYD19$RNL6,VKLXUD0DQG*ROGHQ66 &LUFDGLDQ UK\WKPV LQ UDSLGO\ GLYLGLQJ F\DQREDFWHULD 6FLHQFH .RQGR76WUD\HU&$.XONDPL5'7D\ORU: ,VKLXUD0*ROGHQ 66 DQG -RKQVRQ &+ &LUFDGLDQ UK\WKPV LQ SURNDU\RWHV OXFLIHUDVH DV D UHSRUWHU RI FLUFDGLDQ JHQH H[SUHVVLRQ LQ F\DQREDFWHULD3URF1DWO$FDG6FL86$ .RWDQL+DQG7DEDWD6 /HVVRQVIURPWKHVHTXHQFLQJRIWKH JHQRPH RI D XQLFHOOXODU F\DQREDFWHULXP Synechocystis VS 3&& $QQX5HY3ODQW3K\VLRO3ODQW0RO%LRO .XVWX 6 6DQWHUR ( .HHQHU - 3RSKDP ' DQG :HLVV ' ([SUHVVLRQRI QWU$ GHSHQGHQWJHQHV LV SUREDEO\XQLWHG E\ DFRPPRQPHFKDQLVP0LFURELRO5HY .XWVXQD 6 .RQGR 7 $RNL 6 DQG ,VKLXUD 0 $ SHULRG H[WHQGHUJHQHSH[WKDWH[WHQGVWKHSHULRGRIWKHFLUFDGLDQFORFN LQ WKH F\DQREDFWHULXP Synechococcus 3&& - %DFWHULRO /DEEH-3 +DUULFDQH 0& %R\HU0 'HUDQFRXUW- 5RXVWDQ & DQG %HQ\DPLQ< %LRFKHPLFDOHYLGHQFHIRUWKHSUHVHQFHRIDQ XQFRQYHQWLRQDO DFWLQ SURWHLQ LQ D SURNDU\RWLF RUJDQLVP &RPS %LRFKHPDQG3K\VLRO% /DJDULDV -& DQG /DJDULDV '0 6HOIDVVHPEO\ RIV\QWKHWLF SK\WRFKURPHKRORSURWHLQ in vitro. 3URF1DWO $FDG6FL 86$ /DPSDUWHU 7 0LWWPDQQ ) *lUWQHU : %RPHU +DUWPDQQ ( DQG +XJKHV - &KDUDFWHUL]DWLRQ RIUHFRPELQDQW SK\WRFKURPH IURP WKH F\DQREDFWHULXP Synechocystis. 3URF 1DWO $FDG 6FL 86$ /DXGHQEDFK '( DQG *URVVPDQ $5 &KDUDFWHUL]DWLRQ DQG PXWDJHQHVLV RI VXOIXUUHJXODWHG JHQHV LQ D F\DQREDFWHULXP HYLGHQFHIRUIXQFWLRQLQVXOIDWHWUDQVSRUW-%DFWHULRO /HH+0)ORUHV(+HUUHUR$+RXPDUG-DQG7DQGHDXGH0DUVDF 1 $ UROH IRU WKH VLJQDO WUDQVGXFWLRQ SURWHLQ 3,, LQ WKH FRQWURO RI QLWUDWHQLWULWH XSWDNH LQ D F\DQREDFWHULXP )(%6 /HWWHUV /H[ 0 DQG &DUU 1* 7KH PHWDEROLVP RI JOXFRVH E\ KHWHURF\VWV DQG YHJHWDWLYH FHOOV RI Anabaena cylindrica. $UFK 0LFURELRO /LDQJ - 6FDSSLQR / DQG +DVHONRUQ 5 7KH patA JHQH SURGXFWZKLFKFRQWDLQVDUHJLRQVLPLODUWR&KH<RIEscherichia coli, FRQWUROVKHWHURF\VWSDWWHPIRUPDWLRQLQWKHF\DQREDFWHULXP Anabaena 3URF1DWO$FDG6FL86$ /LRWHQEHUJ6&DPSEHOO'&DVWHWV$0 +RXPDUG-DQG7DQGHDX GH 0DUVDF 1 0RGLILFDWLRQRIWKH3,,SURWHLQ LQUHVSRQVH WRFDUERQDQGQLWURJHQDYDLODELOLW\ LQ ILODPHQWRXV KHWHURF\VWRXV F\DQREDFWHULD)(060LFURELRO/HWW /LX < *ROGHQ66 .RQGR7 ,VKLXUD0DQG-RKQVRQ &+D %DFWHULDOOXFLIHUDVHDV DUHSRUWHURIFLUFDGLDQJHQHH[SUHVVLRQLQ F\DQREDFWHULD-%DFWHULRO /LX < DQG 7VLQRUHPDV 1) $Q XQXVXDO JHQH DUUDQJHPHQW IRU WKH SXWDWLYH FKURPRVRPH UHSOLFDWLRQ RULJLQ DQG FLUFDGLDQ H[SUHVVLRQ RI dnaN LQ Synechococcus 3&& *HQH /LX < 7VLQRUHPDV 1) -RKQVRQ &+ /HEHGHYD 19 *ROGHQ 66 ,VKLXUD0 DQG .RQGR7E &LUFDGLDQRUFKHVWUDWLRQRIJHQH H[SUHVVLRQLQF\DQREDFWHULD*HQHV'HY /RQHWWR 0 *ULEVNRY 0 DQG *URVV &$ 7KH IDPLO\ VHTXHQFH FRQVHUYDWLRQ DQG HYROXWLRQDU\ UHODWLRQVKLSV - %DFWHULRO 0DOLQ*DQG:DOVE\$( &KHPRWD[LV RIDF\DQREDFWHULXP RQ FRQFHQWUDWLRQ JUDGLHQWV RI FDUERQ GLR[LGH ELFDUERQDWH DQG R[\JHQ-*HQ0LFURELRO
1+ 0DQQ 0DQQ 1+ 3URWHLQ SKRVSKRU\ODWLRQ LQ F\DQREDFWHULD 0LFURELRORJ\ 0DQQ 1 &DUU 1* DQG 0LGJOH\ -(0 51$ V\QWKHVLV DQG DFFXPXODWLRQ RIJXDQLQH QXFOHRWLGHV GXULQJ JURZWK VKLIW GRZQ LQ WKH EOXH JUHHQ DOJD Anacystis nidulans. %LRFKLP %LRSK\V $FWD 0DQQ1+5LSSND5DQG+HUGPDQ0 UHJXODWLRQRISURWHLQ SKRVSKRU\ODWLRQ LQ WKH F\DQREDFWHULXP Anabaena VWUDLQ 3&& -*HQ0LFURELRO 0DUFXV )0REHUO\/DQG/DWVKDZ 63 &RPSDUDWLYH DPLQR DFLG VHTXHQFH RIIUXFWRVH ELVSKRVSKDWDVHV LGHQWLILFDWLRQ RI DUHJLRQ XQLTXH WR WKH OLJKWUHJXODWHG FKORURSODVW HQ]\PH 3URF 1DWO$FDG6FL86$ 0DUVGHQ :-1 DQG &RGG *$ 3XULILFDWLRQ DQG PROHFXODU DQG FDWDO\WLF SURSHUWLHV RI SKRVSKRULEXORNLQDVH IURP WKH F\DQREDFWHULXP Chlorogloeopsis fritschii. - *HQ 0LFURELRO 0F&OHDU\:5 6WRFN-% DQG1LQID$- ,VDFHW\OSKRVSKDWH DJOREDOVLJQDOLQEscherichia coli? -%DFWHULRO 0F&ROO 60 DQG (YDQV (+ 7KH ORFDWLRQ RID SXWDWLYH () KDQG ORRS &D ELQGLQJ VLWH LQ WKH SKRWRV\VWHP R[\JHQ HYROYLQJFRPSOH[ $EVWUDFWV RIWKH)LUVW(XURSHDQ:RUNVKRSRQ WKH0ROHFXODU%LRORJ\RI&\DQREDFWHULD 0L +/ (QGR 7 6FKUHLEHU 8 2JDZD 7 DQG $VDGD . (OHFWURQ GRQDWLRQ IURP F\FOLF DQG UHVSLUDWRU\ HOHFWURQ IORZV WR WKH SKRWRV\QWKHWLF LQWHUV\VWHP FKDLQ LV PHGLDWHG E\ S\ULGLQH QXFOHRWLGH GHK\GURJHQDVH LQ WKH F\DQREDFWHULXP Synechocystis 3&& 3ODQW&HOO3K\VLRO 0L + (QGR 7 6FKUHLEHU 8 2JDZD 7 DQG $VDGD . 1$'3 +GHK\GURJHQDVHGHSHQGHQWF\FOLFHOHFWURQIORZDURXQG SKRWRV\VWHP,LQ WKHF\DQREDFWHULXP Synechocystis 3&& DVWXG\RIGDUNVWDUYHGFHOOVDQGVSKHURSODVWV 3ODQW&HOO3K\VLRO 0L + (QGR 7 2JDZD 7 DQG $VDGD . 7K\ODNRLG 1$'3+VSHFLILF S\ULGLQHQXFOHRWLGH PHPEUDQHERXQG GHK\GURJHQDVHFRPSOH[PHGLDWHVF\FOLFHOHFWURQWUDQVSRUWLQWKH F\DQREDFWHULXP Synechocystis VS 3&& 3ODQW&HOO 3K\VLRO 0LOOHU $* (VSLH *6 DQG &DQYLQ '7 $FWLYH WUDQVSRUW RI &2 E\ WKH F\DQREDFWHULXP 6\QHFKRFRFFXV 87(; 3ODQW 3K\VLRO 0LWVXL $ .XPD]DZD 6 7DNDKDVKL $ ,NHPRWR + DQG $UDL 7 6WUDWHJ\ E\ ZKLFK QLWURJHQIL[LQJ XQLFHOOXODU F\DQREDFWHULDJURZSKRWRDXWRWURSKLFDOO\ 1DWXUH 0LWVXL $ 6XGD 6 DQG +DQDJDWD 1 &HOO F\FOH HYHQWV DW GLIIHUHQW WHPSHUDWXUHV LQ DHURELF QLWURJHQIL[LQJ PDULQH XQLFHOOXODU F\DQREDFWHULXP Synechococcus VS 0LDPL %* -0DU%LRWHFKQRO 0L\DFKL 6 %XUJHU - .RW]DEDVLV . 7KLHOPDQQ - DQG 6HQJHU + 3KRWRV\QWKHWLF FKDUDFWHULVWLFV RI VWUDLQV RI F\DQREDFWHULD JURZQ XQGHU ORZ&2 RU KLJK&2 FRQGLWLRQV =HLWVFKULIWIXU1DWXUIRUVFKXQJ 0RUJDQ'* %DXPJDUWQHU-:DQG+D]HOEDXHU*/ 3URWHLQV DQWLJHQLFDOO\UHODWHG WR PHWK\ODFFHSWLQJFKHPRWD[LV SURWHLQV RI Escherichia coli GHWHFWHG LQ DZLGHUDQJHRIEDFWHULDO VSHFLHV - %DFWHULRO 0XOOHU(*'DQG%XFKDQDQ%% 7KLRUHGR[LQLVHVVHQWLDOIRU SKRWRV\QWKHWLFJURZWK -%LRO&KHP 0XWLR]'RUDGR-,QRX\H6DQG,QRX\H0 $JHQHHQFRGLQJ D SURWHLQ VHULQHWKUHRQLQH NLQDVH LV UHTXLUHG IRU QRUPDO GHYHORSPHQWRIM . xanthus, DJUDPQHJDWLYHEDFWHULXP&HOO 1DJD\D0 $LKD+DQG0L]XQR7 7KHsphR SURGXFW D WZRFRPSRQHQW V\VWHP UHVSRQVH UHJXODWRU SURWHLQ UHJXODWHV
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW SKRVSKDWHDVVLPLODWLRQLQ Synechococcus 3&&E\ELQGLQJ WRWZRVLWHV XSVWUHDPIURP WKHphoA SURPRWHU -%DFWHULRO 1DNDPD < .DQHNR 7 +LURVDZD 0 0L\DMLPD 1 DQG 7DEDWD 6 &\DQREDVH D ::: GDWDEDVH FRQWDLQLQJ WKH FRPSOHWH QXFOHRWLGHVHTXHQFH RIWKH JHQRPHRISynechocystis 3&& 1XFOHLF$FLGV5HV 1HZPDQ - .DUDND\D + 6FDQODQ '- DQG 0DQQ 1+ $ FRPSDULVRQ RI JHQH RUJDQL]DWLRQ LQ WKH ]ZI UHJLRQ RI WKH JHQRPHVRIWKHF\DQREDFWHULDSynechococcus VS 3&& DQG Anabaena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hormidium laminosum. 3K\VLRO3ODQW 2JDZD 7 $ JHQH KRPRORJRXV WR WKH VXEXQLW JHQH RI 1$'+ GHK\GURJHQDVHLV HVVHQWLDO WR LQRUJDQLF FDUERQ WUDQVSRUW RI Synechocystis 3&& 3URF 1DWO$FDG6FL 86$ 2JDZD 7 D 1$'+ GHK\GURJHQDVH D FRPSRQHQW RI 3KRWRV\VWHP, F\FOLF HOHFWURQ WUDQVSRUW IORZ GULYLQJ LQRUJDQLF FDUERQWUDQVSRUW3KRWRV\Q5HV 2JDZD 7 E ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WKH ictA/indhL JHQHSURGXFW HVVHQWLDO WR LQRUJDQLF FDUERQ WUDQVSRUW RI Synechocystis 3&& 3ODQW3K\VLRO 2JDZD 7 0L\DQR $ DQG ,QRX\H < 3KRWRV\VWHP ,GULYHQ LQRUJDQLF FDUERQ WUDQVSRUW LQ WKH F\DQREDFWHULXP Anacystis nidulans. %LRFKLP%LRSK\V$FWD 2KPRUL 0 F$03 LQ Anabaena cylindrica: UDSLG FKDQJHV LQ FHOOXODU OHYHOV LQ UHVSRQVH WR FKDQJHV LQ H[WUDFHOOXODU HQYLURQPHQWV 3ODQW&HOO3K\VLRO 2KPRQ . +LURVH 0 DQG 2KPRQ 0 $Q LQFUHDVH LQ WKH LQWUDFHOOXODU FRQFHQWUDWLRQ RI F$03 WULJJHUV IRUPDWLRQ RI DQ DOJDO PDWE\ WKH F\DQREDFWHULXP Spirulina platensis. 3ODQW &HOO 3K\VLRO 2KPRUL 0 .DWD\DPD 0 .DVDKDUD 0 DQG 7HUDXFKL . $GHQ\ODWH F\FODVH JHQHV UHVSRQVLEOH IRU HQYLURQPHQWDO VLJQDOV $EVWUDFWVRIWKH9,&\DQREDFWHULDO:RUNVKRS$ 2KPRUL 0 2KPRUL . DQG +DVXQXPD. 5DSLG FKDQJH LQ F\FOLF$03FRQFHQWUDWLRQWULJJHUHGE\DOLJKWRIIRUOLJKWRQ VLJQDOLQAnabaena cylindrica. $UFK0LFURELRO 2QHN /$ /HD 3- DQG 6PLWK 5- ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI D FDOPRGXOLQOLNH SURWHLQ IURP WKH F\DQREDFWHULXP Nostoc VS 3&& $UFK 0LFURELRO 2QHN/$DQG6PLWK 5- &DOPRGXOLQ DQG FDOFLXPPHGLDWHG UHJXODWLRQLQSURNDU\RWHV-*HQ0LFURELRO 3DFROG 0( 6WHYHQV )- /L ' DQG $QGHUVRQ /( 7KH 1$'3OLQNHG JO\FHUDOGHK\GHSKRVSKDWH GHK\GURJHQDVH RI Anabaena variabilis DQG Synechocystis 3&& ZKLFK ODFN RQH RIWKH F\VWHLQHV IRXQG LQ WKH KLJKHU SODQW HQ]\PH DUH QRW UHGXFWLYHO\DFWLYDWHG3KRWRV\Q5HV
3DUNLQVRQ -6 DQG .RIRLG (& &RPPXQLFDWLRQ PRGXOHV LQ EDFWHULDOVLJQDOOLQJSURWHLQV$QQ5HY*HQHW 3HOUR\ 5$ .LUN 05 DQG %DVVKDP -$ 3KRWRV\VWHP ,, UHJXODWLRQ RI PDFURPROHFXOH V\QWKHVLV LQ WKH EOXHJUHHQ DOJD Aphanocapsa -%DFWHULRO 3HVFKHN *$ ,QWHUDFWLRQ RISKRWRV\QWKHVLV DQG UHVSLUDWLRQ %LRFKHP6RF7UDQV 3HWWHUVVRQ $ DQG %HUJPDQ% &DOPRGXOLQ LQ KHWHURF\VWRXV F\DQREDFWHULDELRFKHPLFDODQGLPPXQRORJLFDOHYLGHQFH )(06 0LFURELRO/HWW 3LWWD 73 DQG %HUJ +& 6HOIHOHFWURSKRUHVLV LV QRW WKH PHFKDQLVP IRU PRWLOLW\ LQ VZLPPLQJF\DQREDFWHULD - %DFWHULRO 3LWWD73 6KHUZRRG(( .REHO$0 DQG %HUJ+& &DOFLXP LV UHTXLUHG IRUVZLPPLQJE\ WKHQRQIODJHOODWHGF\DQREDFWHULXP Synechococcus VWUDLQ:+-%DFWHULRO 3RUDQNLHZLF]- DQG&ODUNH$. ,QGXFWLRQRIWKHKHDW VKRFN SURWHLQ &OS% DIIHFWV FROG DFFOLPDWLRQ LQ WKH F\DQREDFWHULXP Synechococcus 3&& -%DFWHULRO 3RUDQNLHZLF] - 6FKHOLQ - DQG &ODUNH $. 7KH $73 GHSHQGHQW&OSSURWHDVHLVHVVHQWLDOIRUDFFOLPDWLRQWR89% DQG ORZ WHPSHUDWXUH LQ WKH F\DQREDFWHULXP Synechococcus. 0ROHF 0LFURELRO 3RWWV06XQ+0RFNDLWLV..HQQHOO\3-5HHG'DQG7RQNV1. $ SURWHLQW\URVLQHVHULQH SKRVSKDWDVH HQFRGHG E\ WKH JHQRPH RI WKH F\DQREDFWHULXP Nostoc commune 87(; - %LRO&KHP 4XDLO 3+ 3KRWRVHQVRU\ SHUFHSWLRQ DQG VLJQDO WUDQVGXFWLRQ LQSODQWV&XUU2SLQLRQ*HQHW'HYHO 5H\HV -& &UHVSR -/ *DUFLD'RQLLQJXH] 0 DQG )ORUHQFLR )- (OHFWURQ WUDQVSRUW FRQWUROV JOXWDPLQH V\QWKHWDVH DFWLYLW\ LQ WKH IDFXOWDWLYH KHWHURWURSKLF F\DQREDFWHULXP Synechocystis VS3&&3ODQW3K\VLRO 5LFKDUGVRQ // DQG &DVWHQKRO] 5: 'LHO YHUWLFDO PRYHPHQWV RIWKH F\DQREDFWHULXP Oscillatoria terebriformis LQ DVXOILGHULFKKRWVSULQJPLFURELDOPDW $SSO(QYLURQ 0LFURELRO 5LFKDUGVRQ //DQG &DVWHQKRO] 5: &KHPRNLQHWLF PRWLOLW\ UHVSRQVHV RI WKH F\DQREDFWHULXP Oscillatoria terebriformis. $SSO(QYLURQ 0LFURELRO 5RVHQ %3 %DFWHULDO FDOFLXP WUDQVSRUW %LRFKLP %LRSK\V $FWD 6DOODO $.- 7KH SUHVHQFH RI JOXWDPLQH V\QWKHWDVH LQ WKH WK\ODNRLGV RIQLWURJHQIL[LQJ F\DQREDFWHULXPNostoc muscorum. -%DVLF0LFURELRO 6FDQODQ '- %RXUQH - DQG 0DQQ 1+ $ SXWDWLYH WUDQVFULSWLRQDO DFWLYDWRU RI WKH &US)QU IDPLO\ IURP WKH PDULQH F\DQREDFWHULXP Synechococcus VS :+ - $SSO 3K\FRO 6FDQODQ'-DQG0DQQ1+ 6HTXHQFHDQDO\VLV RIDJHQRPLF UHJLRQ $FFHVVLRQ 1R 8 IURP WKH F\DQREDFWHULXP Synechococcus VS 3&&HQFRGLQJD1DU/KRPRORJXH DQGD WZRFRPSRQHQWV\VWHPSDLU3ODQW3K\VLRO3*5 6FKDHIIHU ) DQG 6WDQLHU 5< *OXFRVHSKRVSKDWH GHK\GURJHQDVHRIAnabaena VS $UFK0LFURELRO 6FKLUPHU (& *ORYHU -5 6LQJHU 0$ DQG /LQGTXLVW 6 +63&OS SURWHLQV D FRPPRQ PHFKDQLVP H[SODLQV GLYHUVH IXQFWLRQV7UHQGV%LRO6FLHQFHV 6FKOXFKWHU:0 DQG%U\DQW'$ 0ROHFXODUFKDUDFWHUL]DWLRQ RI IHUUHGR[LQ1$'3 R[LGRUHGXFWDVH LQ F\DQREDFWHULD FORQLQJ DQG VHTXHQFH RIWKHpetH JHQH RISynechococcus VS 3&& DQGVWXGLHVRQWKHJHQHSURGXFW%LRFKHPLVWU\
6FKPHWWHUHU * &\DQREDFWHULDO UHVSLUDWLRQ ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHUDFDGHPLF3XEOLVKHUV'RUGUHFKW7KH1HWKHUODQGV 6FKQHLGHU*-7XUQHU1(5LFKDXG& %RUEHO\*DQG+DVHONRUQ5 3XULILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI 51$ SRO\PHUDVH IURP WKH F\DQREDFWHULXP Anabaena - %LRO &KHP 6FKQHLGHU3RHWVFK +$ %UDXQ % 0DU[ 6 DQG 6FKDXPEXUJ $ 3K\WRFKURPHVDQGEDFWHULDOVHQVRUSURWHLQVDUHUHODWHGE\ VWUXFWXUDODQGIXQFWLRQDOKRPRORJLHV)(%6/HWW 6FKZDU] 5 DQG *URVVPDQ $5 $ UHVSRQVH UHJXODWRU RI F\DQREDFWHULD LQWHJUDWHV GLYHUVH HQYLURQPHQWDO VLJQDOV DQG LV FULWLFDO IRU VXUYLYDO XQGHU H[WUHPH FRQGLWLRQV 3URF 1DWO $FDG 6FL 86$ 6KDUURFN5$DQG4XDLO3+ 1RYHOSK\WRFKURPHVHTXHQFHV LQ Arabidopsis thaliana: VWUXFWXUH HYROXWLRQ DQG GLIIHUHQWLDO H[SUHVVLRQ RI D SODQW UHJXODWRU\ SKRWRUHFHSWRU IDPLO\ *HQHV 'HY 6KDZ'-5LFH':DQG*XHVW-5 +RPRORJ\EHWZHHQ&$3 DQG )1 DUHJXODWRURIDQDHURELFUHVSLUDWLRQLQEscherichia coli. -0RO%LRO 6KL / DQG &DUPLFKDHO :: SSOF\DQR D SURWHLQ VHULQHWKUHRQLQH SKRVSKDWDVH JHQH IURP WKH F\DQREDFWHULXP Microcystis aeruginosa. $UFK0LFURELRO 6KL : DQG =XVPDQ '5 7KHIU]VLJQDO WUDQVGXFWLRQ V\VWHP FRQWUROV PXOWLFHOOXODU EHKDYLRXU LQ Myxococcus xanthus. ,Q +RFK -$ DQG 6LOKDY\ 7- HGV 7ZR&RPSRQHQW 6LJQDO 7UDQVGXFWLRQSS 6LOPDQ 1- &DUU 1* DQG 0DQQ 1+ $'3ULERV\ODWLRQ RI JOXWDPLQHV\QWKHWDVHLQ WKHF\DQREDFWHULXP Synechocystis 3&& -%DFWHULRO 6LPSVRQ**&ODUN*DQG%URZQJWS ,VRODWLRQRIDPDL]H F'1$ HQFRGLQJ D SURWHLQ ZLWK H[WHQVLYH VLPLODULW\ WR DQ LQKLELWRURISURWHLQ NLQDVH & DQG D F\DQREDFWHULDO RSHQ UHDGLQJ IUDPH%LRFKLP%LRSK\V$FWD 6PLWK$- 0RGHVRIF\DQREDFWHULDOFDUERQPHWDEROLVP ,Q &DUU1*DQG :KLWWRQ %$ HGV 7KH %LRORJ\ RI&\DQREDFWHULD SS%ODFNZHOO6FLHQWLILF3XEOLFDWLRQV2[IRUG 6PLWK * DQG 2ZQE\ -' &\FOLF $03 LQWHUIHUHV ZLWK SDWWHUQ IRUPDWLRQ LQ WKH F\DQREDFWHULXP Anabaena variabilis. )(060LFURELRO/HWW 6PLWK + 3K\VLRORJLFDO DQG HFRORJLFDO IXQFWLRQ ZLWKLQ WKH SK\WRFKURPHIDPLO\ $QQ5HY3ODQW3K\VLRO3ODQW0RO %LRO 6PLWK 5- &DOFLXPPHGLDWHG UHJXODWLRQ LQ WKH F\DQREDFWHULD",Q 5RJHUV /, DQG*DOORQ-5HGV %LRFKHPLVWU\ RI $OJDH DQG &\DQREDFWHULD SS &ODUHQGRQ 3UHVV 2[IRUG 6PLWK 5- DQG &DUU 1* 7KH UHJXODWLRQ RI VWDEOH 51$ V\QWKHVLV LQ WKH EOXHJUHHQ DOJD Anacystis nidulans: HIIHFW RI OHXFLQH GHSULYDWLRQ DQG PHWK\OWU\SWRSKDQ DGGLWLRQ - *HQ 0LFURELRO 6PLWK 5- +REVRQ 6 DQG (OOLV ,5 (YLGHQFH IRU FDOFLXP PHGLDWHG UHJXODWLRQ RI KHWHURF\VW IUHTXHQF\ DQG QLWURJHQDVH DFWLYLW\LQNostoc 1HZ3K\WRO 6REF]\N$%HO\$7DQGHDXGH0DUVDF1DQG+RXPDUG- $ SKRVSKRU\ODWHG '1$ELQGLQJ SURWHLQ LV VSHFLILF IRU WKH UHG OLJKW VLJQDO GXULQJ FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ 0ROHF 0LFURELRO 6WDO/,DQG.UXPEHLQ:( 1LWURJHQDVHDFWLYLW\LQWKHQRQ KHWHURF\VWRXV F\DQREDFWHULXP Oscillatoria VS JURZQ XQGHU DOWHUQDWLQJOLJKWGDUNF\FOHV$UFK0LFURELRO
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ynechocystis 3&& UHQGHUV WKH PXWDQWOLJKWVHQVLWLYH-%LRO&KHP 6XPPHUV 0/ 0HHNV -& &KX 6 DQG :ROI 5( -U D 1XFOHRWLGH VHTXHQFH RI DQ RSHURQ LQ Nostoc $7&& HQFRGLQJ IRXU JHQHV RI WKH R[LGDWLYH SHQWRVH SKRVSKDWH F\FOH 3ODQW3K\VLRO 6XPPHUV 0/ :DOOLV -* &DPSEHOO (/ DQG 0HHNV -& E *HQHWLF HYLGHQFH RI D PDMRU UROH IRU JOXFRVHSKRVSKDWH GHK\GURJHQDVH LQ QLWURJHQ IL[DWLRQ DQG GDUN JURZWK RI WKH F\DQREDFWHULXP Nostoc $7&& - %DFWHULRO 6XQGDUDP 6 .DUDND\D + 6FDQODQ '- DQG 0DQQ 1+ 0XOWLSOH ROLJRPHULF IRUPV RI JOXFRVHSKRVSKDWH GHK\GURJHQDVH LQ F\DQREDFWHULD DQG WKH UROH RI 2SF$ LQ WKH DVVHPEO\SURFHVV0LFURELRORJ\ 6XUiQ\L*.RUF]$3iOIL=DQG%RUEHO\* (IIHFWVRIOLJKW GHSULYDWLRQ RQ 51$ V\QWKHVLV DFFXPXODWLRQ RI JXDQRVLQH ¶¶ GLSKRVSKDWH DQG SURWHLQ V\QWKHVLV LQ KHDWVKRFNHG Synechococcus 3&& D F\DQREDFWHULXP - %DFWHULRO 6X]XNL,+RULH1 6XJL\DPD7DQG2PDWD7 ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WZR QLWURJHQUHJXODWHG JHQHV RI WKH F\DQREDFWHULXP Synechococcus 3&& UHTXLUHG IRU PD[LPXP HIILFLHQF\ RI QLWURJHQ DVVLPLODWLRQ - %DFWHULRO 6X]XNL , 6XJL\DUQD7DQG 2PDWD7 UHJXODWLRQE\ F\DQDWH RIWKHJHQHV LQYROYHGLQFDUERQ DQG QLWURJHQ DVVLPLODWLRQLQWKH F\DQREDFWHULXP Synechococcus 3&& - %DFWHULRO 6ZHHQH\%0 DQG%RUJHVH0% $ FLUFDGLDQ UK\WKPLQ FHOO GLYLVLRQ LQ D SURNDU\RWH WKH F\DQREDFWHULXP Synechococcus :+-3K\FRO 7DQ ; 9DUXJKHVH 0 DQG :LGJHU :5 $ OLJKWUHSUHVVHG WUDQVFULSW IRXQG LQ Synechococcus 3&& LV VLPLODU WR D FKORURSODVWVSHFLILF VPDOO VXEXQLW ULERVRPDO SURWHLQ DQG WR D WUDQVFULSWLRQ PRGXODWRU SURWHLQ DVVRFLDWHG ZLWK - %LRO &KHP 7DQDND . 0DVXGD 6 DQG 7DNDKDVKL + D 7KH FRPSOHWH QXFOHRWLGH VHTXHQFH RIWKH JHQH (rpoDI) HQFRGLQJ WKH SULQFLSDO VLJPD IDFWRU RIWKH 51$ SRO\PHUDVH IURP WKH F\DQREDFWHULXP Synechococcus 3&&%LRFKLP%LRSK\V$FWD 7DQDND . 0DVXGD 6 DQG 7DNDKDVKL + E 0XOWLSOH rpoDUHODWHG JHQHV RI F\DQREDFWHULD %LRVFL %LRWHFK %LRFKHP 7DQGHDX GH 0DUVDF 1 DQG +RXPDUG - $GDSWDWLRQ RI F\DQREDFWHULD WR HQYLURQPHQWDO VWLPXOL QHZ VWHSV WRZDUGV PROHFXODUPHFKDQLVPV)(060LFURELRO5HY 7HUDXFKL . DQG 2KPRUL 0 $Q DGHQ\ODWH F\FODVH &\DO PHGLDWHV WKH VLJQDO RI EOXH OLJKW LQ WKH F\DQREDFWHULXP Synechocystis VS3&& $EVWUDFWVRIWKH9,&\DQREDFWHULDO :RUNVKRS6
&KDSWHU 'HWHFWLQJWKH (QYLURQPHQW 7KPPOHU)$OJDUUD3DQG)RER*0 6HTXHQFHVLPLODULWLHV RI SK\WRFKURPHV WR SURWHLQ NLQDVHV LPSOLFDWLRQV IRU WKH VWUXFWXUH IXQFWLRQ DQG HYROXWLRQ RI WKH SK\WRFKURPH JHQH IDPLO\ )(%6 /HWW 7RUUHFLOOD, /HJDQHV)%RQLOOD,DQG)HPDQGH]3LxDV 8VH RIUHFRPELQDQW DHTXRULQ WR PRQLWRU&D WUDQVLHQWV LQAnabaena VS 3&& LQ UHVSRQVH WRHQYLURQPHQWDO FKDQJHV $EVWUDFWV RIWKH9,&\DQREDFWHULDO:RUNVKRS$ 7UDPRQWLQL /6 7KH UROH RI &D LQ SKRWRV\VWHP RI Anacystis nidulans. 3K'WKHVLV/DQFDVKLUH3RO\WHFKQLF8. 7VDL -: 3URWHLQ SKRVSKRU\ODWLRQ LQ F\DQREDFWHULDO OLJKW KDUYHVWLQJFRPSOH[HV3K'WKHVLV8QLYHUVLW\RI:DUZLFN 7VLQRUHPDV1) &DVWHWV$0+DUULVRQ0$ $OOHQ-)DQG7DQGHDX GH 0DUVDF 1 3KRWRV\QWKHWLF HOHFWURQ WUDQVSRUW FRQWUROV QLWURJHQ DVVLPLODWLRQ LQ F\DQREDFWHULD E\ PHDQV RI SRVWWUDQVODWLRQDO PRGLILFDWLRQ RI the glnB JHQH SURGXFW 3URF 1DWO$FDG6FL86$ 7VLQRUHPDV 1) ,VKLXUD 0 .RQGR 7 $QGHUUVRQ &5 7DQDND . 7DNDKDVKL+-RKQVRQ&+DQG*ROGHQ66 $VLJPDIDFWRU WKDW PRGLILHV WKH FLUFDGLDQ H[SUHVVLRQ RI D VXEVHW RI JHQHV LQ F\DQREDFWHULD(0%2- 8GYDUG\ - %RUEHO\ * -XKDV] $ DQG )DUNDV */ 7KLRUHGR[LQV DQG WKH UHGR[ PRGXODWLRQ RIJOXFRVHSKRVSKDWH GHK\GURJHQDVH LQ Anabaena 3&& YHJHWDWLYH FHOOV DQG KHWHURF\VWV-%DFWHULRO 8GYDUG\ - *RGHK 0+ DQG )DUNDV */ 5HJXODWRU\ SURSHUWLHV RI D IUXFWRVH ELVSKRVSKDWDVH IURP WKH F\DQREDFWHULXP Anacystis nidulans. -%DFWHULRO 9HJD3DODV 0$ )ORUHV ( DQG +HUUHUR $ 1WF$ D JOREDO QLWURJHQ UHJXODWRU IURP WKH F\DQREDFWHULXP Synechococcus WKDW EHORQJV WR WKH &US IDPLO\ RI EDFWHULDO UHJXODWRUV 0ROHF 0LFURELRO :DOVE\$( *DVYHVLFOHV0LFURELRO5HY :DUQHU .0 DQG %XOOHMDKQ *6 /LJKWGHSHQGHQW W\URVLQH SKRVSKRU\ODWLRQ LQ WKH F\DQREDFWHULXP Prochlorothrix hollandica. 3ODQW3K\VLRO :DWHUEXU\-%:LOOH\-0)UDQNV'*9DORV):DQG:DWVRQ 6: $ F\DQREDFWHULXP FDSDEOH RI VZLPPLQJ PRWLOLW\ 6FLHQFH :DWVRQ*0) 6FDQODQ '- DQG0DQQ 1+ &KDUDFWHUL]DWLRQ RI WKH JHQHV HQFRGLQJ D SKRVSKDWHUHJXODWHG WZRFRPSRQHQW VHQVRU\V\VWHPLQWKHPDULQHF\DQREDFWHULXPSynechococcus VS :+)(060LFURELRO/HWW :DZU]\QRZ$%DQHFNL% DQG=\OLF]0 7KH&OS$73DVHV GHILQH DQRYHO FODVV RIPROHFXODU FKDSHURQHV 0ROHF 0LFURELRO :HEHU ,7 DQG 6WHLW] 7$ 6WUXFWXUH RI D FRPSOH[ RI FDWDEROLWHJHQHDFWLYDWRUSURWHLQDQGF\FOLF$03UHILQHGDW c UHVROXWLRQ-0RO%LRO :LOGH$&KXULQ<6FKXEHUW+DQG%RPHU7 'LVUXSWLRQRI D Synechocystis VS 3&& JHQH ZLWK SDUWLDO VLPLODULW\ WR SK\WRFKURPHJHQHV DOWHUVJURZWKXQGHUFKDQJLQJOLJKWTXDOLWLHV )(%6/HWW :LOOH\ -0 DQG :DWHUEXU\ -% &KHPRWD[LV WRZDUGV QLWURJHQRXV FRPSRXQGV E\ VZLPPLQJ VWUDLQV RI PDULQH Synechococcus VSS $SSO(QY0LFURELRO :LOOH\ -0 :DWHUEXU\ -% DQG *UHHQEHUJ (3 6RGLXP FRXSOHG PRWLOLW\ LQ D VZLPPLQJ F\DQREDFWHULXP - %DFWHULRO :RON&3(UQVW$DQG(OKDL- +HWHURF\VWGHYHORSPHQWDQG PHWDEROLVP ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU DFDGHPLF 3XEOLVKHUV 'RUGUHFKW7KH1HWKHUODQGV
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hitton and M. Potts (eds), The Ecology of Cyanobacteria, Kluwer Academic Publishers . Printed in the Netherlands
' %KD\D 5 6FKZDU]DQG$5*URVVPDQ
,9
*HQH 5HJXODWLRQ 'XULQJ 3KRVSKRUXV /LPLWDWLRQ 3RO\SKRVSKDWHV ' 6XOIXU /LPLWDWLRQ 6XOIDWH 8SWDNH *HQH 5HJXODWLRQ 'XULQJ 6XOIXU /LPLWDWLRQ &\V5 &KDQJHV LQ WKH 3KRWRV\QWKHWLF $SSDUDWXV D 3KRWRV\QWKHWLF $FWLYLW\ DQG *HQH 5HJXODWLRQ E 3K\FRELOLVRPH 'HJUDGDWLRQ &RQFOXGLQJ 5HPDUNV
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
,QWURGXFWLRQ 7KHUH LV D ZHDOWK RILQIRUPDWLRQ RQ WKH ELRFKHPLFDO DQG SK\VLRORJLFDO UHVSRQVHV RI F\DQREDFWHULD WR HQYLURQPHQWDOVWUHVV 7KHHPSKDVLVLQWKLVFKDSWHULV RQWKH0ROHFXODUUHVSRQVHVWRWZRNH\HQYLURQPHQWDO SDUDPHWHUV OLJKW DQG QXWULHQWV 0RVW F\DQREDFWHULD DUH REOLJDWH SKRWRDXWRWURSKV PDNLQJ LW FULWLFDO IRU WKHPWR VHQVH DQGUHVSRQGWRERWKOLJKWLQWHQVLW\ DQG TXDOLW\ 9DQ /LHUH DQG :DOVE\ 6LQFH OLJKW SULPDULO\ DIIHFWVSKRWRV\QWKHWLF DFWLYLW\ PDQ\ RIWKH $3
3&
Abbreviations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b6f FRPSOH[ WKH $73 V\QWKDVH DQG WKH SK\FRELOLVRPH 3%6 7KH 3%6 LV WKH PDMRU OLJKWKDUYHVWLQJ FRPSOH[ LQ F\DQREDFWHULD ZKLFK HIILFLHQWO\ JDWKHUV OLJKW HQHUJ\ DQG GLUHFWV PRVW RI WKDW HQHUJ\ WR WKH UHDFWLRQ FHQWHUV RI36,, 7KH DQWHQQDH SLJPHQWV RI 36, FRQVLVW PRVWO\ RI FKORURSK\OO FKO PROHFXOHV
:KHQ WKH H[FLWDWLRQ HQHUJ\ UHDFKHV WKH UHDFWLRQ FHQWHUV LW LV FRQYHUWHG LQWR FKHPLFDO HQHUJ\ LQ WKH IRUP RIDFKDUJH VHSDUDWLRQ ,QWKHUHDFWLRQFHQWHURI 36,, ZDWHU PROHFXOHV DUH R[LGL]HG DQG WKH HOHFWURQV DUH WUDQVIHUUHG WR WKH SODVWRTXLQRQH 34 SRRO (OHFWURQVIURPWKH34SRRODUHSDVVHG VHTXHQWLDOO\WR WKH F\WRFKURPH b6f FRPSOH[ SODVWRF\DQLQ RU F\WRFKURPH F GLIIXVLEOH HOHFWURQ FDUULHUV DQG ILQDOO\WR36, 7KH HOHFWURQV IURP 36, DIWHU IXUWKHU HQHUJL]DWLRQ DUH WUDQVIHUUHG WR IHUUHGR[LQ DQG ILQDOO\ WR 1$'3+ 3URWRQV DFFXPXODWH LQ WKH WK\ODNRLG OXPHQDV DFRQVHTXHQFH RIWKH VSOLWWLQJRIZDWHUSOXV HOHFWURQ WUDQVIHU WKURXJK WKH F\WRFKURPH b6fFRPSOH[ WKH SURWRQ JUDGLHQW JHQHUDWHG DFURVV WKH WK\ODNRLG PHPEUDQHV LV FRXSOHG WR $73 SURGXFWLRQ YLD WKH $73 V\QWKDVH %RWK WKH UHGXFWDQW 1$'3+ DQG HQHUJ\ $73 JHQHUDWHG E\ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUWDUHFHQWUDOWRWKHPHWDEROLVPRIWKHFHOO
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µVWDWH WUDQVLWLRQV¶ DQG GR QRW UHTXLUH SURWHLQ V\QWKHVLV +RZHYHU F\DQREDFWHULD DUH DOVR FDSDEOH RIDOWHULQJ L WKH WRWDO QXPEHU RISKRWRV\VWHPV LL WKH UDWLR RI 36, WR 36,, DQG LLL 3%6 VWUXFWXUH LQ UHVSRQVH WR SURORQJHG FKDQJHV LQ WKH OLJKW
HQYLURQPHQW 7KHVH ORQJWHUPDFFOLPDWLRQUHVSRQVHV UHTXLUH WKH LQGXFWLRQ RI JHQHV DQG WKH de novo V\QWKHVLV RI SURWHLQV )XUWKHUPRUH ZKHQ WKH OLJKW LQWHQVLW\LVYHU\KLJKLWFDQFDXVH µSKRWRLQKLELWLRQ¶ RU GDPDJHWRWKHUHDFWLRQFHQWHUVRI36,,WKDWUHVXOWVLQD GHFOLQH LQ WKH TXDQWXP \LHOG RI 2 HYROXWLRQ &\DQREDFWHULDFDQ PRGLI\ WKH SURWHLQFRPSRVLWLRQ RI 36,,LQKLJKOLJKW ZKLFKPD\PDNHWKH36,,UHDFWLRQ FHQWHUV OHVVVXVFHSWLEOHWRSKRWRLQKLELWLRQ 7KHZD\V LQZKLFKWKHLUSKRWRV\QWKHWLF DSSDUDWXV FDQ DGMXVWWR WKHFKDQJLQJOLJKWHQYLURQPHQWDUHGLVFXVVHGEHORZ
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µVWDWHWUDQVLWLRQV¶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b6fEHFRPHUHGXFHG7KH UHGXFWLRQ RI WKH 34 SRRO DSSHDUV WR VWLPXODWH WKH
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µPRELOH DQWHQQD¶ PRGHO K\SRWKHVL]HVWKDWWKH3%6 GLVFRQQHFWV IURP 36,, DQG EHFRPHV SK\VLFDOO\ DVVRFLDWHG ZLWK 36, ,Q WKH µVSLOORYHU¶ PRGHO WKH 3%6 GR QRW SK\VLFDOO\ PRYH KRZHYHU H[FHVV H[FLWDWLRQ HQHUJ\ DFFXPXODWHG LQ 36,,FDQVSLOORYHULQDVRPHZKDWLOOGHILQHGPDQQHU LQWR WKH UHDFWLRQ FHQWHUV RI36, ,Q WKH WKLUG PRGHO WHUPHG WKH µGHWDFKPHQW¶ PRGHO H[FHVV H[FLWDWLRQ RI 36,, UHVXOWV LQ 3%6 GHWDFKPHQW KRZHYHU WKH GHWDFKHG 3%6 GRHV QRW QHFHVVDULO\ UHDVVRFLDWH ZLWK DQG GRQDWH HQHUJ\ WR 36, 1RQH RI WKH PRGHOV H[SODLQV DOO RI WKH H[SHULPHQWDO GDWD DQG QHZ DSSURDFKHV ERWK ELRSK\VLFDO DQG JHQHWLF DUH EHLQJ XVHG WR HOXFLGDWH WKH PROHFXODU GHWDLOV RI VWDWH WUDQVLWLRQV 6RPH H[SHULPHQWV KDYH VXJJHVWHG D FULWLFDO UROH IRU DOORSK\FRF\DQLQ HQFRGHG E\ apcD LQ FRQWUROOLQJ WKH GLVWULEXWLRQ RI H[FLWDWLRQ HQHUJ\EHWZHHQWKHSKRWRV\VWHPV=KDRHWDO
% /RQJ7HUP $FFOLPDWLRQV +LJK/LJKWDQG3KRWRLQKLELWLRQ 7KHZD\VLQZKLFKSKRWRV\QWKHWLFRUJDQLVPVDGMXVWWR H[FHVV H[FLWDWLRQ LV RIPDMRU HFRORJLFDO DQG SRWHQWLDO FRPPHUFLDO LPSRUWDQFH ([FHVV H[FLWDWLRQ RI WKH SKRWRV\QWKHWLF UHDFWLRQ FHQWHUV FDQ OHDG WR SKRWRLQKLELWLRQ $QGHUVRQ DQG 6W\ULQJ $UR HW
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO 6WUHVV DO .LPHWDO 6DPXHOVVRQHWDO RU D GHFOLQH LQ WKH TXDQWXP \LHOG RI SKRWRV\QWKHVLV WKDW PD\ UHIOHFW GDPDJH WR UHDFWLRQ FHQWHUV RI 36,, 7KH 36,, UHDFWLRQ FHQWHUV DUH FRPSULVHG RI IRXU SURWHLQV ' DQG ' ZKLFK IRUPKHWHURGLPHUV LQWKH PHPEUDQHV F\WRFKURPH b559 DQG WKH psb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psbA) ORFDWHG RQ WKH SODVWLG JHQRPH $QGHUVVRQ DQG 6W\ULQJ 6KLSWRQ DQG %DUEHU ,Q F\DQREDFWHULD VXFK DV Synechococcus 3&& WKHUHDUHWZRGLVWLQFWIRUPVRIWKH' SURWHLQHQFRGHG E\DVPDOOpsbA PXOWLJHQHIDPLO\*ROGHQ ,Q FRQWUDVW ZKLOH WKHUH DUH WZR psbD JHQHV IRU ' LQ Synechococcus 3&& WKH SRO\SHSWLGHV HQFRGHG E\ WKHVH JHQHV DUH LGHQWLFDO *LQJULFK HW DO 7KHUHJXODWLRQ RIWKHpsbA DQG psbD JHQHV KDV EHHQ H[WHQVLYHO\ VWXGLHG LQ Synechococcus 3&& *ROGHQHWDO *ROGHQHWDO *ROGHQDQG 6WHDUQV 7KLV RUJDQLVP FRQWDLQV WKUHH psbA JHQHV GHVLJQDWHGpsbAI, psbAII, psbAIII. 7KH psbAI JHQH FRGHV IRU )RUP , RI WKH ' SRO\SHSWLGH DOVR NQRZQ DV ' ZKLOH ERWKpsbAII DQG psbAIII FRGH IRU ' )RUP ,, ' 2I WKH DPLQR DFLG GLIIHUHQFHV EHWZHHQ )RUP , DQG )RUP DUH FOXVWHUHG ZLWKLQ WKH ILUVW DPLQR WHUPLQDO UHVLGXHV RIWKHSRO\SHSWLGH 6HYHQDPLQRDFLGGLIIHUHQFHVDUH DOVRSUHVHQWLQWUDQVPHPEUDQHKHOLFHV,,DQG,,, 7KH psbA JHQHV H[KLELW FOHDU GLIIHUHQFHV LQ WKHLU H[SUHVVLRQSDWWHUQVZLWKUHVSHFWWROLJKWLQWHQVLW\ $W ORZ LUUDGLDQFHV WKHpsbAI WUDQVFULSWSUHGRPLQDWHV ,I WKHFHOOVDUHWUDQVIHUUHGWRKLJKHUOLJKWLQWHQVLWLHVWKH OHYHORIWKHpsbAI WUDQVFULSWUDSLGO\GHFOLQHVZKLOHWKH psbAII DQG psbAIII WUDQVFULSWV DFFXPXODWH %XVWRV HW DO 7KHVHFKDQJHVLQWUDQVFULSWOHYHOVDUHDOVR UHIOHFWHG DW WKH SRO\SHSWLGH OHYHO Synechococcus 3&& JURZQ DW ORZ OLJKWFRQWDLQV SULPDULO\ '
)RUP,ZKLFKLVDOPRVWHQWLUHO\UHSODFHGDIWHUDVKRUW H[SRVXUH WR KLJK OLJKW LQWHQVLWLHV ZLWK ' )RUP ,, &ODUNH.XONDUQLDQG*ROGHQ 7KH KLJK OLJKW LQWHQVLW\WULJJHUHG FKDQJH LQ WKH SRO\SHSWLGH FRPSRVLWLRQ RIWKH 36,, UHDFWLRQ FHQWHUV LQ F\DQREDFWHULD PD\ EH LPSRUWDQW IRU WKH RUJDQLVP WR FRSH ZLWK WKH SRWHQWLDOO\ GDPDJLQJ HIIHFWV RI H[FHVV LUUDGLDWLRQ 7KH ' )RUP ,, SRO\SHSWLGH SUREDEO\ KHOSV SURWHFW WKH FHOOV IURP SKRWRGDPDJH 7KH PRVW FRQYLQFLQJ GDWD FRQFHUQLQJ WKH EHQHILW RI ' )RUP ,, GXULQJ H[SRVXUH RISynechococcus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ynechococcus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ynechococcus 3&& FXOWXUHVWUDQVIHUUHGIURP WRPROSKRWRQP V WKH' )RUP,ZDVUHSODFHGE\' )RUP,,ZLWKLQ
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psbA P51$OHYHOV%XVWRVHWDO 6FKDHIHUDQG *ROGHQ $W ORZ OLJKW PRVW RI WKH psbA WUDQVFULSWZDVIURPpsbAI. 8SRQDVKLIWWRKLJKOLJKW WKHUH ZDV D UDSLG DFFXPXODWLRQ RI WKH psbAII DQG psbAIII WUDQVFULSWV DQG D FRQFRPLWDQW ORVV RI WKH psbAI PHVVDJH 7KLV DFFXPXODWLRQ RIWKHpsbAII DQG psbAIlI PHVVDJHV LV D FRQVHTXHQFH RI DFWLYDWLRQ RI WKHVH JHQHV .XONDUQL HW DO ZKLOH WKH ORVV RI WKH psbAI WUDQVFULSW UHIOHFWV ERWK D GHFUHDVHG UDWH RI WUDQVFULSWLRQDQGDQLQFUHDVHGUDWHRIP51$WXUQRYHU .XONDUQL HW DO %DVHG RQ SURPRWHU DQDO\VHV LQ ZKLFK WKH psbAII DQG psbAIII SURPRWHUV ZHUH IXVHG WR lacZ HQFRGLQJ 3JDODFWRVLGDVH VHTXHQFHV ZLWKLQ WKHXQWUDQVODWHG OHDGHUUHJLRQV RIWKH P51$V ZHUH GHWHUPLQHG WR EH UHVSRQVLEOH IRU LQFUHDVHG H[SUHVVLRQ RI WKH psbAII DQG psbAIII JHQHV LQ KLJK OLJKW /L DQG *ROGHQ 7KHVH VHTXHQFH HOHPHQWVPD\IXQFWLRQDVHQKDQFHUVWKDWELQGSURWHLQV WKDWDUHRQO\SUHVHQWLQH[WUDFWVIURPKLJKOLJKWJURZQ FHOOV/LHWDO $IWHU ORQJ H[SRVXUHV WR KLJK OLJKW WKH psbAI WUDQVFULSW EHJLQV WR LQFUHDVH ZKLOH WKH psbAII DQG psbAIII WUDQVFULSWV UHPDLQUHODWLYHO\KLJK ,QVSLWH RI WKLV ORQJWHUP FKDQJH LQ WKH OHYHO RI WKH GLIIHUHQW psbA WUDQVFULSWV WKH ' )RUP ,, SURWHLQ UHPDLQV DVVRFLDWHG ZLWK 36,, VXJJHVWLQJ SRVWWUDQVFULSWLRQ UHJXODWLRQRISURWHLQDEXQGDQFHHJDIDVWHUWXUQRYHU RIWKH' )RUP, 3RVWWUDQVFULSWLRQDO UHJXODWLRQ LQ WKLV V\VWHP SUREDEO\ RFFXUV DW D QXPEHU RIGLIIHUHQW OHYHOV 7KHNLQHWLFV RIGHJUDGDWLRQ RIDOOWKUHHRIWKH psbA PHVVDJHV DUH GLIIHUHQW .XONDUQL HW DO )XUWKHUPRUH WKH UDWH RI WXUQRYHU RI WKH psbAI WUDQVFULSW LQFUHDVHV XSRQ WUDQVIHU RIWKH FHOOV WR KLJK OLJKW &XULRXVO\ WKH psbAIII WUDQVFULSW DOVR VKRZV HQKDQFHGGHJUDGDWLRQ LQKLJKOLJKWZKLOHWKHVWDELOLW\ RI WKH psbAII WUDQVFULSW LV XQDIIHFWHG E\ OLJKW
' %KD\D 56FKZDU]DQG$5*URVVPDQ LQWHQVLW\ 5HFHQWO\ .XONDUQL DQG *ROGHQ KDYH LGHQWLILHG DQ HOHPHQW WKDW VWDELOL]HV psbA P51$V ZKLFKPDSVZLWKLQWKHUHJLRQRIWKHWUDQVFULSWFRGLQJ IRU WKH ILUVW PHPEUDQH VSDQ RI WKH ' SRO\SHSWLGH 7KLV VWDELOL]DWLRQ HIIHFW LV LQGHSHQGHQW RI OLJKW LQWHQVLW\*ROGHQDQG.XONDUQL 7KH VLJQDO WUDQVGXFWLRQ SDWKZD\ WKDW FRQWUROV WKH UDWHV RI WUDQVFULSWLRQ RI WKH psb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hli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b6f, DV ZRXOGRFFXULQKLJKOLJKWDUHLPSRUWDQWLQFRQWUROOLQJ psbAII DQG psbAIII, +RZHYHU LW LV SRVVLEOH DV VXJJHVWHGE\WKHDXWKRUVWKDWPRGHUDWHFORVXUHRIWKH 36,,UHDFWLRQFHQWHUV LV FUXFLDO LQUHJXODWLQJWKHpsbA JHQHIDPLO\ ,WLV QRWDEVROXWHO\FOHDUWKDW'&08LV FRQWUROOLQJ WKHUDWLR RI' )RUP, WR ' )RUP ,, E\ DOWHULQJWKHOHYHOVRIWKHpsbA WUDQVFULSWV *ROGHQDQG FRZRUNHUV IRXQGQR HIIHFWRISKRWRV\QWKHWLFHOHFWURQ WUDQVSRUW LQKLELWRUV VXFK DV '&08 RQ WKH DFFXPXODWLRQ RIWKHpsbAII DQG psbAIII WUDQVFULSWV 6 *ROGHQ SHUV FRPP 7KHVH REVHUYDWLRQV FDQ EH H[SODLQHGLI'&08ZKLFKELQGVWR'GLIIHUHQWLDOO\ DIIHFWV WKH VWDELOLW\ RI WKH WZR IRUPV RI ' UDWKHU WKDQFDXVLQJDFKDQJHLQWKHWUDQVFULSWLRQRIWKHpsbA JHQHV
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO6WUHVV &RQWURORI3KRWRV\VWHP 6WRLFKLRPHWU\ (DUO\VWXGLHVZLWKF\DQREDFWHULDDQGUHGDOJDHZKLFK OLNH F\DQREDFWHULD FRQWDLQ 3%6 GHPRQVWUDWHG YDULDELOLW\ RIWKH TXDQWXP\LHOG RISKRWRV\QWKHVLV IRU OLJKW DEVRUEHG E\ FKO a. 7KLV YDULDELOLW\ ZDV GHSHQGHQW RQ WKH OLJKW FRQGLWLRQV HJ H[FLWDWLRQ HQHUJ\SUHIHUHQWLDOO\DEVRUEHGE\36,RU36,, GXULQJ JURZWK DQG FRUUHODWHG ZLWK FKDQJHV LQ FHOOXODU SLJPHQW FRPSRVLWLRQ %LJJLQV DQG %UXFH 0\HUV HWDO ZHUH WKH ILUVWWR REVHUYH WKDWWKH 36,36,, UDWLR ZDV KLJKHU LQ Synechococcus 3&& (Anacystis nidulans) FHOOV JURZQ LQ OLJKW DEVRUEHG PDLQO\ E\ WKH 3%6 WKDQ LQ FHOOV JURZQ LQ OLJKW DEVRUEHG PDLQO\ E\ FKO a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a WKH PDMRU OLJKWKDUYHVWLQJ SLJPHQW IRU 36, H[KLELW HOHYDWHG OHYHOV RI3%6 UHODWLYHWRFKOa. 7KLVDOWHUHG UDWLR ZKLFK DJDLQ WHQGV WR IDYRU EDODQFHG OLQHDU HOHFWURQ IORZ LV SULPDULO\ D FRQVHTXHQFH RI D GHFUHDVH LQ WKH QXPEHU RI 36, UHDFWLRQ FHQWHUV $L]DZD HW DO )XMLWD DQG 0XUDNDPL )XMLWDHWDO )XMLWDHWDO ,Q WKH QDWXUDO HQYLURQPHQW F\DQREDFWHULD DUH PRVW IUHTXHQWO\H[SRVHGWRµZKLWHOLJKW¶ZKLFKFRQWDLQVDOO ZDYHOHQJWKV RIWKH YLVLEOH VSHFWUXP 6LQFH WKH 3%6 DEVRUEVDODUJHSURSRUWLRQ RIWKHLQFLGHQWZKLWHOLJKW WKHUH ZLOO EH SUHIHUHQWLDO H[FLWDWLRQ RI 36,, ZKHQ
F\DQREDFWHULD DUH H[SRVHG WR QRQVDWXUDWLQJ ZKLWH OLJKW ZKLFK ZRXOG VHUYH DV D VLJQDO WKDW SURPSWV HOHYDWHG V\QWKHVLV RI 36, UHODWLYH WR 36,, $V WKH LQWHQVLW\ RIZKLWH OLJKW LQFUHDVHV ERWK SKRWRV\VWHPV EHFRPH VDWXUDWHG DQG WKH 36,36,, UDWLR IDOOV DSSURDFKLQJ WKDW PHDVXUHG LQ FHOOV JURZQ LQ FKO a OLJKW 2WKHU OLJKWPHGLDWHG FRQWURO PHFKDQLVPV WKDW DUH GLVWLQFW IURP WKRVH WKDW UHJXODWH SKRWRV\VWHP VWRLFKLRPHWU\ PD\ DOVR LQIOXHQFH SKRWRV\QWKHWLF DFWLYLW\ )RUH[DPSOHSURORQJHGKLJKOLJKWLQWHQVLWLHV PD\ OHDG WR D GHFUHDVH LQ WKH WRWDO QXPEHU RI SKRWRV\VWHPV
RQH RU PRUH SURVWKHWLF JURXS DQG DVVHPEOH LQWR IXQFWLRQDOPXOWLVXEXQLWFRPSOH[HV 7KHFRRUGLQDWLRQ RI WKH GLVWLQFW VWHSV WKDW PXVW EH WHPSRUDOO\ DQG VSDWLDOO\ FRQWUROOHG DQG WKDWPD\ LQYROYH D KLHUDUFK\ RIUHJXODWRU\HYHQWVLVVWLOODP\VWHU\
&RQWURORI3K\FRELOLVRPH %LRV\QWKHVLV E\/LJKW4XDOLW\ D 3K\FRELOLVRPH 6WUXFWXUH $EULHIGHVFULSWLRQRIWKH3%6VWUXFWXUHLVSURYLGHGWR KHOS WKH UHDGHU XQGHUVWDQG WKH ZD\V LQ ZKLFK OLJKW FDQ FRQWURO 3%6 ELRV\QWKHVLV ,Q HDUO\ VWXGLHV HOHFWURQ PLFURJUDSKV RI F\DQREDFWHULD VKRZHG UHJXODUO\ DUUDQJHG SDUDOOHO URZV RI JUDQXOHV RQ WKH VXUIDFH RI WKH WK\ODNRLG PHPEUDQH *DQWW *OD]HU 7KHVHSURIXVHJUDQXOHVZHUH3%6WKDW ZHUH H[WULQVLFDOO\ DWWDFKHG WR WKH PHPEUDQH VXUIDFH 3%6 PD\FRQVWLWXWHDVPXFKDVRIWKHSURWHLQRI WKH FHOO DQG DUH FRPSRVHG RI VHYHUDO GLIIHUHQW SLJPHQWHG ZDWHUVROXEOH SRO\SHSWLGHV WKDW JLYH F\DQREDFWHULD WKHLU FKDUDFWHULVWLF EOXHJUHHQ RU UHG FRORU 7KH 3%6 IXQFWLRQV DV D OLJKWKDUYHVWLQJ DSSDUDWXV WKDW DEVRUEV SKRWRQV DQG WUDQVPLWV WKH H[FLWRQHQHUJ\ WR WKH SKRWRV\QWKHWLF UHDFWLRQ FHQWHUV ZLWKDQH[WUHPHO\KLJKHIILFLHQF\3RUWHUHWDO 6HDUOHHWDO $ V\QWKHVLV RI WKH LQIRUPDWLRQ JHQHUDWHG RYHU D VFRUHRI\HDUVIURPHOHFWURQPLFURVFRS\ ELRFKHPLFDO DQG ELRSK\VLFDO VWXGLHV ;UD\ FU\VWDOORJUDSK\ DQG PROHFXODU DQDO\VHV RI JHQHV HQFRGLQJ 3%6 SRO\SHSWLGHVKDV OHG WRWKHGHYHORSPHQWRIDGHWDLOHG PRGHO RI3%6 VWUXFWXUH %U\DQW %U\DQWHWDO *DQWW*OD]HU *OD]HUHWDO $ GLDJUDPPDWLF UHSUHVHQWDWLRQ RIWKH 3%6 LV VKRZQ LQ)LJ 6HFWLRQ,,EIRUPRUHGHWDLO (DFK3%6LV RUJDQL]HGLQWRURG DQGFRUHVXEVWUXFWXUHV 7KHFRUH PDLQO\ FRPSRVHG RI DOORSK\FRF\DQLQ $3 $PD[ QP WULPHUV LV LQ GLUHFW FRQWDFW ZLWK WK\ODNRLG PHPEUDQHV YLD D ODUJH SLJPHQWHG SRO\SHSWLGH /&0 WKDW VHUYHV DV D WHUPLQDO HQHUJ\ DFFHSWRU RI WKH 3%6 7KHUH DUH XVXDOO\ VL[ VHWV RI URGVWKDWUDGLDWH RXWIURPWKHFRUHLQDIDQOLNHDUUD\ 5RGV DUH FRPSRVHG RI KH[DPHUV RI WKH FKURPRSKRU\ODWHG SK\FRF\DQLQ 3& $PD[ QP DQG VRPHWLPHV SK\FRHU\WKULQ 3( $PD[ QP RU SK\FRHU\WKURF\DQLQ3(&$PD[ QP (DFKRIWKH SK\FRELOLSURWHLQ PRQRPHUV KDV WZR GLIIHUHQW VXEXQLWV DQG ZLWK PROHFXODUPDVVHV RIEHWZHHQ DQG N'D HDFK 7KH FKURPRSKRUHV RU OLJKW DEVRUELQJ PROHFXOHV WKDW DUH DWWDFKHG WR WKH
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a RIWKH SKRWRV\QWKHWLF UHDFWLRQ FHQWHUV E
&RPSOHPHQWDU\ &KURPDWLF $GDSWDWLRQ
7KH FRQVWUXFWLRQ RID 3%6 LV D G\QDPLF SURFHVV WKDW LV UHVSRQVLYH WR FKDQJLQJ HQYLURQPHQWDO FRQGLWLRQV ,Q F\DQREDFWHULD QXWULHQW OHYHOV &ROOLHU DQG *URVVPDQ &ROOLHU DQG *URVVPDQ
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO6WUHVV
5RG VXEXQLW FRPSRVLWLRQ
5RG VXEXQLW FRPSRVLWLRQ
Fig.1. &KURPDWLFDGDSWDWLRQ DQGWKHFRPSRVLWLRQ RIWKH F. diplosiphon 3%6 IURPFHOOV PDLQWDLQHG LQJUHHQ OLJKW*/ DQGUHG OLJKW5/ 7KHFRPSRVLWLRQRIWKHSHULSKHUDOURGV IURP5/JURZQFHOOVDUHJLYHQE\ DQG ZKLOHWKHFRPSRVLWLRQRIWKHURGVLQ*/JURZQ FHOOV DUHJLYHQE\ DQG 7KH3%6URGVIURP5/ JURZQFHOOVDUHFRPSRVHGH[FOXVLYHO\RUDOPRVWH[FOXVLYHO\ RI3&KH[DPHUVZKLOHWKH URGVIURP*/JURZQFHOOVUHWDLQRQO\RQH3&KH[DPHUQHDUHVWWRWKHFRUH DQGFRQWDLQ PRVWO\3(KH[DPHUV 7KHURGVXEXQLWFRPSRVLWLRQ UHIHUVWRWKH DQGVXEXQLWVWKDWFRPSULVHWKH3(KH[DPHU 3& RU3&KH[DPHU 3( (DFK OLQNHU / SRO\SHSWLGH LV DVVRFLDWHG ZLWK DVSHFLILF VHWRIKH[DPHUV DQGLV VKRZQZLWKDVXSHUVFULSWWKDWUHIHUV WRWKHPROHFXODUPDVV DQG DVXEVFULSW WKDWUHIHUV WR LWV SRVLWLRQ ZLWKLQ WKHKH[DPHU5 IRUURGRU 5& IRUURGFRUH )RUIXUWKHUGHWDLOV RQ FRUH FRPSRVLWLRQ DQG JHQHV HQFRGLQJ SK\FRELOLSURWHLQV UHIHU WR *URVVPDQHWDO
7KLV SURFHVV LV YLVXDOO\ PRVW DSSDUHQW ZKHQ WKH DGDSWLQJRUJDQLVPV DUH FRPSDUHG DIWHUJURZWK LQUHG OLJKW5/ RUJUHHQOLJKW*/ ,Q5/F. diplosiphon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
SK\FRELOLSURWHLQ VXEXQLWV DQG DVVRFLDWHG / SURWHLQV )XUWKHUPRUH GHSHQGLQJ XSRQ WKH RUJDQLVP WKH GLIIHUHQW SK\FRELOLSURWHLQV PD\ EH HQFRGHG E\ D PXOWLJHQH IDPLO\ 7KH 3%6 JHQHV WKDW DUH GLIIHUHQWLDOO\ UHJXODWHG GXULQJ &&$ DUH WKRVH HQFRGLQJ 3( 3& DQG WKH / SRO\SHSWLGHV WKDW DVVRFLDWH ZLWKWKH3( DQG 3& KH[DPHUV 1RQH RIWKH JHQHV HQFRGLQJ WKH FRUH FRPSRQHQWV RIWKH 3%6 DUH GLIIHUHQWLDOO\UHJXODWHGGXULQJ&&$ 7KHUH DUH WKUHH GLVWLQFW JHQH VHWV GHVLJQDWHG cpcB1A1, cpcB2A2 and cpcB3A3) WKDWHDFKHQFRGH DQGVXEXQLWVRI3&LQF. diplosiphon &RQOH\HWDO &RQOH\HWDO &RQOH\HWDO 0D]HO HW DO 0D]HO DQG 0DUOLHUH 7KH cpcB1A1 RSHURQ LV FRQVWLWXWLYHO\ H[SUHVVHG DQG WKH 3& WKDW LW HQFRGHV LV UHIHUUHG WR DV 3& VXEVFULSW GHQRWHV FRQVWLWXWLYH ,W DSSHDUV WR EH FRWUDQVFULEHG ZLWK cpcE DQG SHUKDSV cpcF, ZKLFK HQFRGH D O\DVH WKDW DVVLVWV LQ WKH DWWDFKPHQW RI WKH WHWUDS\UUROH FKURPRSKRUHV WR WKH VXEXQLW RI3& )DLUFKLOG DQG *OD]HU )DLUFKLOG HW DO $ VHFRQG
RSHURQ WKDW LQFOXGHV cpcB2A2 EHFRPHV WUDQVFULSWLRQDOO\ DFWLYH RQO\ LQ 5/ LW LV LQDFWLYH LQ */ DQG WKH 3& WKDW LW HQFRGHV LV VSHFLILFDOO\ GHVLJQDWHG 3& VXEVFULSW GHQRWHVLQGXFLEOH &RQOH\ HW DO &RQOH\ HWDO 7KH LQGXFWLRQ RI WKLV RSHURQ LV FULWLFDO IRU &&$ +H[DPHUV RI 3&L FRQVWLWXWH WKH PDMRULW\ RI WKH 3%6 URG VXEVWUXFWXUH ZKHQ F. diplosiphon LV JURZQ LQ 5/ 7KUHH JHQHV HQFRGLQJ WKH / SRO\SHSWLGHV WKDW DVVRFLDWH ZLWK 3& DUH FRWUDQVFULEHG ZLWK cpcB2A2 DQG DUH GHVLJQDWHG cpcH2, cpcI2 DQG cpcD2 /RPD[HWDO 7KH WKLUG 3& RSHURQ FRQWDLQV cpcB3A3 DV ZHOO DV JHQHV HQFRGLQJ DVVRFLDWHG /SRO\SHSWLGHV ,WLV RQO\DFWLYH GXULQJ VXOIXUOLPLWHG JURZWK 0D]HO DQG 0DUOLHUH DQG WKH 3&V VXEVFULSW LQGLFDWLQJ DFWLYDWLRQ GXULQJ 6XOIXUOLPLWDWLRQ HQFRGHG E\ cpcB3A3, KDV QR VXOIXUFRQWDLQLQJ DPLQR DFLGV RWKHU WKDQ WKH F\VWHLQHV WKDW DUH HVVHQWLDO IRU WKH DWWDFKPHQW RIWKH SK\FRELOLQ FKURPRSKRUHV +HQFH ZKHQ VWDUYHG IRU VXOIXU F. diplosiphon FDQ FRQVHUYH WKLV QXWULHQW E\ VXEVWLWXWLQJ 3&L DQG SHUKDSV 3&F ZLWK 3&V 3( LV HQFRGHG E\ WKH cpeBA RSHURQ 0D]HO HW DO DQGWKHDVVRFLDWHGOLQNHUSRO\SHSWLGHVDUHHQFRGHGE\ D VHSDUDWH RSHURQ cpeCDE )HGHUVSLHO DQG *URVVPDQ )HGHUVSLHO DQG 6FRWW 7KH cpeBA DQG cpeCDE RSHURQV DUH FRRUGLQDWHO\ UHJXODWHG WUDQVFULSWV IURP ERWK RIWKHVH RSHURQV DUH DEXQGDQWLQ*/DQGEDUHO\GHWHFWDEOHLQ5/
L 3URPRWHU %LQGLQJ IDFWRUV 7R VWXG\ UHJXODWRU\ HOHPHQWV WKDW PD\ ELQG WR WKH cpeBA SURPRWHU VROXEOHSURWHLQV IURP5/ DQG */ JURZQ F. diplosiphon RU Calothrix 3&& FHOOV ZHUH XVHG LQ PRELOLW\ VKLIW DVVD\V 6ROXEOH F\DQREDFWHULDO SURWHLQV ELQG WR WKH cpeBA SURPRWHU UHWDUGLQJ LWV PRELOLW\ GXULQJ HOHFWURSKRUHVLV LQ D SRO\DFU\ODPLGH JHO 7KUHH SURWHLQV DSSHDUHG WR LQWHUDFW ZLWK VHTXHQFHV SRVLWLRQHG WR UHODWLYH WR WKH WUDQVFULSWLRQ VWDUW VLWH RI WKH cpeBA RSHURQ 6REF]\N HW DO 51$ SRO\PHUDVH LQWHUDFWHG ZLWK WKH VHTXHQFH IURP WR 7ZR RWKHUSURWHLQV GHVLJQDWHG 5FD$ DQG 5FD% DSSHDUHG WREHSUHVHQWRQO\LQ*/JURZQFHOOV 5FD$LQWHUDFWV ZLWKWKH VHTXHQFHIURP WR ZKLFK FRQWDLQV D WDQGHPUHSHDW 77*77$1 77*77$ 6FKPLGW *RIIDQG)HGHUVSLHO XVHG in vivo IRRWSULQWLQJ in vitro HOHFWURSKRUHWLF PRELOLW\ VKLIW DVVD\V DQG GLPHWK\OVXOIDWHDQG'1DVH, in vitro IRRWSULQWLQJ WR VKRZ WKDW D SURWHLQ GHVLJQDWHG 3HS% 3( SURPRWHU ELQGLQJ SURWHLQ ERXQG WR DQG SURWHFWHG WKH 77*77$ WDQGHP UHSHDW ORFDWHG XSVWUHDP RI WKH
'%KD\D56FKZDU]DQG$5*URVVPDQ WUDQVFULSWLRQVWDUWVLWH 6LQFH5FD$DQG3HS%ELQGWR WKH VDPH VHTXHQFHV LW LV SUREDEOH WKDW WKH\ DUH LGHQWLFDOSURWHLQV 7KH WDQGHP UHSHDW SUHVHQW XSVWUHDP RI cpeBA SURPRWHUV ZDV DOVR REVHUYHG LQ RWKHU F\DQREDFWHULD WKDWH[KLELW&&$ +RZHYHU LWZDVQRWSUHVHQWLQWKH cpeCDE RSHURQ LQVSLWH RI WKH IDFW WKDW WKLV RSHURQ ZDVFRQWUROOHGLGHQWLFDOO\WRcpeBA. 7KHUHIRUH LIWKH 77*77$ PRWLILV LPSRUWDQW IRU &&$ DGGLWLRQDO HOHPHQWV PXVW EH LQYROYHG LQ FRRUGLQDWLQJ WKH WUDQVFULSWLRQDOUHVSRQVHVRIcpeBA DQG cpeCDE. 7KH H[DFW SRVLWLRQ RI WKH ELQGLQJ VLWH IRU 5FD% LV QRW FOHDU 3URWHLQV ZHUH UHSRUWHG WR ELQG WR WKH SURPRWHU RI WKH cpcB2A2 RSHURQ ZKLFKHQFRGHV 3& 6REF]\NHW DO 7R IXUWKHU DQDO\]H WKLV SURPRWHU D FKLPHULF UHSRUWHU JHQH ZDV FRQVWUXFWHG LQ ZKLFK WKH cpcB2A2 SURPRWHU ZDV IXVHG WR WKH uidA JHQH HQFRGLQJ JOXFXURQLGDVH RU *86 7KH SURPRWHU UHJLRQ RI WKH FKLPHULF JHQH ZDV PRGLILHG UHLQWURGXFHG LQWR F. diplosiphon, DQG WUDQVIRUPDQWV ZHUH DVVD\HG IRU OLJKWUHVSRQVLYH *86 DFWLYLW\ $ IUDJPHQW H[WHQGLQJ IURP WR UHODWLYH WR WKH cpcB2A2 WUDQVFULSWLRQ VWDUW VLWH FRQIHUUHG 5/*/ UHVSRQVLYHQHVV WR *86 H[SUHVVLRQ 'HOHWLRQV H[WHQGLQJ WR UHGXFHG *86 DFWLYLW\ LQ ERWK 5/ DQG */ VXJJHVWLQJ WKDW WKH VHTXHQFH EHWZHHQ DQG LV HVVHQWLDO IRU WKH DFWLYLW\ RI WKH cpcB2A2 SURPRWHU *HOPRELOLW\VKLIWDVVD\VGHPRQVWUDWHGWKDW D SURWHLQ LQ H[WUDFWV IURP 5/JURZQ EXW QRW */ JURZQ FHOOV ZDV DEOH WR ELQG WR D WR ES IUDJPHQW $'1$IUDJPHQWIURPWRZDVDEOH WR FRPSHWH IRU WKLV ELQGLQJ DFWLYLW\ &DVH\ DQG *URVVPDQ VXJJHVWLQJ WKDW WKH WR UHJLRQ RI WKH cpcB2A2 SURPRWHU LQWHUDFWV ZLWK D SURWHLQWKDWLVHLWKHURQO\SUHVHQWRURQO\ELQGVLQ5/ JURZQ FHOOV 7RJHWKHU WKH UHVXOWV VXJJHVW WKDW WKH UHJLRQ cpcB2A2 RSHURQ IURP WR LV UHTXLUHG IRU RYHUDOO SURPRWHU DFWLYLW\ ZKLOH WKH WR VHTXHQFH ELQGV D UHJXODWRU\ SURWHLQ ZKLFK LQ WXUQ SURPRWHV WUDQVFULSWLRQ IURP cpcB2A2 LQ 5/ 7KLV ELQGLQJ VLWH ZDV VKRZQ WR FRQWDLQ D GLUHFW UHSHDW RI WKH VHTXHQFH $$$777*&$&$$$ &DVH\ DQG *URVVPDQ
LL 0XWDQW&KDUDFWHUL]DWLRQ 2YHU WKH ODVW VHYHUDO \HDUV QXPHURXV PXWDQWV RI F. diplosiphon ZHUH JHQHUDWHG DQG FKDUDFWHUL]HG %UXQV HWDO &KLDQJHWDO &REOH\DQG0LUDQGD .HKRH DQG *URVVPDQ 7DQGHDX GH 0DUVDF 7KHVH PXWDQWV FDQ EH SODFHG LQWR DW
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diplosiphon. 7KH )G%N PXWDQW ZDV FRPSOHPHQWHG ZLWK D JHQH GHVLJQDWHG rcaE ZKLOH GLIIHUHQW )G5 PXWDQWV ZHUH FRPSOHPHQWHGE\ rcaF, rcaE .HKRH DQG *URVVPDQ RUrcaC &KLDQJHWDO 6LJQLILFDQWO\DOO WKH FRPSRQHQWV HQFRGHG E\ WKHVH JHQHV DUH SURWHLQV WKDWKDYH VLPLODULW\WR FRPSRQHQWVRIWZRFRPSRQHQW UHJXODWRU\V\VWHPV 7KH N'DSURWHLQ HQFRGHG E\ rcaE KDVDFDUER[\WHUPLQDOUHJLRQZLWKVLPLODULW\WR RXWSXW GRPDLQV RI KLVWLGLQH NLQDVH VHQVRUV 6XUSULVLQJO\ WKH DPLQRWHUPLQDO KDOI RI WKH SURWHLQ RYHU D UHJLRQ RI DSSUR[LPDWHO\ DPLQR DFLGV LV VLPLODU WR WKH WHWUDS\UUROH FKURPRSKRUH DWWDFKPHQW GRPDLQ RIWKH SK\WRFKURPHV .HKRH DQG *URVVPDQ 5HFHQWO\LWZDVVKRZQXVLQJDFRPELQDWLRQRI DQWLERGLHV DQG ]LQFHQKDQFHG IOXRUHVFHQFH WKDW WKH 5FD( SURWHLQ LV WLJKWO\ DVVRFLDWHG ZLWK D OLQHDU WHWUDS\UUROH .HKRH DQG *URVVPDQ XQSXEOLVKHG GDWD 3K\WRFKURPHV DUH ZHOOFKDUDFWHUL]HG SKRWRUHFHSWRUVWKDWFRQWURO DZLGHUDQJH RIUHVSRQVHV LQ YDVFXODUSODQWV .HQGULFNDQG .URQHQEHUJ 5FD( DOVR KDV VHTXHQFH VLPLODULW\ WR YDVFXODU SODQW HWK\OHQH UHFHSWRUV &KDQJ HW DO +XD HW DO :LONLQVRQHWDO 7KHHWK\OHQHUHFHSWRUV DUH VLPLODU WR VHQVRU NLQDVHV RI EDFWHULDO WZR FRPSRQHQWUHJXODWRU\ V\VWHPV 7KHSKHQRW\SHRIWKH )G%N PXWDQW .HKRH DQG *URVVPDQ WKH VLPLODULW\ RI 5FD( WR D YDVFXODU SODQW SKRWRUHFHSWRU DQG VHQVRU KLVWLGLQH NLQDVHV DQG WKH DVVRFLDWLRQ RI WKH 5FD( SRO\SHSWLGH ZLWK D WHWUDS\UUROH FKURPRSKRUH DUH DOO FRQVLVWHQW ZLWK 5FD( EHLQJ LQYROYHGLQSKRWRSHUFHSWLRQ 7KH 5FD) SURWHLQ HQFRGHG E\ D JHQH ZKLFK LV LQLWLDWHG ES GRZQVWUHDP RI WKH WUDQVODWLRQ WHUPLQDWLRQVLWHRIrcaE, LVSUHGLFWHGWREH DPLQR
DFLGV DQG LV KRPRORJRXV WR UHVSRQVH UHJXODWRUV +RZHYHU OLNH &KH< DQG 6SR) UHVSRQVH UHJXODWRUV LQYROYHG LQ IODJHOODU PRYHPHQW LQ E. coli DQG VSRUXODWLRQ LQ Bacillus subtilis, UHVSHFWLYHO\ &OHJJ DQG .RVKODQG 3DUNLQVRQ DQG .RIRLG 3HUHJRDQG+RFK 5DYLG HWDO :ROIHHW DO
'%KD\D 56FKZDU]DQG$5*URVVPDQ
Fig. 2. 0RGHOIRUFRQWURORISK\FRELOLSURWHLQJHQHVE\UHGOLJKW5/ GXULQJFRPSOHPHQWDU\FKURPDWLFDGDSWDWLRQ 7KHSKRVSKRUHOD\WKDWLV DFWLYDWHGE\5/DQGUHJXODWHVWKHJHQHVIRU3&LcpcB2A2 DQG3(cpe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cpcB2A2 DQG cpeBA RSHURQV WR GLUHFWO\ UHJXODWH WUDQVFULSWLRQ RU ZKHWKHU DGGLWLRQDO UHJXODWRU\IDFWRUVDUHUHTXLUHG :KHQ*/LVDEVRUEHGE\5FD(WKHSKRVSKRUHOD\LV PDLQWDLQHGLQDGHSKRVSKRU\ODWHGVWDWH7KLVFRXOGEH DFFRPSOLVKHG LI 5FD( ELQGV WR DQG EORFNV SURPLVFXRXVSKRVSKRU\ODWLRQRI5FD)WKDWFDQ RFFXU WKURXJK LQWHUDFWLRQV ZLWK RWKHU VHQVRUV LQ WKH FHOO RU VPDOO SKRVSKRGRQRU PROHFXOHV RU LI5FD( DFWV DV D
SKRVSKDWDVH UHPRYLQJ DQ\ SKRVSKDWH WKDW LV WUDQVIHUUHGWR5FD)
,,, 5HVSRQVHVWR 1XWULHQW /LPLWDWLRQ 7KHUHVSRQVHV RIF\DQREDFWHULD WR QXWULHQW GHILFLHQF\ KDYH EHHQ VWXGLHG H[WHQVLYHO\ $OOHQ %U\DQW %U\DQW +HDOH\ 5HLWKPDQ HW DO 6LPRQ 7DQGHDX GH 0DUVDF DQG +RXPDUG 6RPH RI WKH UHVSRQVHV RI WKHVH UHVSRQVHV DUH VSHFLILF WULJJHUHG E\ D GHILFLHQF\ IRU RQH SDUWLFXODU QXWULHQW ZKLOH RWKHUV DUH JHQHUDO DQG RFFXU GXULQJ GHILFLHQF\ IRU DQ\ RI D QXPEHU RI GLIIHUHQW QXWULHQWV 7KH IRUPHU FDWHJRU\ LQFOXGHV LQFUHDVHG V\QWKHVLV RI VSHFLILF WUDQVSRUW V\VWHPV DQG HQ]\PHV WKDW WUDQVIRUP LQDFFHVVLEOH IRUPV RI D QXWULHQW LQWR WKRVH WKDW WKH FHOO FDQ PHWDEROL]H 7KH ODWWHU FDWHJRU\ LQFOXGHV FKDQJHV LQ FHOO PRUSKRORJ\ LQWUDFHOOXODU QXWULHQW UHVHUYHV DQG DOWHUDWLRQV LQ WKH DFWLYLW\ RI QXPHURXV SK\VLRORJLFDO SURFHVVHV LQFOXGLQJ SKRWRV\QWKHVLV *URVVPDQ HW DO D *URVVPDQ HW DO E )RU FRQYHQLHQFH Synechococcus 3&& Anacystis nidulans, Synechococcus 3&& DQG Synechococcus leopoliensis, ZKLFK DUH DOO YHU\ VLPLODU VWUDLQV ZLOO VRPHWLPHV EH UHIHUUHG WR DV Synechococcus VS
&KDSWHU 0ROHFXODU 5HVSRQVHVWR (QYLURQPHQWDO6WUHVV /LNHZLVH D QXPEHU RI Anabaena VWUDLQV DUH DOVR WUHDWHG DV D JURXS 7KH IROORZLQJ GLVFXVVLRQ LV OLPLWHG WR WKH UHVSRQVHV RI F\DQREDFWHULD WR PDFURQXWULHQW OLPLWDWLRQ ZKLFK LQFOXGHV FDUERQ & QLWURJHQ1 SKRVSKRUXV3 DQGVXOIXU6
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ynechococcus :+ GLG QRW LQGXFH D &&0 .DUDJRXQL HW DO $ UH HYDOXDWLRQ RI WKLV LQGLFDWHV WKDW WKLV RUJDQLVP GRHV FRQFHQWUDWH&ZKHQPDLQWDLQHGLQORZLQWHQVLW\OLJKW $ .DSODQ SHUV FRPP LQ WKH LQLWLDO VWXG\ .DUDJRXQL HW DO WKH UHODWLYHO\ KLJK SKRWRQ IOX[ GHQVLWLHV XVHG PD\ KDYH FDXVHG FRQVLGHUDEOH SKRWRGDPDJH 7KH &&0 SURPRWHV WKH IL[DWLRQ RI&2 E\ UDLVLQJ LWVFRQFHQWUDWLRQDURXQG5X%3FDVHDQHQ]\PHZLWKD UHODWLYHO\ORZDIILQLW\ IRU LWV VXEVWUDWH &&0 DFWLYLW\ PD\ DOVR EH SDUWLDOO\ UHVSRQVLEOH IRU GLIIHUHQFHV
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
0HFKDQLVPRI&L 8SWDNH $ UDWLR RI >&L@LQ>&L@RXW WKDWUHDFKHG KDV EHHQ REVHUYHG LQ ORZ&2 DGDSWHG FHOOV 7DNLQJ LQWR DFFRXQW WKH DQG WKH PHPEUDQH SRWHQWLDO DQ DFWLYH SURFHVV PXVW EH UHVSRQVLEOH IRU FRQFHQWUDWLQJ &L DJDLQVW LWV HOHFWURFKHPLFDO SRWHQWLDO 7KH DGGLWLRQ RI +&2 WR FXOWXUHV RI Synechococcus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
' %KD\D 5 6FKZDU]DQG$5*URVVPDQ
)LJ $ VFKHPDWLF SUHVHQWDWLRQ RI&L IOX[HV DVVRFLDWHG ZLWK WKH &&0 LQ F\DQREDFWHULD 7KH F\WRSODVP FDUKR[\VRPDO VSDFH WK\ODNRLG UHJLRQXQVWLUUHGOD\HUVXUURXQGLQJWKHFHOODQGWKHEXONPHGLXPDUHLQGLFDWHG(QHUJ\GHSHQGHQW&L IOX[HVDUHGHVLJQDWHG( DQGSUHVHQWHG DV KHDY\ OLQHV 'DVKHG DUURZV LQGLFDWH WKDW WKH &L VSHFLHV DUH QRW DW FKHPLFDO HTXLOLEULXP LQ WKH F\WRSODVP DQG WKH XQVWLUUHG OD\HU $V LQGLFDWHGLQWKHWH[W WKH&2 JHQHUDWHGLQWKHFDUER[\VRPHV E\ WKHDFWLYLW\RIFDUERQLF DQK\GUDVHLV PRVWO\ IL[HGE\ 5X%3FDVH EXWVRPH PD\GLIIXVHRXWRIWKHFDUER[\VRPHDQGLQWRWKHF\WRSODVPRIWKHFHOO
5HTXLULQJ +&5 PXWDQW ,/ LPSDLUHG LQ +&2 WUDQVSRUW WKDW KDV D OHVLRQ LQ D JHQH HQFRGLQJ D SURWHLQ VLPLODU WR WKH 1DSDQWRWKHQDWH V\PSRUWHU RI E. coli DQG WR D IDPLO\ RI PXOWLGUXJ UHVLVWDQFH SURWHLQV5RQHQ7DUD]LHWDO E 2QH RIWKH PDMRU SURWHLQV LQGXFHG XSRQ H[SRVXUH RI Synechococcus 3&& WR DLUOHYHOV RI&2 LV D N'D PHPEUDQH SURWHLQ 7KH JHQH HQFRGLQJ WKLV SURWHLQ cmpA, LV SDUW RIWKH cmpABCD JHQH FOXVWHU ZKLFK HQFRGHV SURWHLQV KRPRORJRXV WR FRPSRQHQWV RI $%& WUDQVSRUW V\VWHPV 6X]XNL HW DO %DVHG RQWKLVKRPRORJ\ LWZDV SURSRVHG WKDWWKH N'D SRO\SHSWLGH ZDV LQYROYHG LQ &L WUDQVSRUW 6XUSULVLQJO\ LQDFWLYDWLRQ RIWKH cmpA JHQH GLG QRW DIIHFW JURZWK DW DLU OHYHOV RI &2 RU &L WUDQVSRUW
2PDWD HWDO ,W LV SRVVLEOH WKDW WKH WUDQVSRUW FRPSRQHQWV HQFRGHG E\ WKH cmpABCD RSHURQ DUH HVVHQWLDO DW FRQFHQWUDWLRQV RI&2 ORZHU WKDQ WKDW LQ DLU 7KH WUDQVSRUW RI +&2 DQG &2 LQWR F\DQREDFWHULDO FHOOV SUREDEO\ RFFXUV YLD GLVWLQFW WUDQVSRUW V\VWHPV 7KLV LV VXSSRUWHG E\ WKH ILQGLQJ WKDW F\DQREDFWHULDO PXWDQWV FDQ EH VSHFLILFDOO\ LPSDLUHG LQ +&2 5RQHQ7DUD]L HW DO E RU &2 WUDQVSRUW .DWRK HW DO D 7KH GHIHFWLYH JHQH LQ D VWUDLQ XQDEOH WR WUDQVSRUW &2 cotA, LV VLPLODUWR cemA, ZKLFKHQFRGHV DSRO\SHSWLGHWKDW LV ORFDOL]HG WR WKH LQQHU PHPEUDQH RI SHD FKORURSODVWV .DWRKHWDO D 7KH cotA JHQHSURGXFWKDV DPLQR DFLGV DQG LV ORFDWHG LQ ERWK WKH F\WRSODVPLF
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO 6WUHVV DQG WK\ODNRLG PHPEUDQHV 2JDZD 6WUDLQV LQ ZKLFK cot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ynechococcus 3&& DIWHU WKH PHGLXP ZDV GHSOHWHG RI &L 7KHVH H[SHULPHQWV GHPRQVWUDWHG WKDWKLJK &2JURZQFHOOV ZHUH DEOHWR WDNH XS &2 ZKLOH DLUDGDSWHG FHOOV WUDQVSRUWHG ERWK &2 DQG +&2 7KH . IRU WKH WUDQVSRUW RI +&2 ZDV P0 +&2 ZKLOH WKH 9PD[ ZDV SPRO +&2 PJ FKO K 9RORNLWD HW DO 8VLQJPDVVVSHFWURPHWU\(VSLHHWDO WKH&2 WUDQVSRUWHU RISynechococcus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ynechocystis 3&& %pGXHWDO $ F\WRSODVPLFPHPEUDQHDVVRFLDWHGFDUERQLFDQK\GUDVH PD\ DOVR VHUYH WKH LPSRUWDQW IXQFWLRQ RIPLQLPL]LQJ WKH OHDN RIF\WRSODVPLF &2 PROHFXOHV IURP WKH FHOO E\ FRQYHUWLQJ WKHP EDFN WR +&2 )ULGODQG HW DO )XUWKHUPRUH LW ZDV VXJJHVWHG WKDW WKH WUDQVSRUW RI &2 PLJKW RFFXU E\ SDVVLYH GLIIXVLRQ
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ndhB, ndhK DQG ndhL, HQFRGLQJ VXEXQLWV RIWKLV FRPSOH[ UHTXLUH KLJK &2 IRU JURZWK DQG DUH LPSDLUHG LQ &L DFFXPXODWLRQ 0DUFR HW DO 2JDZD SUHVXPDEO\ DV D FRQVHTXHQFH RI LPSDLUHG F\FOLF HOHFWURQIORZ 6XUSULVLQJO\Synechocystis 3&& PXWDQWV LQDFWLYDWHG IRU ndhC 2JDZD DQG ndhF 6FKOXFKWHUHWDO <XHWDO FRXOG JURZ XQGHU ORZ &2 FRQGLWLRQV 7KLV VXJJHVWV WKDW HLWKHU QRW DOO RI WKH VXEXQLWV RI WKH 1$'+ GHK\GURJHQDVH FRPSOH[ DUH HTXDOO\ UHTXLUHG IRU HIILFLHQW F\FOLF HOHFWURQ IORZ DQG JURZWK RQ DLU RU WKDW WKH ndhC DQG ndhF JHQHV DUH IXQFWLRQDOO\ UHGXQGDQW 7KH LPSRUWDQFH RISKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW IRU WKH +&2 XSWDNH LV DOVR VXJJHVWHG E\ WKH ILQGLQJ WKDW D PXWDQW LQ D JHQRPLF UHJLRQ FRQWDLQLQJ WKH psaI/psaL JHQHV ZKLFK HQFRGH VXEXQLWV 9,,, DQG ;, RI 36, LV LPSDLUHG LQ +&2 WUDQVSRUWDQG XQDEOH WR JURZRQ DLU 5RQHQ7DUD]L HW DOE )LQDOO\ D VWURQJ FRUUHODWLRQ EHWZHHQ &L XSWDNH DQG IOXRUHVFHQFH TXHQFKLQJ LQ Synechococcus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nabaena 3&& VXJJHVWHG WKDW 5X%3FDVH DFWLYDVH DQG FKDSHURQLQ DUH DOVR FDUER[\VRPHORFDOL]HG -DJHU DQG %HUJPDQ DOWKRXJK RWKHU VWXGLHV XVLQJ Synechococcus 3&& KDYH ORFDOL]HG 5X%3FDVH DFWLYDVH WR WKH
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hiobacillus neopolitunus WKH FDUER[\VRPHVDUHGHOLQHDWHGE\DSURWHLQDFHRXV VKHOO WKDW DSSHDUV WR EH GHYRLG RIOLSLG +ROWKLM]HQ HW DO WKLV LV SUREDEO\ DOVR WKH FDVH IRU F\DQREDFWHULDOFDUER[\VRPHV 7KH JHQHHQFRGLQJWKH PDMRU FDUER[\VRPH VKHOO SRO\SHSWLGH RI T. neopolitunus ZDV FORQHG DQG WKH GHGXFHG SURWHLQ VHTXHQFH KDV KRPRORJ\ WR WKH SURWHLQV HQFRGHG E\ WKH FDUER[\VRPH UHODWHG JHQHV RI Synechococcus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ynechococcus 3&& ZDV IRXQG WR EH DVVRFLDWHG ZLWK FDUER[\VRPHV 3ULFH HW DO )XUWKHUPRUHSynechococcus 3&&LQ ZKLFKKXPDQ FDUERQLF DQK\GUDVHZDV H[SUHVVHGDV D
'%KD\D 56FKZDU]DQG$5*URVVPDQ VROXEOH F\WRSODVPLF SURWHLQ UHTXLUHG HOHYDWHG OHYHOV RI&2 IRUJURZWK WKLVSKHQRW\SHZDVDFRQVHTXHQFH RID YHU\ KLJK UDWH RI&L OHDNDJH 3ULFH DQG %DGJHU E )LQDOO\ PXWDQWV RI Synechococcus 3&& LPSDLUHG LQ WKH icfA JHQH ZKLFK LV ORFDWHG NESGRZQVWUHDPRIWKH rbc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ynechococcus 3&& ZLWK DOWHUHG FDUER[\VRPHVWUXFWXUHZHUHVKRZQWRKDYHOHVLRQV LQ DQ\ RIILYHGLVWLQFWJHQHV GHVLJQDWHG ccmK, L M, N, O, ZKLFKDUHFOXVWHUHGRQWKHJHQRPHLQWKHUHJLRQRI rbcLS 5RQHQ7DUD]LHWDO D *HQHVFULWLFDOIRU WKH ELRJHQHVLV RI FDUER[\VRPHV ZHUH LVRODWHG IURP Synechocystis 3&& 2QHRIWKHVHJHQHV FFP$ LV XQLTXH ZKLOH WKH RWKHU LV VLPLODU WR WKH ccmM RI Synechococcus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
&L'HSHQGHQW*HQH([SUHVVLRQDQG 6LJQDO7UDQVGXFWLRQ 6HYHUDO F\DQREDFWHULDO JHQHV DUH LQGXFHG XSRQ H[SRVXUHRIF\DQREDFWHULDOFHOOVWRORZOHYHOVRI&2 6RPHRIWKHVHJHQHVDUHH[FOXVLYHO\WUDQVFULEHGLQDQ DWPRVSKHUH RI ORZ &2 WKH JHQHV DUH µVWULQJHQWO\ FRQWUROOHG¶ ZKLOH RWKHUV DUH WUDQVFULEHG LQ ERWK ORZ DQG KLJK DWPRVSKHULF &2 EXW WUDQVFULSW DEXQGDQFH LVHOHYDWHGXQGHUWKHIRUPHUFRQGLWLRQVWKHJHQHVDUH µXSUHJXODWHG¶ 7KH JHQHV WKDW DUH VWULQJHQWO\ FRQWUROOHG LQFOXGH purK HQFRGLQJ VXEXQLW ,, RI SKRVSKRULERV\O DPLQRLPLGD]ROH FDUER[\ODVH DQG cmpA HQFRGLQJ D N'D SURWHLQ LQWHJUDO WR WKH F\WRSODVPLF PHPEUDQHV 'HOHWLRQ DQDO\VLV RI WKH cmpA SURPRWHU LQGLFDWHG WKDW SRVLWLYH DQG QHJDWLYH FLVDFWLQJ HOHPHQWV DUH OLNHO\ WR EH LQYROYHG LQ WKH UHJXODWLRQ E\ DPELHQW &2 .DSODQ 7KH cmpA JHQH LV QRW H[FOXVLYHO\ FRQWUROOHG E\ &2 OHYHOV VLQFH LW WXUQV RXW WR EH LGHQWLFDO WR WKH cbpA JHQH ZKLFK HQFRGHV D SXWDWLYH FDURWHQRLGELQGLQJ SURWHLQ WKDW LV V\QWKHVL]HG GXULQJ H[SRVXUH RI Synechococcus 3&&WRHOHYDWHG OLJKWLQWHQVLWLHV 5HGG\ HW DO +LJK OLJKW DQG ORZ &2 PD\ DIIHFW WKH WUDQVFULSWLRQ RI WKLV JHQH E\ DOWHULQJ WRSRORJ\ RIWKH JHQRPLF '1$ UHDJHQWV WKDW LQKLELW J\UDVH DFWLYLW\ DSSHDU WR SUHYHQW DFWLYDWLRQ RI WKH JHQH LQKLJKOLJKW5HGG\HWDO )XUWKHUPRUH LIORFDOL]HG FKDQJHV LQ '1$ WRSRORJ\ DUH LPSRUWDQW IRU FRQWURO LW PLJKW H[SODLQ ZK\ PDQ\ RIWKH JHQHV WKDW DUH DFWLYDWHG DW ORZ &L FRQFHQWUDWLRQV DUH FOXVWHUHG 7KH JURXS RI JHQHV XSUHJXODWHG LQ DQ DWPRVSKHUH RI ORZ &2 LQFOXGHV ndhB HQFRGLQJ VXEXQLW ,, RI 1$'+ GHK\GURJHQDVH rbcLS DQG WKH ccmNO RSHURQ WKH ODWWHU JHQHV ORFDWHG LPPHGLDWHO\ XSVWUHDP RI rbcLS, HQFRGH SURWHLQV LQYROYHG LQ FDUER[\VRPH ELRJHQHVLV 7KH ccmN JHQH SURGXFW
PD\EHDUHJXODWRU\HOHPHQWVLQFHDSRLQWPXWDWLRQLQ ccm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ynechocystis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hlamydomonas reinhardtii PXWDQW LPSDLUHG LQ WKH FRQYHUVLRQ RI SKRVSKRJO\FRODWH WR JO\FRODWH ZKLFK FDQQRW DFFOLPDWH WR ORZ &L OHYHOV 0DUHNDQG6SDOGLQJ ,QWHUHVWLQJO\ D VLJQLILFDQW LQFUHDVH LQ WKH H[WHQW RI SKRVSKRU\ODWLRQ RI VHYHUDO SRO\SHSWLGHV ZDV QRWHG ZKHQ ORZ&2JURZQ Synechocystis 3&& FHOOV ZHUHVXSSOLHG ZLWKHLWKHUJOXFRVHRU+&2 %OR\H HW DO 7KH QDWXUH RI WKH SKRVSKRU\ODWHG SRO\SHSWLGHV DQG WKHLU SRVVLEOH UROH LQ VXSSUHVVLQJ WKHLQGXFWLRQRIWKH&&0KDVQRWEHHQH[SORUHG 5HFHQWO\ DJHQH GHVLJQDWHG hatR ZDV LGHQWLILHG LQ Synechocystis 3&& ZKLFK ZKHQ LQDFWLYDWHG HOLPLQDWHGWKHLQGXFWLRQRIDKLJKDIILQLW\&L WUDQVSRUW V\VWHP QRUPDOO\ REVHUYHG XSRQ WUDQVIHU RIKLJK &2 JURZQ FHOOV WR DLU OHYHOV RI&2 %pGX HW DO 7KH hatR PXWDQW JUHZ VRPHZKDW VORZHU WKDQ ZLOG W\SH FHOOV LQ DLU VXJJHVWLQJ WKDW WKH &L WUDQVSRUW V\VWHP WKDW FDQQRW EH LQGXFHG LQ WKH hatR PXWDQW LV QRW HVVHQWLDO IRU JURZWK XQGHU WKHVH FRQGLWLRQV EXW PD\ EH UHTXLUHG DW ORZHU &L FRQFHQWUDWLRQV 7KH
GHGXFHG DPLQR DFLG VHTXHQFH RI WKH hatR JHQH SURGXFW H[KLELWV D VLPLODULW\ WR &KH< E. coli DQG 6SR2) Bacillus subtilis VPDOO UHVSRQVH UHJXODWRUV LQYROYHG LQ PRWLOLW\ 5DYLG HW DO
% 1LWURJHQ/LPLWDWLRQ 7KH HQHUJ\ DQG UHGXFWDQW WKDW DUH JHQHUDWHG IURP SKRWRV\QWKHVLV DUH LQ ODUJH SDUW UHTXLUHG IRU WKH DVVLPLODWLRQRIQLWURJHQ1 DQG WKHUHGXFWLRQ RI&2 )ORUHV HW DO +R DQG .URJPDQ 0DQ]DQR HW DO &\DQREDFWHULD FDQ XVH D YDULHW\RILQRUJDQLFDQG RUJDQLFVRXUFHVRI1WRIXOILO WKHLU QXWULWLRQDO UHTXLUHPHQWV 7KH PRVW DEXQGDQW LQRUJDQLF VRXUFHV DUH QLWUDWH DPPRQLXP DQG 1 ZKLOH FRPPRQ RUJDQLF VRXUFHV DUH XUHD DQG FHUWDLQ DPLQRDFLGV +RZHYHUDOOWKHVHVRXUFHVPXVWILUVWEH
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coli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
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO6WUHVV DQG +HUUHUR 6LQJK 5HFHQWO\ &ROOLHU DQG 3DOHQLN FKDUDFWHUL]HG WKHXUHDVH IURP WKH PDULQH6\QHFKRFRFFXV:+ DQGLVRODWHGIRUWKH ILUVWWLPHWKHJHQHVHQFRGLQJXUHDVH 7KLVXUHDVHKDV D . RI 0 DQG D S+ RSWLPXP RI 7KH VWUXFWXUDO JHQHV (ureABC) DQG DFFHVVRU\ JHQHV HQFRGLQJ SURWHLQV UHTXLUHG WR V\QWKHVL]H DQ DFWLYH XUHDVH (ureDEFG), DUH VLPLODU WR WKRVH RI RWKHU EDFWHULD +RZHYHU OLWWOH LV VWLOO NQRZQ DERXW KRZ XUHD XSWDNH DQG DVVLPLODWLRQ DUH UHJXODWHG DOWKRXJK WKH KLJK . RI XUHDVH IRU LWV VXEVWUDWH VXJJHVWV WKDW WKHUH PLJKW EH D XUHD FRQFHQWUDWLQJ PHFKDQLVP ,W KDVUHFHQWO\EHHQVKRZQWKDWSynechococcus VSVWUDLQ 3&&FHOOV JURZQ LQ WKH SUHVHQFH RIP0 XUHD DQG HOHYDWHG &2 OHYHOV XQGHUJR UDSLG FKORURVLV DQG FHOO GHDWK GXULQJ VWDWLRQDU\ SKDVH KRZHYHU D ureC PXWDQW ZKLFK KDV QR XUHDVH DFWLYLW\ GRHV QRWVKRZ WKLV SKHQRW\SH 7KLV SKHQRPHQRQ FRXOG EH PLPLFNHG E\ WKH DGGLWLRQ RI SRO\XQVDWXUDWHG IDWW\ DFLGV RU SDUDTXDW WR WKH PHGLXP ERWK RI ZKLFK LQGXFH VHYHUH R[LGDWLYH VWUHVV 6DNDPRWR HW DO )XUWKHU ZRUN LV UHTXLUHG WR XQGHUVWDQG WKH H[DFW VHTXHQFH RI HYHQWV WKDW WULJJHU H[WHQVLYH FKORURVLV DQG FHOOGHDWK XQGHU FHUWDLQ JURZWK FRQGLWLRQV
E $PLQR$FLGV 7KH DPLQR DFLGV DUJLQLQH JOXWDPLQH DQG DVSDUDJLQH FDQ VHUYH DV 1 VRXUFHV IRU PDQ\ F\DQREDFWHULD )ORUHV DQG +HUUHUR )ORUHV DQG 0XUR3DVWRU )ORUHV DQG 0XUR3DVWRU $UJLQLQH LV WDNHQ XS E\ D WUDQVSRUW V\VWHP VSHFLILF IRU EDVLF DPLQR DFLGV +HUUHUR DQG )ORUHV ,W LV FRQYHUWHG WR XUHD DQG RUQLWKLQH E\ WKH HQ]\PH DUJLQDVH RU WR FLWUXOOLQH DQG DPPRQLXP E\ DUJLQLQH GHLPLQDVH :KLOH VRPH F\DQREDFWHULD OLNH Synechocystis 3&& JURZDV ZHOORQDUJLQLQHDV RQ QLWUDWH RWKHUV OLNH Anabaena 3&& DQG Pseudanabaena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
DO 7KLVLVLQVWULNLQJFRQWUDVWWRWKHVLWXDWLRQ REVHUYHGIRUWKHHQWHULFEDFWHULDZKHUHWKHUHDUHRIWHQ WUDQVSRUW V\VWHPV WDLORUHG IRU D VSHFLILF DPLQR DFLG 7KLV PD\ UHIOHFW D IXQGDPHQWDO GLIIHUHQFH LQ WKH DYDLODELOLW\RIDPLQRDFLGVLQWKHHQWHULFPLOLHXYHUVXV WKDW LQ VRLO RU DTXDWLF HQYLURQPHQWV ,Q WKH KHWHURWURSKLF XQLFHOOXODU Synechocystis 3&& WKUHH DPLQR DFLG WUDQVSRUW V\VWHPV KDYH EHHQ LGHQWLILHG/DEDUUHHWDO 2QH RIWKH V\VWHPV LV VSHFLILF IRU EDVLF DPLQR DFLGV DQG JOXWDPLQH D VHFRQG LV VSHFLILF IRU QHXWUDO DPLQR DFLGV H[FOXGLQJ JOXWDPLQH DQG D WKLUG LV VSHFLILF IRU JOXWDPDWH DQG JOXWDPLQH ,Q WKH KHWHURF\VWIRUPLQJAnabaena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ynechocystis 3&& .DQHNRHW DO 2QH RI WKH JHQHV natA 25) VOU HQFRGHV D SURWHLQ ZLWK KRPRORJ\ WR DQ $%&W\SH SHUPHDVH QXFOHRWLGH ELQGLQJ SURWHLQ ZKLOH WKH VHFRQG natB 25) VOU HQFRGHV D SURWHLQ VLPLODU WR D SHULSODVPLF ELQGLQJ SURWHLQ ,QVHUWLRQDO PXWDQWV RI natA DQG natB ZHUH XQDEOH WR WUDQVSRUW QHXWUDODPLQRDFLGV H[FHSWIRUJOXWDPLQH LQGLFDWLQJ WKDWWKHVHJHQHVHQFRGH FRPSRQHQWV RIQHXWUDO DPLQR DFLGXSWDNHV\VWHPV +RPRORJXHV RInatA DQG natB ZHUH IRXQG LQ DOO WKH F\DQREDFWHULD H[DPLQHG 7KLV ZRUN GHPRQVWUDWHV KRZ WKH LQIRUPDWLRQ DYDLODEOH IURP WKH FRPSOHWH VHTXHQFH RI WKH Synechocystis
3&&JHQRPHRIIHUVWKHSRVVLELOLW\RIFRPELQLQJ D VWURQJ PROHFXODU DSSURDFK ZLWK FODVVLFDO JHQHWLFV DQG SK\VLRORJ\ WR LGHQWLI\ WKH IXQFWLRQV RIPHPEHUV RIVSHFLILFJHQHIDPLOLHV
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nabaena 3&& IRUP VSHFLDOL]HG FHOOV FDOOHG KHWHURF\VWV WKDW GHYHORS DW UHJXODU LQWHUYDOV DORQJ WKH ILODPHQW DQG FRQWDLQWKH1IL[LQJPDFKLQHU\ $XQLTXHFHOO ZDOO WKDW VHUYHV DV D EDUULHU WR DWPRVSKHULF 2 DQG WKH HOLPLQDWLRQ RI2 HYROYLQJ 36,, DFWLYLW\ KHOS SUHYHQW WKH LQDFWLYDWLRQ RI QLWURJHQDVH LQ WKH KHWHURF\VW +DVHONRUQ :RON :RONHWDO ,Q VRPH QRQKHWHURF\VWRXV ILODPHQWRXV F\DQREDFWHULD VXFK DV Trichodesmium, 1 IL[DWLRQ LV GHSHQGHQW RQ WKHSUHVHQFHRIOLJKW2KNLDQG)XMLWD &KDSWHU LPSO\LQJ WKH QHFHVVLW\ IRU SKRWRV\QWKHWLFDOO\ GHULYHG UHGXFWDQW DQG HQHUJ\ DQG H[KLELWV D GLXUQDO UK\WKP:\PDQHWDO 7KHSDUWLDOHOLPLQDWLRQ RI 2 IURP WKH VLWH RI 1 IL[DWLRQ PD\ EH D FRQVHTXHQFH RIFRORQ\ PRUSKRORJ\ DQGRU DQ DELOLW\ RI WKH FHOOV WR HIILFLHQWO\ VFDYHQJH 2 YLD HOHYDWHG UHVSLUDWRU\ DFWLYLW\ 7KH GLIIXVLRQ RI 2 DQG WKH SHQHWUDWLRQ RI OLJKW WR FHOOV QHDU WKH FHQWHU RI D Trichodesmium FRORQ\ PD\ EH H[WUHPHO\ OLPLWHG ZKLFK PD\ KHOS SUHVHUYH QLWURJHQDVH DFWLYLW\ &DUSHQWHUDQG3ULFH +RZHYHU QRW DOO QRQ KHWHURF\VWRXV 1 IL[HUV FDQ IL[ 1 LQ DHURELF RU
' %KD\D 56FKZDU]DQG$5*URVVPDQ PLFURDHURELF FRQGLWLRQV IRU LQVWDQFH Plectonema boryanum IL[HV 1 RQO\ XQGHU FRQGLWLRQV RI DQDHURELRVLV6WHZDUWDQG/H[ 8QLFHOOXODU F\DQREDFWHULD IURP YDULRXV HQYLURQPHQWDO QLFKHV KDYH EHHQ VKRZQ WR IL[ 1 &DUSHQWHUDQG3ULFH +XDQJDQG &KRZ 5HGG\ HW DO :\DWW DQG 6LOYH\ 7KHVH RUJDQLVPV DUH XELTXLWRXV DQG XQLTXH LQ WKHLU DELOLW\ WR IL[ 1 ZLWKLQ DFHOO WKDWPDLQWDLQV DIXQFWLRQDO 2 HYROYLQJ SKRWRV\QWKHWLF DSSDUDWXV ,Q FHUWDLQ VWUDLQV RI Gloeothece DQG Synechococcus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de novo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nifHDK, ZKLFK HQFRGH VWUXFWXUDO FRPSRQHQWV RI WKH QLWURJHQDVH ZDV PDGH VHYHUDO \HDUV DJR 0D]XU HW DO 2QH RI WKH PRVW VWULNLQJ IHDWXUHV RIWKLV UHJLRQ RIWKH Anabaena 3&& JHQRPH ZDV WKH H[LVWHQFH RI D ODUJH LQVHUWLRQ NES LQ WKH FRGLQJ UHJLRQ RI nifD LQ YHJHWDWLYH FHOOV *ROGHQ HW DO +DVHONRUQ :RON HW DO 'XULQJ KHWHURF\VW GHYHORSPHQWWKH NES LQVHUWLRQ LV VSOLFHG IURPWKH JHQRPH FUHDWLQJ D IXQFWLRQDO nifHDK RSHURQ 7KH JHQHHQFRGLQJWKHH[FLVDVHxisA ZKLFKUHPRYHVWKH NESLQVHUWZDVFORQHGDQGVHTXHQFHG/DPPHUVHW DO 6XEVHTXHQWO\ RWKHUH[DPSOHV RIJHQRPLF UHDUUDQJHPHQWV WKDW RFFXU GXULQJ KHWHURF\VW IRUPDWLRQKDYH EHHQFKDUDFWHUL]HG LQAnabaena 3&& Anabaena cylindrica DQG Nostoc 0$&
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO6WUHVV +DVHONRUQ &DUUDVFRHWDO &DUUDVFRDQG *ROGHQ ,QWHUHVWLQJO\WKH NESLQVHUWLRQLV QRW SUHVHQW LQ QRQKHWHURF\VWRXV 1IL[LQJ F\DQREDFWHULD )XMLWD HW DO 5HFHQWO\ LWZDV VKRZQ WKDW LQ DGGLWLRQ WR WKH PDMRU QLWURJHQDVH Anabaena, $7&& V\QWKHVL]HV YDQDGLXP DQG )H QLWURJHQDVH HQ]\PHV WKDW DUH VLPLODU WR WKRVH IRXQG LQ WKH VRLO EDFWHULXP Azotobacter vinelandii .HQWHPLFK HW DO .HQWHPLFK HW DO 7KLHO /LNH RWKHU GLD]RWURSKV F\DQREDFWHULD SUHIHU IL[HG 1VRXUFHV1 IL[DWLRQLVDODVWUHVRUWVLQFHLWUHTXLUHV KLJKOHYHOVRIERWKHQHUJ\DQGUHGXFWDQW 3UHIHUUHG1 VRXUFHV VXFK DV DPPRQLXP RU QLWUDWH LQKLELW WKH H[SUHVVLRQRIWKH 1 IL[DWLRQ V\VWHP SULPDULO\ DW WKH OHYHORIWUDQVFULSWLRQ $PPRQLXPFDQ LQGHSHQGHQWO\ LQKLELW WKH GHYHORSPHQW RI ERWK KHWHURF\VWV LQ Anabaena 3&& DQG QLWURJHQDVH V\QWKHVLV LQ KHWHURF\VW DQG QRQKHWHURF\VWIRUPLQJ 1 IL[HUV 0XOOLJDQ DQG +DVHONRUQ 0XOOLQHDX[ HW DO 5DL HW DO 7KH UHSUHVVLYH HIIHFW RI DPPRQLXP DQG QLWUDWH SUREDEO\ RSHUDWH WKURXJK WKH VDPH SDWKZD\ LH *6 DFWLYLW\ LV UHTXLUHG IRU UHSUHVVLRQ )XMLWD HW DO +XDQJ DQG &KRZ 0DUWLQ1LHWRHWDO 7KH UHJXODWLRQ RI WKH nif genes LQ Azotobacter vinelandii DQG Klebsiella pneumoniae LV ZHOO XQGHUVWRRG DQG LV XQGHU WKH FRQWURO RIWKH nifLA JHQH SURGXFWV 0HUULFN 6RIDUQRKRPRORJXHVRInifLA KDYHEHHQUHSRUWHGLQ F\DQREDFWHULD DOWKRXJK D JHQH HQFRGLQJ D SRO\SHSWLGHZLWK LGHQWLW\WRnifL IURP Klebsiella pneumoniae ZDVQRWHGE\.DQHNRHWDO LQWKH Synechocystis 3&&JHQRPH
LL 3URWHLQ7XUQRYHU 3URWHLQ WXUQRYHU DQ LPSRUWDQW DVSHFW RI WKH F\DQREDFWHULDO UHVSRQVHV WR QXWULHQW GHILFLHQF\ KDV EHHQ H[DPLQHG PRVW WKRURXJKO\ LQ 1 GHILFLHQW FHOOV $ ODUJH SURSRUWLRQ RI FHOOXODU SURWHLQ LV GHJUDGHG LQ Anabaena variabilis GXULQJ WKH GLIIHUHQWLDWLRQ RI KHWHURF\VWV 2ZQE\HWDO 7KLHO %RWK SK\FRELOLSURWHLQV DQG RWKHU FHOOXODU SURWHLQV DUH GHJUDGHGLQ1GHSULYHGSynechococcus DQG Spirulina %RXVVLED HW DO %RXVVLED DQG 5LFKPRQG :\PDQHWDO 7KHDPLQRDFLGVUHOHDVHG IURPWKHGHJUDGDWLRQ RISURWHLQVWKDWDUHQRWHVVHQWLDO GXULQJ 1 OLPLWDWLRQ PD\ SURYLGH VXEVWUDWHV IRU de novo V\QWKHVLV RI SURWHLQV WKDW DUH UHTXLUHG IRU WKH DFFOLPDWLRQ SURFHVV 7KHUH DUH UHSRUWV RI JHQHUDO SURWHDVHV DQG WKRVH WKDW VSHFLILFDOO\ GHJUDGH 3& %RXVVLED DQG 5LFKPRQG (OPRUMDQL DQG
+HUGPDQ )RXOGV DQG &DUU :RRG DQG +DVHONRUQ :RRG DQG +DVHONRUQ
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
DVVLPLODWLRQ LQ F\DQREDFWHULD LV YLD WKH *6*2*$7 V\VWHP0HHNVHWDO
L *6*2*$7 7KH *6 HQ]\PH HQFRGHG E\ glnA, LV DQ DEXQGDQW SURWHLQ LQ WKH F\DQREDFWHULDO FHOO DQG LWV VWUXFWXUH LV VLPLODUWRWKDWRI*6LQRWKHUSURNDU\RWHV ,WUHTXLUHV 0J IRU DFWLYLW\ DOWKRXJK RWKHU GLYDOHQWFDWLRQV FDQ VXEVWLWXWHIRU0J 0EULGDHWDO ,QE. coli, PXOWLOHYHO FRQWUROV GLFWDWH *6 DFWLYLW\ )LUVW LW LV FRQWUROOHG DW WKH WUDQVFULSWLRQDO OHYHO 6HFRQG WKH HQ]\PH DFWLYLW\ LV UHJXODWHG E\ UHYHUVLEOH FRYDOHQW PRGLILFDWLRQ YLD DGHQ\O\ODWLRQ VXFK WKDW WKH GRGHFDPHULF HQ]\PH LV OHVV DFWLYH ZKHQ 1 LV LQ H[FHVV 7KLUG WKHUH LV DOORVWHULF IHHGEDFN LQKLELWLRQ H[HUWHG E\ VHYHUDO GLIIHUHQW VPDOO PROHFXOHV LQFOXGLQJ DODQLQH JO\FLQH WU\SWRSKDQ &73 $03 FDUEDPR\O SKRVSKDWH DQG JOXFRVDPLQH SKRVSKDWH 7KHUHJXODWLRQ RIglnA H[SUHVVLRQ LV GLVFXVVHG LQ WKH QH[WVHFWLRQ,,,%ELL 6RIDUWKHUHLVQRHYLGHQFH WKDW*6LVDGHQ\O\ODWHGLQF\DQREDFWHULDEXWWKHUHDUH REVHUYDWLRQVWKDWFHUWDLQDPLQRDFLGVDQG QXFOHRWLGHV PD\ ELQG *6 DQG LQKLELW LWV DFWLYLW\ %ODQFR HW DO 0EULGD HW DO $PPRQLXP UDSLGO\ LQKLELWV*6LQSynechocystis 3&&EXWDSSHDUVWR KDYH QR HIIHFW RQ WKH *6 RI Anabaena 3&& +RZHYHU ZKHQ WKH *6 RI Anabaena 3&& LV KHWHURORJRXVO\H[SUHVVHGLQSynechocystis 3&& LW LV LQDFWLYDWHG E\ DPPRQLXP 0pULGD HW DO 0EULGD HW DO 7KH PHFKDQLVP E\ ZKLFK DPPRQLXP LQKLELWV WKH F\DQREDFWHULDO *6 LV QRW \HW XQGHUVWRRG 5H\HV DQG )ORUHQFLR E KDYH VXJJHVWHG WKDW *6 LQDFWLYDWLRQ RFFXUV YLD WKH QRQFRYDOHQW ELQGLQJ RI D SKRVSKRU\ODWHG SURWHLQ LQ FHOOV JURZLQJ LQ PHGLXP FRQWDLQLQJ DPPRQLXP %DVHG RQ FURVVOLQNLQJ VWXGLHV WKLV SURWHLQ ZDV FDOFXODWHGWRKDYHDPROHFXODUPDVVRIN'DEXWQR IXUWKHULGHQWLILFDWLRQKDV\HWEHHQPDGH $ VHFRQG JHQH HQFRGLQJ *6 KDV EHHQ IRXQG LQ Synechocystis 3&& 5H\HV DQG )ORUHQFLR 7KH JHQH GHVLJQDWHGglnN, HQFRGHV DSURWHLQ WKDWLVYHU\GLIIHUHQWIURP*6,N'D DQG*6,, N'D WKDW KDYH EHHQ SUHYLRXVO\ LGHQWLILHG LQ SURNDU\RWHV DQG LV FRQVLGHUHG WR FRQVWLWXWH D QHZ JHQH IDPLO\ *6,,, N'D $ glnA PXWDQW RI Synechocystis 3&& ZDV IRXQG WR EH YLDEOH VXJJHVWLQJ WKDW *OQ1 HQFRGHV D IXQFWLRQDO *6 HYHQ WKRXJKLWDFFRXQWVIRURQO\RIWKHWRWDO*6DFWLYLW\ LQZLOGW\SHFHOOV 7KHSUHVHQFHRIDVHFRQG*6PD\ DOVRH[SODLQWKH UHVXOWV RI:DJQHUHWDO ZKR IRXQG WKDW D glnA PXWDQW RI Agmenellum
'%KD\D 56FKZDU]DQG$5*URVVPDQ quadruplicatum (Synechococcus 3&& ZDV YLDEOH DOWKRXJK LWGLG QRWJURZ DV ZHOO DV ZLOGW\SH FHOOV 7KLV VXJJHVWV WKDW *6,,, DFWLYLW\ FDQQRW EH IXOO\ FRPSHQVDWHG IRU E\ *6, ,QWHUHVWLQJO\ LQ Synechocystis 3&& glnN ZDV VWURQJO\ LQGXFHG LQ 1IUHH PHGLXP DQG WKH glnN JHQH DSSHDUHG WR EH SUHVHQW RQO\ LQ QRQ1 IL[LQJ F\DQREDFWHULD 5H\HV DQG)ORUHQFLR 0RUHLQIRUPDWLRQLV QHHGHGWR XQGHUVWDQG WKH UROH RI WKH GLIIHUHQW *6 SURWHLQV LQ F\DQREDFWHULDDQGWKHQDWXUHRIWKHPRGLILFDWLRQVWKDW PD\DOWHUWKHLUDFWLYLWLHV *2*$7 KDV EHHQ SXULILHG IURP Synechococcus 3&& 0DUTXEV HW DO DQG DSSHDUV PRUH VLPLODU WR WKH SODQW HQ]\PH WKDQ WR WKH EDFWHULDO FRXQWHUSDUW LQ WKDW LW FDQ DFFHSW HOHFWURQV IURP UHGXFHG IHUUHGR[LQ ,W LV D VLQJOH SRO\SHSWLGH FRQWDLQLQJ RQH PROHFXOH RI )01 DQG PD\ FRQWDLQ QRQKHPH LURQ 7ZR GLIIHUHQW JHQHV gltB DQG gltS, HQFRGLQJ IHUUHGR[LQGHSHQGHQW *2*$7 KDYH EHHQ LVRODWHG LQ Synechocystis 3&& 1DYDUUR HW DO 7KH GHGXFHG DPLQR DFLG VHTXHQFH RI gltB VKRZV LGHQWLW\ WR WKH DOIDOID 1$'+*2*$7 DQG DERXW LGHQWLW\ WR WKH SODQW IHUUHGR[LQ *2*$7DQGWRWKH EDFWHULDO *2*$7 *OW%DSSHDUV WRKDYHD)H6 FOXVWHUDQGD)01ELQGLQJGRPDLQ ,QVHUWLRQDO LQDFWLYDWLRQ RI HLWKHU gltB RU gltS GHPRQVWUDWHGWKDWQHLWKHUJHQHDORQHZDVHVVHQWLDOIRU YLDELOLW\ KRZHYHU DGRXEOH PXWDQWRIgltB, gltS ZDV QRQYLDEOH
LL 5HJXODWLRQRIJOQ$ ([SUHVVLRQRIglnA LVUHJXODWHGE\WKH1VRXUFHLQWKH PHGLXP VXFK WKDW LW LV ORZHU LQ DPPRQLXP WKDQ LQ QLWUDWH RU1JURZQFHOOV&RKHQ.XSLHFHWDO 0pULGD HW DO :DJQHU HW DO ,Q Synechococcus 3&& D VLQJOH glnA WUDQVFULSW LV LQGXFHG ZKHQ DPPRQLXP LV UHPRYHG IURP WKH PHGLXP LQGXFWLRQGRHV QRWDSSHDUWRUHTXLUH QLWUDWH 0RUHRYHUWKHSUHVHQFHRIDW\SLFDO1WF$ELQGLQJVLWH LQ WKH UHJLRQ RI WKH SURPRWHU VXJJHVWV WKDW WKLV WUDQVFULSWLRQDO DFWLYDWRU LV UHTXLUHG IRU glnA H[SUHVVLRQ 6HFWLRQ ,,,%D ,Q Anabaena 3&& IRXU P51$V ZLWK GLIIHUHQW ¶ HQGV DUH WUDQVFULEHG IURPWKHglnA JHQH 7XUQHUHW DO 7KUHH RI WKHVH WUDQVFULSWV DUH V\QWKHVL]HG LQ DPPRQLXPJURZQ FHOOV ZKLOH RQH LV LQGXFHG GXULQJ 1GHILFLHQWFRQGLWLRQV :KLOH1WF$LVNQRZQWRELQG WR WKH glnA SURPRWHU IXUWKHU DQDO\VLV LV UHTXLUHG WR HVWDEOLVK KRZWKH VSHFLILFLW\ RIWKH WUDQVFULSWLRQ VWDUW VLWHV LV HVWDEOLVKHG 7KHSDWWHUQ RIglnA WUDQVFULSWV LQ Calothrix 3&& DOVR DSSHDUV WR GHSHQG RQ WKH
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO 6WUHVV VRXUFHRI1SUHVHQWLQWKHJURZWKPHGLXP(OPRUMDQL HWDO ,Q Synechocystis 3&& WKH H[SUHVVLRQ RIglnA PD\ EH XQGHU UHGR[ FRQWURO 5H\HV DQG )ORUHQFLR D 7KHUH ZDV D GUDPDWLF GHFUHDVH LQ glnA P51$ LQ FHOOV WUDQVIHUUHG IURP OLJKW WR GDUN RU WUHDWHG ZLWK LQKLELWRUV RI SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW'&08RU'%0,% 7KLVFRQWURO VHHPV WR EHH[HUWHG DWWKH OHYHO RIWUDQVFULSWLRQ VLQFH WKHKDOI OLIH RI WKH glnA WUDQVFULSW ZDV WKH VDPH LQ WKH OLJKW DQG LQ WKH GDUN *6 OHYHOV KRZHYHU GLG QRW IOXFWXDWH VR WKH VLJQLILFDQFH RI WKHVH UHVXOWV LV QRW FOHDU
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xR HWDO 5HFHQW HYLGHQFH IURP2PDWD¶VJURXSLQGLFDWHVWKDWWKHUHPD\DOVREHD GLVWLQFWQLWULWHWUDQVSRUWHU 0DHGDDQG 2PDWD DOWKRXJKLWLV VWLOOXQFKDUDFWHUL]HG 7KHLGHQWLILFDWLRQ RI WUDQVSRUWHUV IRU QLWUDWH DQG QLWULWH LQYROYHG WKH LVRODWLRQ DQG FKDUDFWHUL]DWLRQ RIPXWDQWV LPSDLUHG LQ YDULRXV DVSHFWV RI 1 PHWDEROLVP 0DGXHxR HW DO 2PDWD 2PDWDHWDO /XTXHHWDO 2PDWD 7KH DQDO\VHV RIWKHVH PXWDQWV DOVR OHG WR WKH LGHQWLILFDWLRQ RI JHQHV HQFRGLQJ SRO\SHSWLGHVLQYROYHG LQ1PHWDEROLVP/XTXHHWDO 2PDWD 2PDWDHWDO 6X]XNLHWDO 7R GDWH HLJKW JHQHV IURP WKLV FOXVWHU KDYH EHHQFKDUDFWHUL]HG 6L[ RIWKH JHQHV DUH LQ WKH nirA RSHURQ ZKLFK LQFOXGHV nirA, nrtA, nrtB, nrtC, nrtD DQG narB. $VHFRQGRSHURQ RULHQWHGLQWKH RSSRVLWH GLUHFWLRQ WR WKH nirA RSHURQ FRQWDLQV nirB DQG ntcB 6X]XNL HW DO D 7KH PRVW FRPSOHWH LQIRUPDWLRQLVDYDLODEOHDERXWWKHJHQHVHQFRGLQJWKH QLWUDWH UHGXFWDVH (narB) DQG WKH QLWULWH UHGXFWDVH (nirA) /XTXHHWDO 2PDWDHWDO
L 1LWUDWH DQG 1LWULWH 5HGXFWDVH 1LWULWH UHGXFWDVH 1L5 HQFRGHG E\ nirA, WKH ILUVW JHQH RIWKH nirA RSHURQ LV D SRO\SHSWLGH RI N'D ZKLFK KDV VWURQJ VLPLODULW\ WR SODQW QLWULWH UHGXFWDVH /XTXH HW DO 6X]XNL HW DO 1L5 LV SURSRVHG WR FRQWDLQ DVLURKHPHSURVWKHWLF JURXS DQG D)H6 FOXVWHU ,QPlectonema boryanum, ZKLFK LV D QRQKHWHURF\VWRXV 1IL[LQJ F\DQREDFWHULXP 6X]XNL HW DO E WKH nirA JHQH HQFRGHV D QRYHO QLWULWH UHGXFWDVHLQZKLFKWKHUHLVD&WHUPLQDOH[WHQVLRQWKDW LV DEVHQW LQ WKH Synechococcus 3&& DQG SODQW 1L5V ,QWULJXLQJO\ WKLV H[WUD GRPDLQ KDV VWURQJ KRPRORJ\ WR IHUUHGR[LQ $GGLWLRQDO VWXGLHV DUH UHTXLUHG WR GHWHUPLQH WKH IXQFWLRQ RI WKLV GRPDLQ 6X]XNL HW DO VXJJHVWHG WKDW WKH IHUUHGR[LQOLNH GRPDLQ PD\ EH LQYROYHG LQ WUDQVIHUULQJ HOHFWURQV WR WKH 1L5 FDWDO\WLF GRPDLQ ZKHUH WKH HOHFWURQV FRXOG EH XVHG WR UHGXFH QLWULWH ,W LV DOVR SRVVLEOH WKDW WKH IHUUHGR[LQOLNH GRPDLQ FRQWUROV WKH DFWLYLW\ RI WKH HQ]\PHE\VHQVLQJWKHUHGR[VWDWHRIWKHFHOO 7KLVLV DQ DWWUDFWLYH LGHD VLQFH LW ZRXOG DOORZ IRU D PHFKDQLVPE\ZKLFKWKHFHOOUDSLGO\FRRUGLQDWHV1L5 DFWLYLW\ ZLWK WKH UDWH RI SKRWRV\QWKHVLV DQG &2 IL[DWLRQ 7KH F\DQREDFWHULDO QLWUDWH UHGXFWDVH 15 LV D SRO\SHSWLGH RI DERXW N'D DQG FRQWDLQV PRO\EGHQXPDQG >)H6@ FOXVWHUVZKLFKPDNHLWPRUH VLPLODU WR WKH EDFWHULDO W\SH RI 15 WKDQ WR WKH HXNDU\RWLF 15 ZKLFK FRQWDLQV )$' DQG KHPH DV SURVWKHWLF JURXSV 5HGXFHG IHUUHGR[LQ ZDV VKRZQ WR WUDQVIHUHOHFWURQV WR15 7KLV LV IXUWKHUHYLGHQFHWKDW 1DVVLPLODWLRQ LV FRXSOHG WR SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW)ORUHVDQG +HUUHUR $QGULHVVHHWDO VHTXHQFHG D JHQH GHVLJQDWHG narB, WKDW ZDV VXJJHVWHG WR HQFRGH WKH 15 RI Synechococcus 3&& EDVHGRQ LWV VLPLODULW\ WR RWKHU EDFWHULDO 15V :KHQ narB ZDVH[SUHVVHGLQE. coli XQGHUWKHFRQWURO RI DQ LQGXFLEOH SURPRWHU SURWHLQ H[WUDFWV H[KLELWHG WKH FRQFXUUHQW DFFXPXODWLRQ RID N'DSURWHLQ DQG 15 DFWLYLW\ 5XELR HW DO 7KH FKDUDFWHULVWLFV RIWKLV KHWHURORJRXVO\ H[SUHVVHG 15 ZHUH VLPLODU WR WKRVH RIWKH Synechococcus 3&& HQ]\PH ERWK LQWHUPV RIWKH .P YDOXHDQGUHTXLUHPHQWIRUUHGXFHG IHUUHGR[LQ )LQDOO\ WKH narB JHQH ZDV VKRZQ WR EH WKH WHUPLQDO JHQH RI WKH nirA RSHURQ /XTXH HW DO 6X]XNLHWDO 5HFHQWO\ 6X]XNL HW DO D FKDUDFWHUL]HG D VHFRQG RSHURQ FRQWDLQLQJ WKH JHQHV nirB DQG ntcB, WKDWLV FORVH WR EXWLQ WKHRSSRVLWH RULHQWDWLRQ RIWKH nirA RSHURQ 1LU% LV D SURWHLQ RI DPLQR DFLGV ZLWK QR NQRZQ KRPRORJ\ WR DQ\ SURWHLQV LQ WKH
GDWDEDVH ,QVHUWLRQDO LQDFWLYDWLRQ RI WKH nirB JHQH FDXVHV D GHFUHDVH LQ 1L5 DFWLYLW\ ZKLOH WKH DFWLYLW\ RI 15 DQG *6*2*$7 ZDV RQO\ VOLJKWO\ KLJKHUWKDQLQZLOGW\SHFHOOV 7KHDXWKRUVVXJJHVWHG WKDW 1LU% LV UHTXLUHG IRU PD[LPDO 1L5 DFWLYLW\ DV D FRQVHTXHQFHRIL DFWLYDWLRQRIJHQHVUHTXLUHGIRUWKH DVVHPEO\RI1L5RULWVFRIDFWRUVRULL VWDELOL]DWLRQRI WKH 1L5 SURWHLQ 1WF% ZDV UHFHQWO\ VKRZQ WR EH LQYROYHGLQWKHSRVLWLYHUHJXODWLRQRIWKH nirA RSHURQ 6HFWLRQ%FLLL $LFKLDQG2PDWD
LL 7UDQVSRUW6\VWHPV 7KH FOXVWHU RI JHQHV GRZQVWUHDP RI nirA LQ Synechococcus 3&& LQFOXGHV nrtABCD, ZKLFK HQFRGHFRPSRQHQWVRIWKHQLWUDWHDQG QLWULWHWUDQVSRUW V\VWHPV $nrtA PXWDQWZDVXQDEOHWRJURZRQQLWUDWH DVDVROH1VRXUFH2PDWD 2PDWD 7KH nrtA JHQH SURGXFW LV N'D DQG DSSHDUV WR EH D PHPEUDQHERXQG VXEVWUDWH ELQGLQJ SURWHLQ ,W ELQGV ERWK QLWULWH DQG QLWUDWH ZLWK D .G YDOXH RI DSSUR[LPDWHO\0IRUERWKVXEVWUDWHV0DHGDDQG 2PDWD 0RVW VXEVWUDWHELQGLQJ SURWHLQV RI *UDP QHJDWLYH EDFWHULD DUH VROXEOH SHULSODVPLF SURWHLQVZLWKDKLJKDIILQLW\IRUDSDUWLFXODUVXEVWUDWH ,QWHUHVWLQJO\ 1UW$ PD\ EH D OLSRSURWHLQ WKDW LV DQFKRUHG WR WKH FHOO PHPEUDQH 0DHGD DQG 2PDWD *ORERP\FLQ ZKLFK VSHFLILFDOO\ EORFNV FOHDYDJH RI WKH VLJQDO SHSWLGH RI OLSRSURWHLQ SUHFXUVRUV FDXVHV WKH LQWUDFHOOXODU DFFXPXODWLRQ RI WKH 1UW$ SUHFXUVRU )XUWKHUPRUH PDQ\ RI WKH SXWDWLYH SHULSODVPLF VXEVWUDWH ELQGLQJ SURWHLQV RI Synechocystis VS VWUDLQ 3&& GHULYHG IURP DQDO\VLV RI WKH FRPSOHWH JHQRPH VHTXHQFH KDYH W\SLFDO OLSRSURWHLQ FRQVHQVXV VHTXHQFHV 0DHGD DQG 2PDWD KDYH VXJJHVWHG WKDW DQFKRUHG VXEVWUDWH ELQGLQJSURWHLQVPD\EHDJHQHUDOIHDWXUHRIWUDQVSRUW V\VWHPV LQ FHUWDLQ F\DQREDFWHULD 6XEVWUDWHELQGLQJ OLSRSURWHLQV DUH UDUH LQ *UDP QHJDWLYH EDFWHULD DOWKRXJK LQ *UDP SRVLWLYH EDFWHULD WKH\ DUH FULWLFDO IRUDQFKRULQJSURWHLQV WR WKH F\WRSODVPLF PHPEUDQH 7KH QHFHVVLW\ IRU DQFKRULQJ WKHVH SURWHLQV WR WKH PHPEUDQH LQ Synechococcus 3&& LV SX]]OLQJ VLQFH WKH ELQGLQJSURWHLQV RIF\DQREDFWHULD ZRXOG EH FRQILQHG WR WKH SHULSODVP E\ D FHOO ZDOO DQG WKH SHULSODVPLF VXEVWUDWH ELQGLQJ SURWHLQV RIRWKHU *UDP QHJDWLYHEDFWHULD VXFKDVE. coli, IXQFWLRQHIILFLHQWO\ ZLWKRXWDPHPEUDQHDQFKRU 2WKHU FRPSRQHQWV RI WKH QLWUDWHQLWULWH WUDQVSRUW V\VWHP DUH QRW DV ZHOO FKDUDFWHUL]HG DV 1UW$ 7KH 1UW% SURWHLQ KDV VLPLODULW\ WR WKH LQWHJUDO PHPEUDQH FRPSRQHQWV RI $%& WUDQVSRUWHUV ZKLOH 1UW& DQG
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ynechococcus 3&& 5RGULJXH]HWDO
LLL 5HJXODWLRQ RI*HQHV OQYROYHGLQ 1LWURJHQ $VVLPLODWLRQ $PPRQLXP LQKLELWV QLWUDWHQLWULWH XSWDNH LQ DOO F\DQREDFWHULD H[DPLQHG VR IDU +HUUHUR DQG *XHUUHUR /DUDHWDO 0DUWLQ1LHWRHWDO ZKLFK LVSDUWO\DFRQVHTXHQFHRIUHSUHVVLRQ RI WKH nirA RSHURQ 6X]XNL HW DO 7UDQVFULSWLRQ LQLWLDWLRQIURPnirA LVDOPRVWLPPHGLDWHIROORZLQJWKH VKLIW WR DPPRQLXPIUHH QLWUDWHFRQWDLQLQJ PHGLXP DQG UHTXLUHV WKH DFWLYDWRU 1WF$ 6HFWLRQ ,,,%D ZKLFK ELQGV WR WKH nirA SURPRWHU 7KH WUDQVFULSWV WKDW DUH V\QWKHVL]HG ! NE FRYHU WKH VL[ JHQHV RI WKH RSHURQ DFFXPXODWH WR D PD[LPXP DIWHU DSSUR[LPDWHO\PLQEXWDUHXQVWDEOH DQG PLQ DIWHU WKH LQLWLDO LQFUHDVH WKH\ GURS WR RI WKH PD[LPDOYDOXH 7KLVVXJJHVWVWKDWWUDQVFULSWLRQRIWKH RSHURQ LV FRQWUROOHG E\ IHHGEDFN UHJXODWLRQ WKURXJK WKH DFFXPXODWLRQ RID PHWDEROLWHV JHQHUDWHG E\ 1 DVVLPLODWLRQ 6X]XNL HW DO 7KH nirA WUDQVFULSWVDOVR DFFXPXODWH ZKHQ FHOOV DUHWUDQVIHUUHG IURP DPPRQLXPFRQWDLQLQJ WR 1IUHH PHGLXP LPSO\LQJ WKDW DPPRQLXP DORQH FDQ UHSUHVV WUDQVFULSWLRQ DQG WKDW QLWUDWH LV QRW UHTXLUHG IRU LQGXFWLRQ +RZHYHU .LNXFKLHWDO H[DPLQHG WKHHIIHFWRIQLWUDWHDQGQLWULWHLQ06;WUHDWHGFHOOVLQ ZKLFK WKHUH LV QR QHJDWLYH IHHGEDFN E\ WKH DPPRQLXP JHQHUDWHG LQWHUQDOO\ YLD WKH UHGXFWLRQ RI QLWUDWH 8QGHU WKHVH FRQGLWLRQV QLWULWH DFWLYDWHG WUDQVFULSWLRQ RIWKH nirA RSHURQ 7KLV DFWLYDWLRQKDV EHHQ VKRZQ WR ZRUN WKURXJK QWF% D SURWHLQ RIWKH /\V5IDPLO\HQFRGHGE\WKHQLU%QWF% RSHURQ$LFKL DQG2PDWD 7KHVHUHVXOWVVXJJHVWWKDWSRVLWLYH UHJXODWLRQ RI WKH RSHURQ E\ QLWULWH LV PDVNHG E\ DPPRQLXPUHSUHVVLRQ 7KXV WKHUH DUH OLNHO\WREHD
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO 6WUHVV QXPEHURIIDFWRUVWKDWOHDGWRILQHO\WXQHG H[SUHVVLRQ RIWKHnirA RSHURQ $PPRQLXP GRHV QRW FDXVH UHSUHVVLRQ RIWKH nirA RSHURQ LQ WKH SUHVHQFH RI06; DQ LQKLELWRU RI *6 DFWLYLW\ GHPRQVWUDWLQJ WKDW D PHWDEROLWH GRZQVWUHDP RI DPPRQLXP LQ WKH 1DVVLPLODWLRQ SDWKZD\ LV UHVSRQVLEOH IRU UHSUHVVLRQ 6X]XNL HW DO *OXWDPLQH LV QRW OLNHO\ WR EH WKH UHJXODWRU VLQFH WUHDWPHQW RI WKH FHOOV ZLWK GLD]RR[R/ QRUOHXFLQH '21 ZKLFK LQKLELWV JOXWDPLQH DPLGR WUDQVIHUDVHFDXVLQJWKHOHYHORILQWUDFHOOXODUJOXWDPLQH WR ULVH GRHV QRW UHSUHVV WKH nirA RSHURQ +HQFH D PHWDEROLWH RI JOXWDPLQH PXVW VLJQDO UHSUHVVLRQ DQG HIIRUWV KDYH EHHQ PDGH WR LGHQWLI\ WKLV PROHFXOH 6X]XNLHWDO 7KHPRVWGLUHFWZD\WRLGHQWLI\ FRQWUROOLQJ PROHFXOHV LV WR PRQLWRU WKH UHSUHVVLYH HIIHFWV RI PHWDEROLWHV RQ WKH OHYHO RI WKH nirA WUDQVFULSW $V PHQWLRQHG DERYH ZKHQ nirA LV LQGXFHG LQ WKH DEVHQFH RIDPPRQLXP WKH WUDQVFULSW OHYHO LQFUHDVHV WR DSHDNDQG WKHQGURSV UDSLGO\ WR D YHU\ ORZ OHYHO PDNLQJ LW GLIILFXOW WR HYDOXDWH DQ\ LQKLELWRU\ HIIHFWWKDW D PHWDEROLWH PLJKW KDYH RQ WKH VWHDG\ VWDWH OHYHO RIWKH nirA WUDQFULSW 7R RYHUFRPH WKLVSUREOHP6X]XNLHWDO XVHG DPXWDQW WKDW ZDVGHIHFWLYHLQDFWLYHQLWUDWHWUDQVSRUWDQGWKHUHIRUH ZDV LQ D VWDWH RI DSSDUHQW 1 GHILFLHQF\ $V D FRQVHTXHQFH LW DFFXPXODWHG KLJK OHYHOV RI WKH nirA WUDQVFULSW 7KLV PXWDQW ZDV XVHG WR WHVW YDULRXV PHWDEROLWHV RI 1 PHWDEROLVP IRU WKHLU DELOLW\ WR UHSUHVV nirA, DQG DWWKHVDPHWLPH WR DFWLYDWH rbcLS, ZKLFKZDVVKRZQWREHXSUHJXODWHGE\WKHDGGLWLRQRI DPPRQLXP ,QLWLDOO\ RI WKH PHWDEROLWHV WHVWHG FDUEDPR\O SKRVSKDWH &3 DQG LWV GHFRPSRVLWLRQ SURGXFW F\DQDWH ZHUH IRXQG WR SUHYHQW DFFXPXODWLRQ RIERWK nirA DQG glnA WUDQVFULSWV 7KHVH PHWDEROLWHV DOVR WULJJHUHG DFFXPXODWLRQ RI WKH rbcLS WUDQVFULSW DQG WUDQVFULSWVIURPccmKLMNO DQG icfA, ZKLFKHQFRGH FDUER[\VRPHUHODWHG SURWHLQV DQG FDUERQLF DQK\GUDVHUHVSHFWLYHO\2PDWD (YLGHQFHWKDW F\DQDWHLVWKHPHWDEROLWHUHVSRQVLEOHIRUUHSUHVVLRQRI WKH nirA RSHURQ UDWKHU WKDQ DPPRQLXP ZKLFK DFFXPXODWHV LQ FHOOV DV D FRQVHTXHQFH RI F\DQDWH EUHDNGRZQ ZDV EDVHG RQ H[SHULPHQWV WKDW GHPRQVWUDWHG WKDW HYHQ ZKHQ DPPRQLXP UHDVVLPLODWLRQ ZDV EORFNHG E\ WKH DGGLWLRQ RI06; F\DQDWHVWLOOUHSUHVVHGnirA DQGDFWLYDWHGrbcLS. 7KHSUHVHQFHRIF\DQDVHDFWLYLW\ZDVGHPRQVWUDWHGLQ Synechococcus 87(; Synechococcus VS VWUDLQ 3&&DQGSynechocystis 3&& DQGWKHJHQH HQFRGLQJ F\DQDVH cynS, ZDV VKRZQ WR EH UHSUHVVHG E\DPPRQLXP0LOOHUDQG(VSLH +DUDQRHWDO
6WXGLHV RQ WKHUHJXODWLRQRIcyn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cynS PXWDQW ZKHUH F\DQDWH LV SUHVHQW DW HOHYDWHG OHYHOV WKHUH LV OLWWOHLQKLELWLRQRIWKHDFFXPXODWLRQRInirA WUDQVFULSWV DOWKRXJKDFWLYDWLRQRIRIrbcLS, ccmKLMNO DQG icfA LV VWLOOREVHUYHG 7KHVHGDWDSODFH VRPH GRXEWRQWKH GHWDLOV RI WKH PRGHO DQG VXJJHVW WKDW D QLWURJHQ PHWDEROLWHRWKHUWKDQF\DQDWHPD\ EH LQYROYHGLQWKH FRQWURORInirA H[SUHVVLRQ
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
D 1WF$ 7KHH[LVWHQFH RIDJOREDO FRQWUROOHURI1 DVVLPLODWLRQ ZDV KLQWHG DW E\ WKH LVRODWLRQ RI VHYHUDO SOHLRWURSLF PXWDQWV WKDWZHUH XQDEOHWR JURZ RQ QLWUDWH DQG DOVR ODFNHG H[SUHVVLRQ RI DOO NQRZQ 1UHJXODWHG JHQHV /XTXHHW DO D 9HJD3DODV HWDO 9HJD 3DODV HW DO 7KH ntcA JHQH LVRODWHG E\ FRPSOHPHQWDWLRQ RIWKHVH SOHLRWURSLF PXWDQWV /XTXH
' %KD\D 56FKZDU]DQG$5*URVVPDQ
Fig. 4. 0RGHOIRUWKHUHJXODWLRQRIFDUERQDQGQLWURJHQDVVLPLODWLRQJHQHV 7KHWUDQVSRUWRIDPPRQLXP 1+ QLWUDWH12 ELFDUERQDWH +&2 DQG&2 LQWRWKHFHOOWKURXJK VSHFLILFWUDQVSRUWHUV LV VKRZQ 7KHPHWDEROLFFRQYHUVLRQ RI+&2 WRSKRVSKRJO\FHUDWH3*$ DQG WULRVHSKRVSKDWHWULRVH3 YLDWKHDFWLYLW\RI&$icfA DQG5X%3FDVH (rbcLS) LV VKRZQ7KH1 DVVLPLODWLRQSDWKZD\IURPQLWUDWH12 WR F\DQDWH 1&2 ZLWK WKH LQWHUPHGLDWHV LV VKRZQ 7KH HQ]\PHV LQYROYHG DUH QLWUDWH UHGXFWDVH 15 HQFRGHG E\ narB, QLWULWH UHGXFWDVH 1L5 HQFRGHGE\nirA. $PPRQLXPSOXVJOXWDPDWH*OX LVFRQYHUWHGWRJOXWDPLQH*OQ E\JOXWDPLQHV\QWKHWDVH *6 HQFRGHGE\glnA. *OQLVFRQYHUWHGWRFDUEDPR\OSKRVSKDWHZKLFKPD\GHFRPSRVHWRF\DQDWHZKLFKLQWXUQLVFRQYHUWHGWR&2DQG1+ E\F\DQDVHcynS 7KHSRVLWLYHUHJXODWLRQRIWKHnirA DQG nirB RSHURQV glnA, QWF$DQGF\Q6JHQHVE\1WF$DQGWKHLUSRVVLEOH UHSUHVVLRQE\1&2 LV VKRZQ 7KHSRVLWLYHHIIHFWRI1&2 RQWKH&DVVLPLODWLRQRSHURQV rbcLS, ccmKLMNO DQG icfA LVDOVRVKRZQ 7KHTXHVWLRQPDUNLQGLFDWHVWKDWLW LVQRWNQRZQZKHWKHUDFWLYDWLRQUHSUHVVLRQE\1&2 GLUHFWO\LQYROYHV1WF$ 6HHWH[WIRUTXDOLILFDWLRQRIWKHPRGHO
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
VLWH ZDV ORFDWHG XSVWUHDP RIWKH nirA, glnA DQG nirB WUDQVFULSWLRQ LQLWLDWLRQ VLWHV DQG VXEVWLWXWHV IRU WKH FDQRQLFDO PRWLI 7KHUH DUH WKUHH SRWHQWLDO 1WF$ ELQGLQJ VLWHV QLU, QLU,, DQG QLU,,, V\PPHWULFDOO\ SODFHG LQ WKH ES VHTXHQFH EHWZHHQ WKH nirA DQG nirB FRGLQJ UHJLRQV (nirA DQG nirB DUH RULHQWHG LQ RSSRVLWH GLUHFWLRQ WR HDFK RWKHU 2IWKHVH QLU, DQG QLU,,, DUH OLNHO\ WR EH WKH 1WF$ ELQGLQJ VLWHV DVVRFLDWHG ZLWK WKH nirA DQG nirB RSHURQV UHVSHFWLYHO\ 7KH VLJQLILFDQFH RI QLU,, ORFDWHG EHWZHHQWKH WZR RWKHU VLWHV LV QRW NQRZQ +RZHYHU WKH SUHVHQFH RI PXOWLSOH ELQGLQJ VLWHV VXJJHVWV WKDW
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO 6WUHVV WUDQVFULSWLRQ IURP nirA DQG nirB PD\ EH XQGHU FRPSOH[1PRGXODWHGFRQWURO $ WUDQVFULSWLRQ IDFWRU GHVLJQDWHG %LI$ RU 9), ZDV LGHQWLILHGRQWKHEDVLVRILWVLQWHUDFWLRQVZLWKWKH nifH DQG xisA SURPRWHUV ERWK JHQHV DUH RQO\ H[SUHVVHG GXULQJ KHWHURF\VW GHYHORSPHQW 7KLV SURWHLQZDVDOVRVKRZQWRELQGXSVWUHDPRIWKHrbcLS RSHURQ 5DPDVXEUDPDQLDQ HW DO :HL HW DO %LI$LVQRZNQRZQWREHWKHVDPHSURWHLQDV 1WF$ 7KXV 1WF$VHHPVWRUHJXODWHJHQHVHQFRGLQJ SURWHLQV FULWLFDO IRU 1DVVLPLODWLRQ &DVVLPLODWLRQ DQGKHWHURF\VWGHYHORSPHQW)ULDVHWDO 7KH ntcA JHQH ZDV VKRZQ WR EH PRUH DFWLYH LQ Synechococcus 3&& FHOOV JURZQ LQWKHDEVHQFH RIDPPRQLXP/XTXHHWDO D ,WKDV DW\SLFDO 1WF$ ELQGLQJ VLWH FHQWHUHG DW UHODWLYH WR WUDQVFULSWLRQLQLWLDWLRQ VXJJHVWLQJWKDWLWFDQUHJXODWH LWV RZQH[SUHVVLRQ 7KHntcA JHQHLQAnabaena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ntcB JHQH ZKLFK LV FRWUDQVFULEHG ZLWK WKH nirB JHQHHQFRGHVDSURWHLQRIDPLQRDFLGVZLWK VLPLODULW\WR/\V5WUDQVFULSWLRQDO DFWLYDWRUV $Q ntcB PXWDQW H[KLELWHG DQ LQWULJXLQJ SKHQRW\SH LQ WKDW LW JUHZVORZHUWKDQZLOGW\SH FHOOV RQQLWUDWHRU QLWULWH EXWDVIDVWDVZLOGW\SHFHOOVRQDPPRQLXP6X]XNLHW DO D 7KLV VXJJHVWV WKDW WKH PXWDQW KDV D UHGXFHG DELOLW\ WR DVVLPLODWH QLWUDWH DQG QLWULWH )XUWKHUPRUHWKHntcB PXWDQWVKRZHGVLJQVEDVHGRQ LWV EOHDFKHG DSSHDUDQFH RI EHLQJ 1OLPLWHG HYHQ ZKHQPDLQWDLQHG RQ DPPRQLXP +HQFH 1WF% PD\ SOD\DQLPSRUWDQWUROHLQUHJXODWLQJWKHSDUWLWLRQLQJRI
IL[HG 1 E\ FRQWUROOLQJ FHUWDLQ ELRV\QWKHWLF DFWLYLWLHV XQGHUERWK1VXIILFLHQWDQGGHILFLHQWFRQGLWLRQV
E 3,,SURWHLQ (DUO\VWXGLHV$OOHQHWDO 6DQGHUVHWDO RQ WKH SRWHQWLDO UROH RISKRVSKRU\ODWLRQ RIWK\ODNRLG PHPEUDQH SURWHLQV LQ Synechococcus 3&& UHYHDOHG WKH SUHVHQFH RI WZR VROXEOH SURWHLQV WKDW ZHUHSKRVSKRU\ODWHG XQGHUFRQGLWLRQV RILOOXPLQDWLRQ WKDWSUHIHUHQWLDOO\H[FLWHG 36,, 2QH RIWKHVH SURWHLQV PROHFXODU PDVV N'D ZDV VKRZQ WR KDYH DQ 1 WHUPLQDOVHTXHQFHZLWKVLPLODULW\WRWKH3,,SURWHLQRI E. coli. 7DQGHDX GH 0DUVDF V JURXS LVRODWHG D JHQH IURP Synechococcus 3&& GHVLJQDWHGglnB, WKDW HQFRGHVD3,, KRPRORJXH 7KLVSURWHLQFRQWDLQV DPLQR DFLGV LV LGHQWLFDO WR WKH SURWHREDFWHULDO 3,, SURWHLQ DQG ZDV SUHVHQW LQ DOO F\DQREDFWHULDO VSHFLHVH[DPLQHG7VLQRUHPDVHWDO 3,,SOD\VDFRPSOH[DQGFHQWUDOUROHLQ1FRQWUROLQ HQWHULF EDFWHULD 1LQID HW DO $ ELIXQFWLRQDO XULG\O\O WUDQVIHUDVH HQFRGHG E\ glnD) FDWDO\]HV WKH UHYHUVLEOH XULG\O\ODWLRQ RI 3,, DW D VSHFLILF W\URVLQH UHVLGXH LQ WKH XULG\ODWHG IRUP 3,, VLJQDOV 1 OLPLWDWLRQ 7KH XULG\O\O WUDQVIHUDVH LV VHQVLWLYH WR FKDQJHV LQ WKHUDWLR RIJOXWDPLQH DQG R[RJOXWDUDWH ZKLFK UHIOHFWV WKH 1 VWDWXV RI WKH FHOO 7KXV 3,, IXQFWLRQV DV D VHFRQG PHVVHQJHU DQG LQWHUDFWV ZLWK 1WU%1WU& WKH VHQVRUUHJXODWRU SDLU WKDW FRQWUROV WKH 175 UHJXORQ DQG GLUHFWO\ UHJXODWHV *6 DFWLYLW\ E\ DOWHULQJ LWV DGHQ\O\ODWLRQ VWDWXV YLD WKH DGHQ\OWUDQVIHUDVHHQFRGHG E\ glnE). 3,,KRPRORJXHV ZHUHIRXQG LQ VHYHUDO JURXSV RIEDFWHULD EXW LWV UROH LQUHJXODWLRQ LV QRWDOZD\V WKH VDPH DV WKDWRI3,, LQ WKH HQWHULF EDFWHULD )RU LQVWDQFH LQ WKH IUHHOLYLQJ 1IL[LQJ EDFWHULXP Azospirillum brasilense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glnB 3,, PXWDQW 7KLV PXWDQW H[KLELWHG QRUPDO DFFXPXODWLRQ RI WUDQVFULSWV HQFRGLQJ 1L5 DQG 15 ZKHQ LW ZDV WUDQVIHUUHG IURP DPPRQLXP WR QLWUDWH FRQWDLQLQJ PHGLXP XQOLNH DQ ntcA PXWDQW ZKLFK VKRZV ORZ OHYHO FRQVWLWXWLYH DFFXPXODWLRQ RI WKHVH WUDQVFULSWV 7KXV WKH UROH RI 3,, LQ UHJXODWLQJ 1 DQG & DVVLPLODWLRQDZDLWVIXUWKHUDQDO\VLV
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
' %KD\D 56FKZDU]DQG$5*URVVPDQ ORQJHUWHUP 3 GHSULYDWLRQ WKH\ QHHG WR LQFUHDVH WKHLU FDSDFLW\ WR WDNH XS 3L DQG VFDYHQJH 3 IURP DQ\ DYDLODEOH VRXUFH ZKLFK PD\ UHTXLUH WKH LQGXFWLRQ RI VHYHUDOJHQHV ,Q WKH ODVWIHZ\HDUV WKHUHKDVEHHQD JURZLQJ LQWHUHVW LQ VRPH RIWKH PROHFXODU UHVSRQVHV WR 3 VWUHVV LQ ERWK PDULQH DQG IUHVKZDWHU F\DQREDFWHULD&DUUDQG0DQQ 7KHUHVSRQVHV RI E. coli WR3VWUHVVKDYHEHHQVWXGLHGLQJUHDWGHWDLO DQG SURYLGH D XVHIXO SDUDGLJP IRU H[SORULQJ F\DQREDFWHULDO UHVSRQVHV 7KHUHIRUH D EULHI DFFRXQW RI WKH DFFOLPDWLRQ RI E. coli WR 3 OLPLWDWLRQ LV LQFOXGHGKHUH ,I E. coli LV SODFHG LQ PHGLXP ODFNLQJ 3 JHQHV UHTXLUHG IRU WKH XWLOL]DWLRQ RIDOWHUQDWH 3 VRXUFHV DQG HQFRGLQJ D KLJKDIILQLW\ 3LVSHFLILF WUDQVSRUWHU PD\ EH LQGXFHG E\ PRUH WKDQ IROG 7KHVH JHQHV DUH SDUW RI D 3+2 UHJXORQ WKDW FRPSULVHV JHQHV RUJDQL]HG LQWR HLJKW GLIIHUHQW WUDQVFULSWLRQDO XQLWV :DQQHU 7KH FRQFHSW RI D UHJXORQ LPSOLHV WKH SUHVHQFH RI D PDVWHU UHJXODWRU WKDW FRQWUROV VHYHUDO JHQHV WKDW DUH FRRUGLQDWHO\ DFWLYDWHG LQ UHVSRQVH WR D VLQJOH H[WHUQDO FXH 7KH 3+2 UHJXORQ LQFOXGHV L WKHphn RSHURQ ZKLFK HQFRGHV SURWHLQV WKDW IDFLOLWDWH WKH EUHDNGRZQ DQG XSWDNH RI SKRVSKRQDWHVLL phoA, ZKLFKHQFRGHV DSHULSODVPLF DONDOLQHSKRVSKDWDVHWKDWFOHDYHV3L IURPDYDULHW\ RI RUJDQLF SKRVSKDWHV LLL phoBR, ZKLFK HQFRGHV WKH VHQVRUUHJXODWRUSDLU WKDW FRQWURO H[SUHVVLRQ IURP WKH RSHURQV WKDW FRPSULVH WKH 3+2 UHJXORQ LY phoE, ZKLFK HQFRGHV DSRO\DQLRQLF SRULQ Y WKHpstSCABphoU RSHURQ RI ZKLFK pstSCAB HQFRGHV DQ $%& WUDQVSRUW V\VWHP IRU 3L XSWDNH DQG phoU ZKLFK HQFRGHV D SRO\SHSWLGH LQYROYHG LQ UHSUHVVLRQ RI WKH 3+2 UHJXORQ YL ugpBAECQ, ZKLFK HQFRGHV SURWHLQV UHTXLUHG IRU WKH XSWDNH RI VQJO\FHURO SKRVSKDWH DQGSKRVSKRGLHVWHUDVH (ugpQ) DFWLYLW\ DQG YLL phoH DQGYLLL psiE RIXQNQRZQIXQFWLRQ 7KH PDMRU FRQWURO RI WKH 3+2 UHJXORQ LQYROYHV VHQVLQJ H[WUDFHOOXODU OHYHOV RI 3L E\ WKH 3LVSHFLILF WUDQVSRUW V\VWHP HQFRGHG E\ pst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
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO 6WUHVV VXEMHFWWRFURVVUHJXODWLRQE\WZRRWKHUFHOOVLJQDOLQJ SDWKZD\V :DQQHU D:DQQHU E :DQQHU 7KHVH3LLQGHSHQGHQWFRQWUROV DUHFRXSOHG WR ODWHUVWHSVLQ3PHWDEROLVPDQGPD\FRQVWLWXWHDIRUP RIFURVVWDON WKDW OLQNV FHQWUDO 3 PHWDEROLF SDWKZD\V WR3LXSWDNH
3KRVSKRUXV$FTXLVLWLRQ D 3KRVSKDWH8SWDNH6\VWHPV Synechococcus VS JURZQ LQ 3GHILFLHQW PHGLXP H[KLELWV D IROG LQFUHDVH LQ 9PD[ IRU 3L WUDQVSRUW DOWKRXJK WKH . UHPDLQV XQFKDQJHG LPSO\LQJ WKDW WKHUH LV XSUHJXODWLRQ RI D VLQJOH 3L WUDQVSRUW V\VWHP *ULOORDQG*LEVRQ 0RUH WKDQ RQH WUDQVSRUW V\VWHP PD\ EH RSHUDWLYH LQ Synechococcus VS EXW FKDUDFWHUL]DWLRQV RI WKHVH SXWDWLYH WUDQVSRUWHUV KDYH QRWEHHQSHUIRUPHG +HDOH\ ,QE. coli WKH 3L WUDQVSRUW V\VWHP LV FRPSULVHG RI IRXU SRO\SHSWLGHV 3VW6 WKH SHULSODVPLF 3LELQGLQJ SURWHLQ 3VW& DQG 3VW$ WKH LQWHJUDO PHPEUDQH SURWHLQV DQG 3VW% WKH SHUPHDVHQXFOHRWLGHELQGLQJSURWHLQ 7KLVLVW\SLFDO RIWKH IDPLO\ RI$%& WUDQVSRUWHUV $PHV ,Q F\DQREDFWHULD RQO\WKH JHQHHQFRGLQJ WKH SKRVSKDWH ELQGLQJSURWHLQZDVLGHQWLILHGDQGFKDUDFWHUL]HG3VW6 IURP PDULQH Synechococcus :+ KDV D PROHFXODU ZHLJKW RI DERXW N'D DQG LV LGHQWLWLFDOWRWKHE. coli 3VW6SURWHLQ pstS WUDQVFULSWV DFFXPXODWHG XQGHU 3LGHSOHWHG FRQGLWLRQV &DUU DQG 0DQQ 6FDQODQHWDO ,QWKHIUHVKZDWHU VWUDLQ Synechococcus 3&& $LED DQG 0L]XQR FKDUDFWHUL]HGDQRYHOSURWHLQGHVLJQDWHG6SK; N'D ZKLFK ZDV SDUWLFXODUO\ DEXQGDQW LQ 3 OLPLWHGFHOOV $OWKRXJKWKH\GLGQRWLGHQWLI\WKLVDVD 3L ELQGLQJSURWHLQ VXEVHTXHQWFRPSDUDWLYH DQDO\VHV KDYH VKRZQ WKDW 6SK; LV PRVW OLNHO\ WR EH WKH KRPRORJXH RI3VW6 $LED DQG 0L]XQR 0DQQ DQG6FDQODQ :DJQHUHWDO $QDO\VLVRI WKH HQWLUHSynechocystis 3&& JHQRPH VHTXHQFH KDVUHYHDOHGDQRSHURQWKDWPLJKWHQFRGHPHPEHUVRI WKH 3L WUDQVSRUW V\VWHP .DQHNR HW DO 7KLV RSHURQ FRQWDLQV WZR SVW6OLNH JHQHV RQH ZKLFK HQFRGHV D SRO\SHSWLGH ZLWK JUHDWHU KRPRORJ\ WR WKH PDULQHW\SH 3VW6 DQG WKH RWKHU PRUH VLPLODU WR WKH IUHVK ZDWHUW\SH 3VW6 1 + 0DQQ SHUV FRPP 7KH VLJQLILFDQFH RI WKLV ILQGLQJ DZDLWV ELRFKHPLFDO FKDUDFWHUL]DWLRQRIWKHSXWDWLYH3VW6SRO\SHSWLGHV
E
3KRVSKDWDVHV
0RVW F\DQREDFWHULD DUH XQDEOH WR WDNH XS ODUJH RUJDQLF 3FRQWDLQLQJPROHFXOHV EXWFDQ FOHDYH WKH 3L PRLHW\IURPDYDULHW\RIVXEVWUDWHVXVLQJH[WUDFHOOXODU SKRVSKDWDVHV ZLWK EURDG VXEVWUDWH VSHFLILFLWLHV 6R IDU WKUHH GLVWLQFW SKRVSKDWDVHV KDYH EHHQ LGHQWLILHG LQGLIIHUHQWF\DQREDFWHULDO VSHFLHV 2IWKHVH WKH EHVW FKDUDFWHUL]HG LV DQ DW\SLFDO DONDOLQH SKRVSKDWDVH 3KR$ RI Synechococcus 3&& 7KLV HQ]\PH DFFXPXODWHV LQ 3GHILFLHQWFXOWXUHV RISynechococcus 3&& DQG LV D SHULSODVPLF SURWHLQ ZLWK D PROHFXODUPDVVRIDSSUR[LPDWHO\ N'D%ORFNDQG *URVVPDQ 7KH pho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coli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pho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ynechococcus 3&& :DJQHU HW DO 7KLV HQ]\PH OLNH 3KR$ LV SHULSODVPLFDOO\ ORFDWHG DQG KDV ERWK D EURDG S+
RSWLPXP LQ WKH DONDOLQH UDQJH DQG VXEVWUDWH VSHFLILFLW\IRUSKRVSKRPRQRHVWHUV ,WKDVDPROHFXODU PDVV RI N'D UHTXLUHV =Q IRU DFWLYLW\ DQG H[KLELWV LGHQWLW\ WR WKH DONDOLQH SKRVSKDWDVH 3KR$ IURP Zymomonas mobilis. 3UHOLPLQDU\ HYLGHQFH VXJJHVWVWKDWWKHphoV JHQHLV QRWUHJXODWHG E\ 3 OHYHOV DQG PD\ EH D FRQVWLWXWLYH SHULSODVPLF SKRVSKDWDVH $ PRUH UHFHQW FRPSDULVRQ RI 3KR9 ZLWK SURWHLQV LQ WKH GDWDEDVH VKRZV LWV VWULNLQJ VLPLODULW\ WR D ZHOOGHILQHG &D $73DVH IURP Flavobacterium odoratum 3HLIIHUHWDO 7KH VLJQLILFDQFHRIWKLVVLPLODULW\LVXQFOHDU 3RWWV DQG FRZRUNHUV LGHQWLILHG DQG FORQHG D JHQH ihpP IURP Nostoc commune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
*HQH5HJXODWLRQ'XULQJ3KRVSKRUXV /LPLWDWLRQ 7KH UHJXODWLRQ RI JHQHV WKDW DUH LQGXFHG GXULQJ 3 GHSULYDWLRQKDVEHHQVWXGLHGLQWKHIUHVKZDWHUVSHFLHV Synechococcus 3&& DQG WKH PDULQH XQLFHOOXODU F\DQREDFWHULXP Synechococcus :+ ,Q ERWK VSHFLHV KRPRORJXHV RI WKH WZRFRPSRQHQW VHQVRUUHJXODWRU SDLU 3KR53KR% ZHUH LGHQWLILHG +HWHURORJRXV FRPSOHPHQWDWLRQ RI DQ E. coli VWUDLQ
' %KD\D 5 6FKZDU] DQG $5 *URVVPDQ ODFNLQJ 3KR5 ZLWK D OLEUDU\ RI Synechococcus 3&& '1$ZDVXVHGWRLVRODWHsphS DQG sphR, ZKRVH JHQHSURGXFWVKDYHVLJQLILFDQWKRPRORJ\WR3KR5DQG 3KR% UHVSHFWLYHO\ $LEDHW DO :DWVRQ HW DO 6SK6 DQG 6SK5 DUH DOPRVW FHUWDLQO\ WKH VHQVRUUHJXODWRU SDLU WKDW FRQWUROV WKH UHVSRQVH RI Synechococcus 3&& WR 3 OLPLWDWLRQ 7KLV LV VXSSRUWHG E\ WKH ILQGLQJ WKDW D PXWDQW RI Synechococcus 3&& LQZKLFKWKH sphRS RSHURQ ZDV LQDFWLYDWHGFRXOG QRWLQGXFHSHULSODVPLFDONDOLQH SKRVSKDWDVH DFWLYLW\ LQ UHVSRQVH WR 3 OLPLWDWLRQ QRU ZDV WKHUH WKH LQGXFWLRQ RIWKH DEXQGDQW SHULSODVPLF 3L ELQGLQJ SURWHLQ 7KLV UDLVHV WKH TXHVWLRQ RI ZKHWKHU WKH HTXLYDOHQW RI D 3+2 UHJXORQ H[LVWV LQ F\DQREDFWHULD DQG ZKHWKHU LW LV XQGHU WKH FRQWURO RI WKH 6SK56SK6 VHQVRUUHJXODWRU SDLU ,Q E. coli, DOO 3+2UHJXORQSURPRWHUVH[FHSWRQH KDYHDQ EDVH FRQVHQVXV VHTXHQFH FRPSULVHG RIWZR EDVH GLUHFW UHSHDWVVHSDUDWHGE\EDVHV FDOOHGWKH SKRER[ WKDW UHSODFHV WKH W\SLFDO VHTXHQFH 3KRVSKRU\ODWHG 3KR% ELQGV WR WKLV VHTXHQFH :DQQHU 7KH SKRVSKRU\ODWHG IRUP RI 6SK5 ELQGV WZR VSHFLILF UHJLRQVXSVWUHDPRIWKHphoA SURPRWHUDWSRVLWLRQV WRDQGWR DQGWRWKHSURPRWHURIWKH sphX JHQH HQFRGLQJ WKH SXWDWLYH 3LELQGLQJ SURWHLQ $LED DQG 0L]XQR 1DJD\D HW DO )XUWKHU ZRUN RQ RWKHU 3UHJXODWHG SURPRWHUV ZLOO EH UHTXLUHG WR HVWDEOLVK WKH H[LVWHQFH RI D SRVVLEOH SKR ER[ LQSynechococcus 3&& &KLPHULFJHQHVLQ ZKLFK WKHphoA SURPRWHU IURP WR UHODWLYH WR WKH WUDQVFULSWLRQ VWDUW VLWH ZHUH IXVHG WR WKH uidA JHQHWRFUHDWHDWUDQVODWLRQDOIXVLRQVKRZHG LQGXFLEOH 3JOXFXURQLGDVH DFWLYLW\ *86 ZKHQ WKH FHOOV ZHUH JURZQLQPHGLXPODFNLQJ3 ,QVLWHGLUHFWHGPXWDQWV RIWKLVFRQVWUXFWLQZKLFKEDVHVLQWKHSRVLWLRQV ZHUHDOWHUHGWKHLQGXFLEOH*86 DFWLYLW\ZDV FRPSOHWHO\ ORVW ' %KD\D DQG $ *URVVPDQ XQSXEOLVKHG GDWD VXJJHVWLQJ WKDW WKHUH LV SRVVLEOH VHTXHQFHVSHFLILF ELQGLQJ RI 6SK5 WR WKH phoA SURPRWHU
3RO\SKRVSKDWHV 3RO\SKRVSKDWH JUDQXOHV DUH XVHG E\ F\DQREDFWHULD DQG RWKHU RUJDQLVPV DV LQWUDFHOOXODU3UHVHUYHV ,Q DGGLWLRQ SRO\SKRVSKDWHV PD\ VHUYH D QXPEHU RI IXQFWLRQV LQ PLFURRUJDQLVPV 7KH\ PD\ DFW DV L FKHODWRUV LL EXIIHUV DJDLQVW DGYHUVH FRQGLWLRQV VXFK DV DONDOLQH HQYLURQPHQWV KHDW DQG R[LGDWLYH VWUHVV LLL UHJXODWRUV RIWUDQVFULSWLRQ DQG LY PROHFXOHV WKDW DOORZ VXUYLYDO GXULQJ VWDWLRQDU\ SKDVH .RUQEHUJ &KDSWHU
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
RYHUSOXV SKHQRPHQRQ UHVXOWV LQ DQ LQLWLDO PDVVLYH DFFXPXODWLRQ RI SRO\SKRVSKDWH JUDQXOHV ZKLFK DUH VXEVHTXHQWO\GHJUDGHG DV WKH FHOOVUHWXUQWR QXWULHQW UHSOHWH H[SRQHQWLDO JURZWK 7KH DELOLW\ WR RYHUDFFXPXODWH SRO\SKRVSKDWH PD\ DOORZ F\DQREDFWHULDWRWDNHDGYDQWDJHRISXOVHVRIDYDLODEOH QXWULHQWV WKDW DUH FRPPRQ LQ SDWFK\ QXWULHQWSRRU HQYLURQPHQWV ,W KDV DOVR EHHQ VXJJHVWHG WKDW WKH EXR\DQF\ RI PDULQH Trichodesmium PD\ EH FRQWUROOHG LQ SDUW E\ GLXUQDO FKDQJHV LQ SRO\SKRVSKDWH FRQWHQW 5RPDQV HW DO 6HH &KDSWHU 7KH LGHQWLILFDWLRQ RI SXWDWLYH JHQHV LQ Synechocystis 3&& HQFRGLQJ D SRO\SKRVSKDWH NLQDVH DQG D SRO\SKRVSKDWDVH .DQHNR HW DO VKRXOG OHDG WR IXUWKHU DQDO\VLV RI WKH UROH RI SRO\SKRVSKDWHV LQ WKH SK\VLRORJ\ DQG PHWDEROLVP RI WKHFHOO
' 6XOIXU/LPLWDWLRQ &\DQREDFWHULD WKDW H[SHULHQFH VXOIXU OLPLWDWLRQ FHDVH JURZWK DOWHU SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW GHJUDGH WKHLU 3%6 DQG LQFUHDVH WKH HIILFLHQF\ ZLWK ZKLFK WKH\ WUDQVSRUW VXOIDWH DQG RWKHU 6FRQWDLQLQJ PROHFXOHV ,QGHHG Synechococcus 3&& LV DEOH WR XVH D QXPEHU RI RUJDQLF DQG LQRUJDQLF 6 FRPSRXQGV LQFOXGLQJ VXOIDWH WKLRVXOIDWH F\VWHLQH F\VWLQH UHGXFHG JOXWDWKLRQH DQG WKLRF\DQDWH /DXGHQEDFKDQG*URVVPDQ /DZU\DQG-HQVHQ 6FKPLGW HW DO ,Q WKLV VHFWLRQ WKH WUDQVSRUW RI VXOIDWH LQWR F\DQREDFWHULDO FHOOV LV FRQVLGHUHG VLQFH ERWK WKH FKDQJH LQ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUWDQG WKHGHJUDGDWLRQRIWKH3%6 DUH GLVFXVVHGHOVHZKHUHLQWKLVUHYLHZ6HFWLRQ,,,'
6XOIDWH8SWDNH 6XOIDWH XSWDNH LV DQ DFWLYH SURFHVV GHSHQGHQW XSRQ OLJKW DQG YDULHV ZLWK ERWK WHPSHUDWXUH DQG S+ LV
VHQVLWLYH WR RVPRWLF VKRFN DQG LV LQKLELWHG E\ WKLRVXOIDWH DQG VXOILWH -HDQMHDQ DQG %URGD 8WNLOHQ HW DO 8WNLOHQ DQG FRZRUNHUV UHSRUWHG D . RI 0 DQG D 9PD[ RI PROFHOOV [ PLQ IRUVXOIDWH WUDQVSRUWDW& LQ 6 VWDUYHG Synechococcus. 6LPLODU SK\VLRORJLFDO VWXGLHV *UHHQ DQG *URVVPDQ GHPRQVWUDWHG WKDW ZKLOH WKH 9PD[ IRU VXOIDWH WUDQVSRUW LQFUHDVHG EHWZHHQ DQG IROG GXULQJ 6GHSULYDWLRQ WKH . ZDVDSSUR[LPDWHO\ 0LQERWK6 VXIILFLHQWDQG 6GHILFLHQWPHGLXP 7KLVVXJJHVWHGWKDW6GHILFLHQF\ FDXVHG HOHYDWHG DFFXPXODWLRQ RI D VLQJOH VXOIDWH WUDQVSRUW V\VWHP 2QFH VXOIDWH LV WUDQVSRUWHG LQWR Synechococcus FHOOV LW LV UHGXFHG YLD WKH 3$36 VXOIRWUDQVIHUDVH SDWKZD\ 6FKPLGW DQG &KULVWHQ 2WKHUF\DQREDFWHULDFDQUHGXFHVXOIDWHYLDWKH $36 VXOIRWUDQVIHUDVH SDWKZD\ 7VDQJ DQG 6FKLII
*HQH5HJXODWLRQ 8SRQ 6XOIXU/LPLWDWLRQ $ UHJLRQ RI WKH Synechococcus 3&& JHQRPH NE ZDV LGHQWLILHG WKDW FRQWDLQHG JHQHV LQYROYHG LQVXOIDWHDFTXLVLWLRQ*UHHQHWDO /DXGHQEDFK HW DO /DXGHQEDFK DQG *URVVPDQ $ ODUJHLQFUHDVHLQWKHOHYHO RIP51$WUDQVFULEHG IURP WKHVH JHQHV RFFXUV ZKHQ WKH FHOOV DUH GHSULYHG RI6 6HTXHQFH FKDUDFWHUL]DWLRQ RI WKLV UHJLRQ RI WKH F\DQREDFWHULDOJHQRPHUHYHDOHGWKHSUHVHQFHRIJHQHV HQFRGLQJ IRXU FRPSRQHQWV RI D W\SLFDO SHULSODVPLF WUDQVSRUWV\VWHPcysA, CYST, F\V:DQGsbp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sbpA GLG QRWSUHYHQWWKHFHOOVIURPXWLOL]LQJ VXOIDWH DV WKH VROH 6 VRXUFH 7KHVH UHVXOWV VXJJHVWHG WKDW HLWKHU 6ES$ ZDV QRW DEVROXWHO\ UHTXLUHG IRU
JURZWK RQ VXOIDWH RU WKDW D VHFRQG JHQH H[LVWHG ZKRVHJHQHSURGXFWFRXOGVXEVWLWXWHIRU6ES$ 7KH VXOIDWH SHUPHDVH PD\ DOVR EH XWLOL]HG IRU WKH XSWDNHRIWKLRVXOIDWH 7KLRVXOIDWH WUDQVSRUWPD\ DOVR EH DXJPHQWHG E\ WKH ELQGLQJ RI WKLRVXOIDWH WR D VSHFLILF VXEVWUDWHELQGLQJSURWHLQ GHVLJQDWHG &\V3 LQ E. coli +U\QLHZLF]HWDO %DVHGRQKRPRORJ\ WR WKHE. coli JHQH WKH cysP JHQH LQ Synechococcus 3&& LV LPPHGLDWHO\ GRZQVWUHDPRIcysW. 7KH SXWDWLYH cysP JHQHLVIROORZHGE\WZRJHQHVWKDWKDYH KRPRORJLHVWRcysT DQGF\V: 7KHVH PD\ EH SDUW RI D WUDQVSRUW V\VWHP WKDW KDV D KLJKHU VSHFLILFLW\ IRU WKLRVXOIDWHWKDQIRUVXOIDWH 7KHUHGRHV QRWDSSHDUWR EH D VHSDUDWH QXFOHRWLGH ELQGLQJ SURWHLQ DVVRFLDWHG ZLWK WKLV V\VWHP VR &\V$ PD\ IXQFWLRQ IRU ERWK WUDQVSRUWV\VWHPV 'RZQVWUHDP RI WKH SXWDWLYH WKLRVXOIDWH WUDQVSRUW JHQHV LV D JHQH GHVLJQDWHG rhdA, ZKLFK HQFRGHV D SURWHLQZLWKVRPHVLPLODULW\WRWKHHQ]\PHUKRGDQHVH ZKLFK WUDQVIHUV WKH WKLRO JURXS RI WKLRVXOIDWH WR WKLRSKLOLF DFFHSWRU PROHFXOHV /DXGHQEDFK HW DO 7KLVSURWHLQDFFXPXODWHVWRKLJKOHYHOVLQWKH SHULSODVPLF VSDFH XSRQ 6 VWDUYDWLRQ +RZHYHU LQDFWLYDWLRQ RI rhdA GRHV QRW SUHYHQW XWLOL]DWLRQ RI YDULRXV LQRUJDQLF 6 VRXUFHV 5KG$ PD\ KDYH D FDWDO\WLF UROH LQ DFFOLPDWLRQ RU PD\ VLPSO\ ELQG FHUWDLQ 6FRQWDLQLQJ FRPSRXQGV DQG SUHYHQW WKHLU OHDNDJHIURPWKHFHOO
&\V5 $JHQH ORFDWHG LQWKHPLGGOH RIWKH VXOIDWHSHUPHDVH RSHURQ GHVLJQDWHG cysR, HQFRGHV D SURWHLQ WKDW UHVHPEOHVWUDQVFULSWLRQDOUHJXODWRUV VXFKDV)L[.DQG &53 /DXGHQEDFK DQG *URVVPDQ :KHQ WKH cysR JHQH LV LQWHUUXSWHG WKLRF\DQDWH FDQ QR ORQJHU VHUYHDVDVROH6VRXUFHDOWKRXJKRWKHU6 VRXUFHVFDQ VXSSRUWJURZWK RIWKHcysR PXWDQW 5HFHQWO\ LWZDV IRXQG WKDW &\V5 ZDV LQYROYHG LQ WKH UHJXODWLRQ RI VHYHUDO JHQHV RQ WKH S$1/ SODVPLG RI Synechococcus. 7KHVH JHQHV HQFRGH D N'D SHULSODVPLF SURWHLQ 6US$ ZKLFK LV VLPLODU WR FDWDODVH HQ]\PHV LQYROYHG LQ F\VWHLQH ELRV\QWKHVLV 6US* DQG 6US+ DQG D SURWHLQ &KU$ ZLWK UHPDUNDEOHKRPRORJ\WRWKHFKURPDWHLQGXFLEOHchrA JHQH SURGXFW RI Alcaligenes eutrophus. 7KH chrA JHQH SURGXFW PD\ EH LQYROYHG LQ WUDQVSRUW RU HIIOX[ RIKHDY\ PHWDOV VXFK DV FKURPDWH 7KLV SURYLGHV WKH ILUVW LQGLFDWLRQ RI WKH UROH RI WKH KLWKHUWR FU\SWLF SODVPLGV RI F\DQREDFWHULD 1LFKROVRQ HW DO 1LFKROVRQ DQG /DXGHQEDFK DQG VXJJHVWV
'%KD\D56FKZDU]DQG$5*URVVPDQ WHPSRUDO FRQWURO RI JHQH UHJXODWLRQ GXULQJ 6 OLPLWDWLRQ )ROORZLQJVXOIXUGHSULYDWLRQWKHJHQHVHQFRGLQJWKH VXOIDWHDQG WKLRVXOIDWHWUDQVSRUWV\VWHPV DQG cysR DUH DFWLYDWHG WKH DFWLYDWRU RI WKH RSHURQV HQFRGLQJ WKH VXOIDWH DQG WKLRVXOIDWH WUDQVSRUW V\VWHPV KDYH QRW EHHQ LGHQWLILHG ,QFUHDVHG V\QWKHVLV RI WKH &\V5 SURWHLQ FDXVHV WKH DFWLYDWLRQ RI QXPHURXV JHQHV RQ WKH ODUJH Synechococcus 3&& SODVPLG )RU VRPHRIWKHSODVPLG ORFDOL]HG JHQHVWKHIXQFWLRQZDV HVWDEOLVKHG KRZHYHU RWKHUV HQFRGH SURWHLQV ZLWK LQWULJXLQJ VLPLODULWLHV WR SURWHLQV WKDW KDYH GHILQHG FDWDO\WLF IXQFWLRQ EXW WKH UROH WKDW WKH\ SOD\ LQ WKH UHVSRQVH RI F\DQREDFWHULD WR VXOIXU OLPLWDWLRQ LV QRW FOHDU
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
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO6WUHVV ZRXOG SURYLGH HQHUJ\ WR GULYH PHWDEROLF SURFHVVHV WKDW PXVW EH PDLQWDLQHG LQ QXWULHQW GHILFLHQW FHOOV 7KLV HQHUJ\ ZRXOG DOVR EH UHTXLUHG IRU WUDQVSRUWLQJ WKHOLPLWLQJQXWULHQWLQWRWKHFHOOZKHQLWGRHVEHFRPH DYDLODEOH 7KH OHYHOV RI WUDQVFULSWV HQFRGLQJ VSHFLILF FRPSRQHQWV RI WKH SKRWRV\QWKHWLF DSSDUDWXV ZHUH VKRZQ WR GHFOLQH GXULQJ QXWULHQW OLPLWDWLRQ ,Q Synechococcus 3&& WUDQVFULSWV IURP WKH cpcBACDEF RSHURQZHUHHVVHQWLDOO\XQGHWHFWDEOH 5 K DIWHU WKH LQLWLDWLRQ RI 1 GHSULYDWLRQ %U\DQW GH /RULPLHU HW DO 'DWD REWDLQHG E\ *DVSDULFK HW DO XVLQJ D cpcB-lacZ FKLPHULF JHQH VXJJHVWHG WKDW 1 GHSOHWLRQ FDXVHG D PDUNHG GHFUHDVH LQ WUDQVFULSWLRQ RI WKH cpcBA RSHURQ ,Q Synechococcus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nabaena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
$73 E\ F\FOLF HOHFWURQ IORZ LQ KHWHURF\VWV ZRXOG KHOS VXSSRUW WKH HQHUJHWLFDOO\ H[SHQVLYH SURFHVV RI 1 IL[DWLRQ
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
PD\ DOVR KHOS SURWHFW WKH FHOOV IURP SKRWRWR[LFLW\ GLVFXVVHG DERYH DOWKRXJK WKHUH PD\ EH RWKHU PHFKDQLVPV WKDW GUDPDWLFDOO\ UHGXFH WKH WUDQVIHU RI HQHUJ\IURPWKH3%6WRFKO 7KHGHJUDGDWLRQRIWKH3%6 RI6\QHFKRFRFFXV3&& LVYLUWXDOO\LGHQWLFDOGXULQJ1DQG6GHSULYDWLRQ ,WLV DQ RUGHUHG SURFHVV LQ ZKLFK GHJUDGDWLRQ EHJLQV DWWKHSHULSKHU\ RIWKH FRPSOH[ HOLPLQDWLQJ WKH FRUH GLVWDO 3& KH[DPHUV ZLWK VXEVHTXHQW HOLPLQDWLRQ RI KH[DPHUV FORVHU WR WKH FRUH &ROOLHU DQG *URVVPDQ 'XNH HW DO
' %KD\D 56FKZDU]DQG$5*URVVPDQ SUHVHQW RQ WKH JHQRPHV RI RWKHU F\DQREDFWHULD 6\QHFKRF\VWLV3&&6\QHFKRFRFFXV3&& F. GLSORVLSKRQ DQG UHG DOJDH XQSXEOLVKHG GDWD $QDO\VLV RIWKH JHQRPH RI6\QHFKRF\VWLV 3&& UHYHDOHG WZR nbl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nbl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
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO6WUHVV QRWNQRZQ DOWKRXJKWKHUHGR[VWDWHRIWKHFHOO ZRXOG EH D FRPPRQ OLQN DPRQJ WKH FXHV VHQVHG E\ WKLV UHVSRQVHUHJXODWRU
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ynechocystis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
$FNQRZOHGJPHQWV :HDUHYHU\JUDWHIXO WR $DURQ .DSODQ 'DYLG .HKRH DQG7DWVXR2PDWDIRUDOORZLQJXVWRXVHDQG PRGLI\ WKHILJXUHVSUHVHQWHGKHUH :HZRXOGOLNHWRWKDQN( )ORUHV 6 6 *ROGHQ $ .DSODQ 1 + 0DQQ 7 2PDWD 6 7DEDWD IRU VKDULQJ XQSXEOLVKHG GDWD SUHSULQWV DQG KHOSIXO GLVFXVVLRQV .DWKL %XPS ZDV LQYDOXDEOHLQW\SLQJDQGIRUPDWWLQJWKLVFKDSWHU :H DFNQRZOHGJH VXSSRUWIURP1,+ 86'$16)DQG WKH +XPDQ)URQWLHUV3URJUDPIRUPRUHWKDQWZRGHFDGHV RIVXSSRUW 7KLVLV&,:SXEOLFDWLRQ1R
5HIHUHQFHV $LED + DQG 0L]XQR 7 $ QRYHO JHQH ZKRVH H[SUHVVLRQ LV UHJXODWHG E\ WKH UHVSRQVHUHJXODWRU 6SK5 LQ UHVSRQVH WR SKRVSKDWH OLPLWDWLRQ LQ Synechococcus VSHFLHV 3&& 0RO 0LFURELRO $LED + 1DJD\D 0 DQG 0L]XQR 7 6HQVRU DQG UHJXODWRU SURWHLQV IURP WKH F\DQREDFWHULXP Synechococcus VSHFLHV 3&& WKDW EHORQJ WR WKH EDFWHULDO VLJQDOWUDQVGXFWLRQ SURWHLQ IDPLOLHV LPSOLFDWLRQ LQ WKH DGDSWLYH UHVSRQVH WR SKRVSKDWH OLPLWDWLRQ0RO0LFURELRO $LFKL0DQG2PDWD7 ,QYROYHPHQWRI1WF% D/\V5IDPLO\ WUDQVFULSWLRQ IDFWRU LQ QLWULWH DFWLYDWLRQ RI WKH QLWUDWH DVVLPLODWLRQ RSHURQ LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ3&&-%DFWHULRO $L]DZD . DQG 0L\DFKL 6 &DUERQLF DQK\GUDVH DQG &2 FRQFHQWUDWLQJ PHFKDQLVP LQ PLFURDOJDH DQG F\DQREDFWHULD )(060LFURELRO5HY $L]DZD.6KLPL]X7+L\DPD76DWRK.1DNDPXUD<DQG)XMLWD < &KDQJHV LQ FRPSRVLWLRQ RI PHPEUDQH SURWHLQV DFFRPSDQ\LQJWKHUHJXODWLRQRI36,36,,VWRLFKLRPHWU\REVHUYHG ZLWK Synechocystis 3&&3KRWRV\Q5HV $OOHQ -) 3URWHLQ SKRVSKRU\ODWLRQ LQ UHJXODWLRQ RI SKRWRV\QWKHVLV%LRFKLP%LRSK\V$FWD $OOHQ -) 6DQGHUV &( DQG +ROPHV 1* &RUUHODWLRQ RI PHPEUDQH SURWHLQ SKRVSKRU\ODWLRQ ZLWK H[FLWDWLRQ HQHUJ\ GLVWULEXWLRQ LQ WKH F\DQREDFWHULXP Synechococcus )(%6 /HWW $OOHQ 0 0 &\DQREDFWHULDO FHOO LQFOXVLRQV $QQX 5HY 0LFURELRO $PHV *)/ %DFWHULDO SHULSODVPLF WUDQVSRUW V\VWHPV VWUXFWXUH PHFKDQLVP DQG HYROXWLRQ $QQX 5HY %LRFKHP $QGHUVVRQ % DQG 6W\ULQJ 6 3KRWRV\VWHP ,, 0ROHFXODU RUJDQL]DWLRQ IXQFWLRQ DQG DFFOLPDWLRQ &XUU7RS%LRHQHUJHWLFV $QGULHVVH; YDQ$UNHO*DQG:HLVEHHN3 &ORQLQJRIWKH QLWUDWH UHGXFWDVH JHQH RI Synechococcus 3&& ,Q 7KLUG ,QWHUQDWLRQDO 6\PSRVLXP RQ 12$VVLPLODWLRQ 0ROHFXODU DQG *HQHWLF$VSHFWV%RPEDQHV $SSOHE\ -/ 3DUNLQVRQ -6 DQG %RXUUHW 5% 7KH PXOWLVWHS SKRVSKRUHOD\1RWQHFHVVDULO\DURDGOHVVWUDYHOHG&HOO $UJDOO 0( 6PLWK *' 6WDPIRUG 13- DQG
SK\VLRORJLFDO DQG JHQHWLF HYLGHQFH IRU D KLJKDIILQLW\ XSWDNHV\VWHP0RO0LFURELRO %HONQDS:5DQG+DVHONRUQ5 &ORQLQJDQGOLJKWUHJXODWLRQ RIH[SUHVVLRQ RIWKH SK\FRF\DQLQ RSHURQ RIWKH F\DQREDFWHULXP Anabaena. (0%2- %HQQHWW - 3URWHLQ SKRVSKRU\ODWLRQ LQ JUHHQ SODQW FKORURSODVWV$QQX5HY3ODQW3K\VLRO %KD\D ' 0ROHFXODU UHVSRQVHV RI F\DQREDFWHULD WR PDFURQXWULHQWOLPLWDWLRQ-6FL,QG5HV %LJJLQV - DQG %UXFH ' 5HJXODWLRQ RI H[FLWDWLRQ HQHUJ\ WUDQVIHULQRUJDQLVPV FRQWDLQLQJSK\FRELOLQV 3KRWRV\Q 5HV %ODQFR ) $ODQD $ /ODPD 0- DQG 6HUUD -/ 3XULILFDWLRQ DQG SURSHUWLHV RIJOXWDPLQH V\QWKHWDVH IURP WKH QRQ1IL[LQJ F\DQREDFWHULXP Anabaena laminosum. - %DFWHULRO %ORFN 0$ DQG *URVVPDQ $5 ,GHQWLILFDWLRQ DQG SXULILFDWLRQ RI D GHUHSUHVVLEOH DONDOLQH SKRVSKDWDVH IURP Anacystis nidulans 5 3ODQW3K\VLRO %OR\H6$6LOPDQ1-0DQQ1+DQG&DUU1* %LFDUERQDWH FRQFHQWUDWLRQ E\ Synechocystis 3&& 3ODQW 3K\VLRO %RJRUDG / 3K\FRELOLSURWHLQV DQG FRPSOHPHQWDU\ FKURPDWLFDGDSWDWLRQ$QQX5HY3ODQW3K\VLRO %RXVVLED 6 $PPRQLD XSWDNH LQ WKH DONDORSKLOLF F\DQREDFWHULXP Spirulina platensis. 3ODQW&HOO3K\VLRO %RXVVLED65HVFK&0DQG*LEVRQ- $PPRQLDXSWDNHDQG UHWHQWLRQLQVRPHF\DQREDFWHULD$UFK0LFURELRO %RXVVLED6 DQG5LFKPRQG$( &SK\FRF\DQLQDV DVWRUDJH SURWHLQ LQ WKH EOXHJUHHQ DOJD Spirulina platensis. $UFK 0LFURELRO %URXQ ,, )DGH\HY 6, .LULN ,, 6HYHULQD ,, DQG 6NXODFKHY 93 /LJKWGHSHQGHQWGHOWD1DJHQHUDWLRQ DQG XWLOL]DWLRQ LQ WKHPDULQHF\DQREDFWHULXP Oscillatoria brevis. )(%6 /HWW %UXQV % %ULJJV :5 DQG *URVVPDQ $5 0ROHFXODU FKDUDFWHUL]DWLRQ RI SK\FRELOLVRPH UHJXODWRU\ PXWDQWV LQ Fremyella diplosiphon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
'%KD\D 56FKZDU]DQG$5*URVVPDQ %XVE\ 6 DQG (EULJKW 5+ 3URPRWHU VWUXFWXUH SURPRWHU UHFRJQLWLRQ DQGWUDQVFULSWLRQDFWLYDWLRQLQSURNDU\RWHV &HOO %XVWRV 6$ 6FKDHIHU 05 DQG *ROGHQ 66 'LIIHUHQW DQG UDSLG UHVSRQVHV RI IRXU F\DQREDFWHULDO psbA WUDQVFULSWV WR FKDQJHVLQOLJKWLQWHQVLW\-%DFWHULRO &DPSEHOO ' &ODUNH $. *XVWDIVVRQ 3 DQG 2TXLVW * ' H[FKDQJH DQG WKH 3KRWRV\VWHP ,, UHSDLU F\FOH LQ WKH F\DQREDFWHULXP Synechococcus. 3ODQW6FLHQFH &DPSEHOO ' =KRX * *XVWDIVVRQ 3 2TXLVW * DQG &ODUNH $. (OHFWURQ WUDQVSRUW UHJXODWHV H[FKDQJH RI WZR IRUPV RI SKRWRV\VWHP ,, ' SURWHLQ LQ WKH F\DQREDFWHULXP Synechococcus. (0%2- &DUSHQWHU ( DQG 3ULFH && 0DULQH Oscillatoria (Trichodesmium): ([SODQDWLRQ IRU DHURELF QLWURJHQ IL[DWLRQ ZLWKRXWKHWHURF\WV6FLHQFH &DUUDVFR&'DQG*ROGHQ-: 7ZRKHWHURF\VWVSHFLILF'1$ UHDUUDQJHPHQWV RI nif RSHURQV LQ Anabaena cylindrica DQG Nostoc VSVWUDLQ0$&0LFURELRO &DUUDVFR &' %XHWWQHU -$ DQG *ROGHQ -: 3URJUDPPHG '1$ UHDUUDQJHPHQW RI D F\DQREDFWHULDO hupL JHQH LQ KHWHURF\VWV3URF1DWO$FDG6FL86$ &DUU 1* DQG 0DQQ 1+ 2FHDQLF F\DQREDFWHULDO SLFRSODQNWRQ ,Q %U\DQW '$ HG 7KH 0ROHFXODU%LRORJ\ RI &\DQREDFWHULDSS .OXZHU'RUGUHFKW &DVH\ (6 DQG *URVVPDQ $5 ,Q YLYR DQG LQ YLWUR FKDUDFWHUL]DWLRQ RI WKH OLJKWUHJXODWHG cpcB2A2 SURPRWHU RI Fremyella diplosiphon. -%DFWHULRO &DVH\ (6 .HKRH '0 DQG *URVVPDQ $5 6XSSUHVVLRQ RI PXWDQWV DEHUUDQW LQ OLJKW LQWHQVLW\ UHVSRQVHV RI FRPSOHPHQWDU\ FKURPDWLFDGDSWDWLRQ-%DFWHULRO &KDQJ & .ZRN 6) %OHHFNHU $% DQG 0H\HURZLW] (0 Arabidopsis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ynechococcus VS 3&& 3ODQWD &ODUNH $. +XUU\ 90 *XVWDIVVRQ 3 DQG 2TXLVW * E 7ZR IXQFWLRQDOO\ GLVWLQFWIRUPV RIWKHSKRWRV\VWHP ,,UHDFWLRQFHQWHU SURWHLQ' LQWKHF\DQREDFWHULXPSynechococcus VS3&& 3URF1DWO$FDG6FL86$ &OHJJ'DQG.RVKODQG' 7KHUROHRIDVLJQDOLQJSURWHLQ LQ EDFWHULDO VHQVLQJ %HKDYLRUDO HIIHFWV RI LQFUHDVHG JHQH H[SUHVVLRQ3URF1DWO$FDG6FL86$ &REOH\-*DQG0LUDQGD5' 0XWDWLRQV DIIHFWLQJ FKURPDWLF DGDSWDWLRQ LQ WKH F\DQRKDFWHULXP Fremyella diplosiphon. %DFWHULRO &RGG *$ &DUKR[\VRPHV DQG ULEXORVH ELVSKRVSKDWH FDUER[\ODVHOR[\JHQDVH ,Q 5RVV $+ 7HPSHVW ': HGV $GYDQFHV LQ PLFURELDO SK\VLRORJ\ 9RO SS $FDGHPLF3UHVV/RQGRQ &RKHQ.XSLHF5*XUHYLW]0DQG=LOEHUVWHLQ$ ([SUHVVLRQ RI glnA LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& LV LQLWLDWHG IURP D VLQJOH nifOLNH SURPRWHU XQGHU YDULRXV QLWURJHQFRQGLWLRQV-%DFWHULRO
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO 6WUHVV &ROHPDQ -5 7KH PROHFXODU DQG ELRFKHPLFDO DQDO\VHV RI &2FRQFHQWUDWLQJ PHFKDQLVPV RI F\DQREDFWHULD DQG JUHHQ DOJDH3ODQW&HOO (QYLURQPHQW &ROOLHU-/DQG*URVVPDQ$5 &KORURVLVLQGXFHGE\QXWULHQW GHSULYDWLRQ LQ Synechococcus VS VWUDLQ 3&& QRW DOO EOHDFKLQJLVWKHVDPH-%DFWHULRO &ROOLHU-/DQG*URVVPDQ $5 $ VPDOOSRO\SHSWLGH WULJJHUV FRPSOHWH GHJUDGDWLRQ RI OLJKWKDUYHVWLQJ SK\FRELOLSURWHLQV LQ QXWULHQWGHSULYHGF\DQREDFWHULD(0%2- &ROOLHU -/ +HUEHUW 6. )RUN '& DQG *URVVPDQ $5 &KDQJHV LQ WKH F\DQREDFWHULDO SKRWRV\QWKHWLF DSSDUDWXV LQ UHVSRQVH WR PDFURQXWULHQW GHSULYDWLRQ 3KRWRV\Q 5HV &ROOLHU -/ DQG 3DOHQLN %3 8UHD XWLOL]DWLRQ E\ PDULQH Synechococcus. -3K\FRO6XSSO &RQOH\3%/HPDX[3*DQG*URVVPDQ $5 &\DQREDFWHULDO OLJKWKDUYHVWLQJ FRPSOH[ VXEXQLWV HQFRGHG LQ WZR UHG OLJKW LQGXFHGWUDQVFULSWV 6FLHQFH &RQOH\ 3% /HPDX[ 3* DQG *URVVPDQ $5 0ROHFXODU FKDUDFWHUL]DWLRQ DQG HYROXWLRQ RI VHTXHQFHV HQFRGLQJ OLJKW KDUYHVWLQJ FRPSRQHQWV LQ WKH FKURPDWLFDOO\ DGDSWLQJ F\DQREDFWHULXP Fremyella diplosiphon. - 0RO %LRO &RQOH\ 3% /HPDX[ 3* /RPD[ 7/ DQG *URVVPDQ $5 *HQHV HQFRGLQJ PDMRU OLJKWKDUYHVWLQJ SRO\SHSWLGHV DUH FOXVWHUHG RQ WKH JHQRPH RI WKH F\DQREDFWHULXP Fremyella diplosiphon. 3URF1DWO$FDG6FL86$ 'H /DV 5LYDV - $QGHUVVRQ % DQG %DUEHU - 7ZR VLWHV RI SULPDU\ GHJUDGDWLRQ RI WKH 'SURWHLQ LQGXFHG E\ DFFHSWRU RU GRQRU VLGH SKRWRLQKLELWLRQ LQ SKRWRV\VWHP ,, FRUH FRPSOH[HV )(%6/HWW GH /RULPLHU 5 %U\DQW '$ 3RUWHU 5' /LX :< -D\ ( DQG 6WHYHQV 6( *HQHV IRU WKH DQG VXEXQLWV RI SK\FRF\DQLQ3URF1DWO$FDG6FL86$ 'HORVPH 5 2OLYH - DQG :ROOPDQ )$ &KDQJHV LQ OLJKW HQHUJ\ GLVWULEXWLRQ XSRQ VWDWH WUDQVLWLRQV DQ LQ YLYR SKRWRDFRXVWLFVWXG\RIWKHZLOGW\SHDQGSKRWRV\QWKHVLVPXWDQWV IURP Chlamydomonas reinhardtii. %LRFKLP%LRSK\V$FWD GH =DPDURF]\ 0 3DTXHOLQ $ DQG (OPHULFK & )XQFWLRQDO RUJDQL]DWLRQRIWKHglnB-glnA FOXVWHURIAzospirillum brasilense. -%DFWHULRO 'HPPLJ$GDPV % &DURWHQRLGV DQG SKRWRSURWHFWLRQ LQ SODQWV $UROHIRUWKH[DQWKRSK\OO ]HD[DQWKLQ %LRFKLP%LRSK\V $FWD 'ROJDQRY 1$0 %KD\D ' DQG *URVVPDQ $5 &\DQREDFWHULDO SURWHLQ ZLWK VLPLODULW\ WR WKH FKORURSK\OO a/bELQGLQJSURWHLQVRIKLJKHUSODQWV(YROXWLRQDQGUHJXODWLRQ3URF 1DWO$FDG6FL86$ 'ROJDQRY 1$0 DQG *URVVPDQ $5 $ SRO\SHSWLGH ZLWK VLPLODULW\ WR SK\FRF\DQLQ a VXEXQLW SK\FRF\DQRELOLQ O\DVH LQYROYHG LQ WKH GHJUDGDWLRQ RIWKH SK\FRELOLVRPHV - %DFWHULRO 6XEPLWWHG 'XNH &6 &H]HDX[ $ DQG $OOHQ 00 &KDQJHV LQ SRO\SHSWLGH FRPSRVLWLRQ RI Synechocystis VS VWUDLQ SK\FRELOLVRUQHVLQGXFHGE\QLWURJHQVWDUYDWLRQ -%DFWHULRO 'XPHU -% DQG %RHJHU 3 8ELTXLWLQ LQ WKH SURNDU\RWH Anabaena variabilis. -%LRO&KHP (EEROH ' DQG =DONLQ + &ORQLQJ DQG FKDUDFWHUL]DWLRQ RID JHQH FOXVWHU IURP Bacillus subtilis HQFRGLQJ QLQH HQ]\PHV IRU de novo SXULQHQXFOHRWLGHV\QWKHVLV-%LRO&KHP
(OPRUMDQL . DQG +HUGPDQ 0 0HWDEROLF FRQWURO RI SK\FRF\DQLQ GHJUDGDWLRQ LQ WKH F\DQREDFWHULXP Synechocystis 3&&$JOXFRVHHIIHFW-*HQ0LFURELRO (OPRUMDQL./LRWHQEHUJ6 +RXPDUG-DQG7DQGHDXGH0DUVDF1 0ROHFXODU FKDUDFWHUL]DWLRQ RI WKH JHQH HQFRGLQJ JOXWDPLQH V\QWKHWDVH LQ WKH F\DQREDFWHULXP Calothrix VS 3&& %LRFKHP%LRSK\V5HV&RPPXQ (QJOLVK56 /RUEDFK 6' 4LQ; DQG 6KLYHO\-0 ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI D FDUER[\VRPH VKHOO JHQH IURP Thiobacillus neapolitanus. 0RO0LFURELRO (VFRXEDV -0 /RPDV 0 /D5RFKH - DQG )DONRZVNL 3* /LJKW LQWHQVLW\ UHJXODWLRQ RI cab JHQH WUDQVFULSWLRQ LV VLJQDOHG E\WKHUHGR[VWDWHRIWKHSODVWRTXLQRQHSRRO 3URF 1DWO$FDG 6FL 86$ (VSLH *6 DQG .DQGDVDP\ 5$ 1DLQGHSHQGHQW +&2 WUDQVSRUW DQG DFFXPXODWLRQ LQ WKH F\DQREDFWHULXP Synechococcus 87(; 3ODQW3K\VLRO (VSLH *6 0LOOHU $* DQG &DQYLQ '7 +LJK DIILQLW\ WUDQVSRUW RI &2 LQ WKH F\DQREDFWHULXP Synechococcus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remyella diplosiphon. -%DFWHULRO )HGHUVSLHO 1$ DQG 6FRWW / &KDUDFWHUL]DWLRQ RI D OLJKW UHJXODWHG JHQH HQFRGLQJ D QHZ SK\FRHU\WKULQDVVRFLDWHG OLQNHU SURWHLQ IURP WKH F\DQREDFWHULXP Fremyella diplosiphon. %DFWHULRO )ORUHV ( *XHUUHUR 0* DQG /RVDGD 0 6KRUWWHUP DPPRQLXP LQKLELWLRQ RI QLWUDWH XWLOL]DWLRQ E\ Anacystis nidulans DQGRWKHUF\DQREDFWHULD$UFK 0LFURELRO )ORUHV ( *XHUUHUR 0* DQG /RVDGD 0 3KRWRV\QWKHWLF QDWXUH RI QLWUDWH XSWDNH DQG UHGXFWLRQ LQ WKH F\DQREDFWHULXP Anacystis nidulans. %LRFKLP%LRSK\V $FWD )ORUHV (DQG +HUUHUR $ $VVLPLODWRU\ QLWURJHQ PHWDEROLVP DQGLWVUHJXODWLRQ ,Q %U\DQW'$ HG 7KH0ROHFXODU%LRORJ\ RI&\DQREDFWHULDSS .OXZHU'RUGUHFKW )ORUHV ( DQG 0XUR3DVWRU 0, 8SWDNH RI JOXWDPLQH DQG JOXWDPDWH E\ WKH GLQLWURJHQIL[LQJ F\DQREDFWHULXP Anabaena VS3&&)(060LFURELRO/HWW )ORUHV ( DQG 0XUR3DVWRU 0, 0XWDWLRQDO DQG NLQHWLF DQDO\VLV RI EDVLF DPLQR DFLG WUDQVSRUW LQ WKH F\DQREDFWHULXP Synechocystis VS3&&$UFK0LFURELRO )RUFKKDPPHU.DQG7DQGHDXGH0DUVDF 1 7KH3,, SURWHLQ LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& LV PRGLILHG E\ VHULQH SKRVSKRU\ODWLRQ DQG VLJQDOV WKH FHOOXODU 1 VWDWXV-%DFWHULRO )RUFKKDPPHU . DQG 7DQGHDX GH 0DUVDF 1 D )XQFWLRQDO DQDO\VLV RI WKH SKRVSKRSURWHLQ 3,, (glnB JHQH SURGXFW LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ3&& -%DFWHULRO )RUFKKDPPHU . DQG 7DQGHDX GH 0DUVDF 1 E 3KRVSKRU\ODWLRQ RI WKH 3,, SURWHLQ (glnB JHQH SURGXFW LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& $QDO\VLV RILQYLWURNLQDVHDFWLYLW\-%DFWHULRO
)RXOGV ,- DQG &DUU 1* $ SURWHRO\WLF HQ]\PH GHJUDGLQJ SK\FRF\DQLQ LQ WKH F\DQREDFWHULXP Anabaena cylindrica. )(060LFURELRO/HWW )ULDV -( )ORUHV ( DQG +HUUHUR $ 5HTXLUHPHQW RI WKH UHJXODWRU\ SURWHLQ 1WF$ IRU WKH H[SUHVVLRQ RI QLWURJHQ DVVLPLODWLRQ DQG KHWHURF\VW GHYHORSPHQW JHQHV LQ WKH F\DQREDFWHULXP Anabaena VS 3&& 0RO 0LFURELRO )UHVQHGR 2 *RPH] 5 DQG 6HUUD -/ &DURWHQRLG FRPSRVLWLRQ LQ WKH F\DQREDFWHULXP Phormidium laminosum: (IIHFWRIQLWURJHQVWDUYDWLRQ)(%6/HWW )ULGODQG / .DSODQ $ DQG 5HLQKROG / 4XDQWLWDWLYH HYDOXDWLRQRIWKHUROH RIDSXWDWLYH&2VFDYHQJLQJHQWLW\LQ WKH F\DQREDFWHULDO &2FRQFHQWUDWLQJ PHFKDQLVP %LRV\VWHP )ULHGEHUJ ' -DJHU .0 .HVVHO 0 6LOPDQ 1- DQG %HUJPDQ % 5XELVFR EXW QRW 5XELVFR DFWLYDVH LV FOXVWHUHG LQ WKH FDUER[\VRPHV RI WKH F\DQREDFWHULXP Synechococcus VS 3&& 0XGLQGXFHGFDUER[\VRPHOHVVPXWDQWV0RO0LFURELRO )ULHGEHUJ' .DSODQ$ $ULHO5 .HVVHO 0DQG 6HLMIIHUV - 7KH IODQNLQJUHJLRQRIWKHJHQHHQFRGLQJWKHODUJHVXEXQLWRI ULEXORVHELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH LV FUXFLDO IRU JURZWK RI WKH F\DQREDFWHULXP Synechococcus 3&& LQ DLU OHYHORI&2-%DFWHULRO )XMLWD< DQG 0XUDNDPL $ 5HJXODWLRQ RIHOHFWURQ WUDQVSRUW FRPSRVLWLRQ LQ F\DQREDFWHULDO SKRWRV\QWKHWLF V\VWHP 6WRLFKLRPHWU\DPRQJSKRWRV\VWHP ,DQG ,,FRPSOH[HV DQG WKHLU OLJKWKDUYHVWLQJ DQWHQQDH DQG F\WRFKURPH b6f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nifH DQG QLI+OLNH frxC JHQHV IURP WKH ILODPHQWRXV QLWURJHQIL[LQJ F\DQREDFWHULXP Plectonema boryanum. 3ODQW&HOO3K\VLRO )XNX]DZD+6X]XNL(.RPXNDL<DQG0L\DFKL6 $JHQH KRPRORJRXVWRFKORURSODVWFDUERQLFDQK\GUDVH(icfA) LVHVVHQWLDO WR SKRWRV\QWKHWLF FDUERQ IL[DWLRQ LQ WKH F\DQREDFWHULXP Synechococcus 3&& 3URF 1DWO $FDG 6FL 86$ *DQWW( 3K\FRELOLVRPHV $QQX5HY3ODQW3K\VLRO
' %KD\D 56FKZDU]DQG$5*URVVPDQ *DVSDULFK*(%X]E\-6%U\DQW'$3RUWHU5'DQG6WHYHQV-U6( 7KH HIIHFWV RIOLJKW LQWHQVLW\ DQG QLWURJHQ VWDUYDWLRQ RQ WKHSK\FRF\DQLQSURPRWHULQWKHF\DQREDFWHULXPSynechococcus VS 3&& ,Q %LJJLQV - HG 3URJUHVV LQ 3KRWRV\QWKHVLV 5HVHDUFK 9RO ,9 SS 0DUWLQXV1LMKRII 3XEOLVKHUV 'RUGUHFKW *LQJULFK -& *DVSDULFK *( 6DXHU . DQG %U\DQW '$ 1XFOHRWLGH VHTXHQFH DQG H[SUHVVLRQ RIWKH WZR JHQHV HQFRGLQJ ' SURWHLQ DQG WKH VLQJOH JHQH HQFRGLQJ WKH &3 SURWHLQ RI SKRWRV\WHP ,, LQ WKH F\DQREDFWHULXP Synechococcus VS 3&& 3KRWRV\Q5HV *OD]HU$1 /LJKWKDUYHVWLQJE\SK\FRELOLVRPHV $QQX 5HY %LRSK\V%LRSK\V&KHP *OD]HU$1 3K\FRELOLVRPHV DVVHPEO\ DQG DWWDFKPHQW ,Q )D\ 3 9DQ %DDOHQ & HGV 7KH &\DQREDFWHULD SS (OVHYLHU%LRPHGLFDO$PVWHUGDP *OD]HU $1 /XQGHOO '-
&KDSWHU 0ROHFXODU5HVSRQVHVWR (QYLURQPHQWDO 6WUHVV *URVVPDQ $5 %KD\D ' $SW .( DQG .HKRH '0 /LJKW KDUYHVWLQJ FRPSOH[HV LQ R[\JHQLF SKRWRV\QWKHVLV 'LYHUVLW\ FRQWURODQGHYROXWLRQ$QQX5HY*HQHW *URVVPDQ $5 %KD\D ' DQG &ROOLHU -/ D 6SHFLILF DQG JHQHUDOUHVSRQVHVRIF\DQREDFWHULDWRPDFURQXWULHQWGHSULYDWLRQ ,Q 7RUULDQL*RULQL$
+RZDUG -% DQG 5HHV '& 1LWURJHQDVH $ QXFOHRWLGH GHSHQGHQWPROHFXODUVZLWFK$QQX5HY%LRFKHP +RZHOO /' *ULIILWKV & 6ODGH /: 3RWWV 0 DQG .HQQHOO\ 3- 6XEVWUDWH VSHFLILFLW\ RI ,SK3 D F\DQREDFWHULDO GXDO VSHFLILFLW\ SURWHLQ SKRVSKDWDVH ZLWK 0$3 NLQDVH SKRVSKDWDVH DFWLYLW\%LRFKHP +U\QLHZLF] 0 6LUNR $ 3DOXFKD $ %RFN $ DQG +XODQLFND ' 6XOIDWH DQG WKLRVXOIDWH WUDQVSRUW LQ Escherichia coli . LGHQWLILFDWLRQRIDJHQHHQFRGLQJDQRYHOSURWHLQLQYROYHG LQ WKLRVXOIDWHELQGLQJ-%DFWHULRO +XD - &KDQJ & 6XQ 4 DQG 0H\HURZLW] (0 (WK\OHQH LQVHQVLWLYLW\ FRQIHUUHG E\ Arabidopsis ETS JHQH 6FLHQFH +XDQJ 5& DQG &KRZ 7- &KDUDFWHUL]DWLRQ RIWKH UK\WKPLF QLWURJHQIL[LQJ DFWLYLW\ RI Synechococcus VS 5), DW WKH WUDQVFULSWLRQOHYHO&XUU0LFURELRO +XDQJ 7& DQG &KRZ 7- &RPSDUDWLYH VWXGLHV RI VRPH QLWURJHQIL[LQJ XQLFHOOXODU F\DQREDFWHULD LVRODWHG IURP ULFH ILHOGV-*HQ0LFURELRO ,VKLJH.1DJDVDZD6 7RNLVKLWD6,DQG0L]XQR7 $QRYHO GHYLFHRIEDFWHULDOVLJQDOWUDQVGXFHUV(0%2- ,XFKL 6 DQG /LQ (&& $GDSWDWLRQ RI Escherichia coli WR UHGR[HQYLURQPHQWVE\JHQHH[SUHVVLRQ0RO0LFURELRO -DJHU.0DQG%HUJPDQ% /RFDOL]DWLRQRIDPXOWLIXQFWLRQDO FKDSHURQLQ *UR(/ SURWHLQ LQ QLWURJHQIL[LQJ Anabaena 3&& SUHVHQFH LQ YHJHWDWLYH FHOOV DQG KHWHURF\VWV 3ODQWD -HDQMHDQ 5 DQG %URGD( 'HSHQGHQFH RIVXOIDWH XSWDNH E\ Anacystis nidulans RQ HQHUJ\ RVPRWLF VKRFN DQG RQ VXOIDWH VWDUYDWLRQ$UFK0LFURELRO -HUORY1* 0DULQH2SWLFV(OVHYLHU$PVWHUGDP -RKQVRQ$' 7KHSULFHRIUHSUHVVLRQ&HOO -RKQVRQ 75 +D\QHV ,, -, :HDODQG -/
.DSODQ$ =HQYLUWK' 0DUFXV < 2PDWD7 DQG 2JDZD7 (QHUJL]DWLRQ DQG DFWLYDWLRQ RILQRUJDQLF FDUERQ XSWDNH E\ OLJKW LQF\DQREDFWHULD 3ODQW3K\VLRO .DUDJRXQL$' %OR\FH6$DQG&DUU1* 7KHSUHVHQFHDQG DEVHQFHRILQRUJDQLF FDUERQFRQFHQWUDWLQJV\VWHPVLQXQLFHOOXODU F\DQREDFWHULD)(060LFURELRO/HWW .DWRK$/HH.6)XNX]DZD+2K\DPD.DQG2JDZD7D FHP$ KRPRORJXH HVVHQWLDO WR &2 WUDQVSRUW LQ WKH F\DQREDFWHULXP Synechocystis 3&& 3URF 1DWO $FDG 6FL 86$ .DWRK$ 6RQRGD0 .DWRK + DQG 2JDZD 7E $EVHQFH RI OLJKWLQGXFHG SURWRQ H[WUXVLRQ LQ D cotAOHVV PXWDQW RI Synechocystis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nabaena variabilis. )(060LFURELRO/HWW .HQWHPLFK 7 +DYHUNDPS * DQG %RWKH + 7KH H[SUHVVLRQ RI D WKLUG QLWURJHQDVH LQ WKH F\DQREDFWHULXP Anabaena variabilis. =1DWXURIRUVFKF .H\VH 60 $Q HPHUJLQJ IDPLO\ RI GXDO VSHFLILFLW\ 0$3 NLQDVHSKRVSKDWDVHV%LRFKLP%LRSK\V$FWD .LNXFKL + $LFKL 0 6X]XNL , DQG 2PDWD 7 3RVLWLYH UHJXODWLRQ E\ QLWULWH RI WKH QLWUDWH DVVLPLODWLRQ RSHURQ LQ WKH F\DQREDFWHULD Synechococcus VS VWUDLQ 3&& DQG Plectonema boryanum. -%DFWHULRO .LP-+ *OLFN 5(DQG0HOLV $ '\QDPLFV RISKRWRV\VWHP VWRLFKLRPHWU\ DGMXVWPHQW E\ OLJKW TXDOLW\ LQ FKORURSODVWV 3ODQW 3K\VLRO .RED\DVKL 0 5RGULJXH] 5 /DUD & DQG 2PDWD 7 ,QYROYHPHQW RI WKH &WHUPLQDO GRPDLQ RI DQ $73ELQGLQJ VXEXQLW LQ WKH UHJXODWLRQ RI WKH $%&W\SH QLWUDWHQLWULWH WUDQVSRUWHURIWKHF\DQREDFWHULXPSynechococcus VS VWUDLQ3&& -%LRO&KHP .ROE $ %XVE\ 6 %XF + *DUJHV 6 DQG $GE\D 6 7UDQVFULSWLRQDO UHJXODWLRQ E\ F\FOLF $03 DQG LWV UHFHSWRU SURWHLQ$QQX5HY%LRFKHP .RUQEHUJ $ ,QRUJDQLF SRO\SKRVSKDWH $ PROHFXODU IRVVLO FRPHWR OLIH ,Q 7RPDQL*RULQL $
' %KD\D 56FKZDU]DQG$5*URVVPDQ .UXSD = RTXLVW * DQG *XVWDIVVRQ 3 3KRWRLQKLELWLRQ RI SKRWRV\QWKHVLV DQG JURZWK UHVSRQVHV DW GLIIHUHQW OLJKW OHYHOV LQ psbA JHQH PXWDQWV RI WKH F\DQREDFWHULXP Synechococcus. 3K\VLRO3ODQW .XONDUQL 5' DQG *ROGHQ 66 $GDSWDWLRQ WR KLJK OLJKW LQWHQVLW\ LQ Synechococcus VS VWUDLQ 3&& 5HJXODWLRQ RI WKUHH psbA JHQHV DQG WZR IRUPV RIWKH ' SURWHLQ - %DFWHULRO .XONDPL 5' DQG *ROGHQ 66 )RUP ,, RI ' LV LPSRUWDQW GXULQJ WUDQVLWLRQ IURP VWDQGDUG WR KLJK OLJKW LQWHQVLW\ LQ Synechococcus VSVWUDLQ3&&3KRWRV\Q5HV .XONDPL5' 6FKDHIHU05DQG*ROGHQ6 6 7UDQVFULSWLRQDO DQG SRVWWUDQVFULSWLRQDO FRPSRQHQWV RI psbA UHVSRQVH WR KLJK OLJKW LQWHQVLW\ LQ Synechococcus VS VWUDLQ 3&& - %DFWHULRO /DEDUUH - 7KXULDX[ 3 DQG &KDXYDW ) *HQHWLF DQDO\VLV RI DPLQR DFLG WUDQVSRUW LQ WKH IDFXOWDWLYHO\ KHWHURWURSKLF F\DQREDFWHULXP Synechocystis VS VWUDLQ - %DFWHULRO /DPPHUV 3- *ROGHQ -: DQG +DVHONRUQ 5 ,GHQWLILFDWLRQ DQGVHTXHQFHRIDJHQHUHTXLUHGIRUDGHYHORSPHQWDOO\UHJXODWHG '1$H[FLVLRQLQAnabaena. &HOO /DUD & 5RPHUR -0 DQG *XHUUHUR 0* 5HJXODWHG QLWUDWH WUDQVSRUW LQ WKH F\DQREDFWHULXP Anacystis nidulans. - %DFWHULRO /DX 5- 0DF.HQ]LH 00 DQG 'RROLWWOH :) 3K\FRF\DQLQ V\QWKHVLV DQG GHJUDGDWLRQLQWKHEOXHJUHHQ EDFWHULXPAnacystis nidulans. -%DFWHULRO /DXGHQEDFK '( (KUKDUGW ' *UHHQ /DQG *URVVPDQ $5 ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI D VXOIXUUHJXODWHG JHQH HQFRGLQJ D SHULSODVPLFDOO\ ORFDOL]HG SURWHLQ ZLWK VHTXHQFH VLPLODULW\WRUKRGDQHVH-%DFWHULRO /DXGHQEDFK '( DQG *URVVPDQ $5 &KDUDFWHUL]DWLRQ DQG PXWDJHQHVLV RI VXOIXUUHJXODWHG JHQHV LQ D F\DQREDFWHULXP (YLGHQFHIRUIXQFWLRQLQVXOIDWHWUDQVSRUW-%DFWHULRO /DZU\ 1+ DQG -HQVHQ 7( &RQGHQVHG SKRVSKDWH GHSRVLWLRQ VXOIXU DPLQR DFLG XVH DQG XQLGLUHFWLRQDO WUDQVXOIXUDWLRQ LQ Synechococcus leopoliensis. $UFK 0LFURELRO /HRQJ7< DQG $QGHUVRQQ-0 /LJKWTXDOLW\DQG LUUDGLDQFH DGDSWDWLRQ RI WKH FRPSRVLWLRQ DQG IXQFWLRQ RI SHD WK\ODNRLG PHPEUDQHV %LRFKLP%LRSK\V$FWD /L5'LFNHUVRQ16 0XHOOHUUW DQG *ROGHQ 66 6SHFLILF ELQGLQJ RI Synechococcus VS VWUDLQ 3&& SURWHLQV WR WKH HQKDQFHU HOHPHQW RI psbAII UHTXLUHG IRU KLJKOLJKWLQGXFHG H[SUHVVLRQ-%DFWHULRO /L 5 DQG *ROGHQ 66 (QKDQFHU DFWLYLW\ RIOLJKWUHVSRQVLYH UHJXODWRU\ HOHPHQWV LQ WKH XQWUDQVODWHG OHDGHU UHJLRQV RI F\DQREDFWHULDO psbA JHQHV3URF1DWO$FDG6FL86$ /RFNDX : 0DVVDOVN\ % DQG 'LUPHLHU $ 3XULILFDWLRQ DQG SDUWLDO FKDUDFWHUL]DWLRQ RI D FDOFLXPVWLPXODWHG SURWHDVH IURP WKH F\DQREDFWHULXP Anabaena variabilis. (XU - %LRFKHP /RPD[ 7/ &RQOH\ 3% 6FKLOOLQJ - DQG *URVVPDQ $5 ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI OLJKWUHJXODWHG SK\FRELOLVRPH OLQNHU SRO\SHSWLGH JHQHV DQG WKHLU WUDQVFULSWLRQ DV D SRO\FLVWURQLFP51$-%DFWHULRO /XTXH , )ORUHV ( DQG +HUUHUR $ 1LWUDWH UHGXFWDVH JHQH IURP Synechococcus VS 3&& +RPRORJ\ EHWZHHQ F\DQREDFWHULDODQGKLJKHUSODQWQLWUDWHUHGXFWDVH 3ODQW0RO%LRO
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO 6WUHVV /XTXH,)ORUHV(DQG+HUUHUR$D 0ROHFXODUPHFKDQLVPIRU WKH RSHUDWLRQ RIQLWURJHQ FRQWUROLQF\DQREDFWHULD (0%2- /XTXH , )ORUHV ( DQG +HUUHUR $ E 1LWUDWH DQG QLWULWH WUDQVSRUW LQ WKH F\DQREDFWHULXP Synechococcus VS 3&& DUH PHGLDWHG E\ WKH VDPH SHUPHDVH %LRFKLP %LRSK\V $FWD /XTXH ,+HUUHUD$ )ORUHV(DQG0DGXHLLR) &OXVWHULQJRI JHQHV LQYROYHG LQ QLWUDWH DVVLPLODWLRQ LQ WKH F\DQREDFWHULXP Synechococcus. 0RO*HQ*HQHW 0DFNHUUDV $+ GH &KD]DO 10 DQG 6PLWK *' 7UDQVLHQW DFFXPXODWLRQV RI F\DQRSK\FLQ LQ Anabaena cylindrica DQG Synechocystis -*HQ0LFURELRO 0DGXHLLR ) %RUULDV:( YDQ$UNHO*$DQG*XHUUHUR0* ,VRODWLRQ DQGFKDUDFWHUL]DWLRQ RIAnacystis nidulans 5 PXWDQWV DIIHFWHG LQ QLWUDWH DVVLPLODWLRQ (VWDEOLVKPHQW RI WZR QHZ PXWDQWW\SHV0RO*HQ*HQHW 0DGXHLLR ) )ORUHV ( DQG *XHUUHUR 0* &RPSHWLWLRQ EHWZHHQ QLWUDWH DQG QLWULWH XSWDNH LQ WKH F\DQREDFWHULXP Anacystis nidulans. %LRFKLP%LRSK\V$FWD 0DHGD 6, DQG 2PDWD 7 6XEVWUDWHELQGLQJ OLSRSURWHLQ RI WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& LQYROYHGLQWKHWUDQVSRUWRIQLWUDWHDQGQLWULWH -%LRO&KHP 0DNLQR . $PHPXUD 0 .LP 6. 1DNDWD $ DQG 6KLQDJDZD + 0HFKDQLVPRIWUDQVFULSWLRQDODFWLYDWLRQRIWKHSKRVSKDWH UHJXORQ LQ Escherichia coli. ,Q 7RPDQL*RULQL $
0D]HO ' *XJOLHOPL * +RXPDUG + 6LGOHU : %U\DQW '$ DQG 7DQGHDX GH 0DUVDF 1 *UHHQ OLJKW LQGXFHV WUDQVFULSWLRQ RI WKH SK\FRHU\WKULQ RSHURQ LQ WKH F\DQREDFWHULXP Calothrix 1XFOHLF$FLGV5HV 0D]HO ' +RXPDUG - DQG 7DQGHDX GH 0DUVDF 1 $ PXOWLJHQH IDPLO\ LQ Calothrix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ynechococcus 87(; $UFK0LFURELRO 0HHNV -& :RON &3 /RFNDX : 6FKLOOLQJ 1 6KDIIHU 3: DQG &KLHQ :6 3DWKZD\V RI DVVLPLODWLRQ RI >1@1 DQG 1+ E\ F\DQREDFWHULD ZLWK DQG ZLWKRXW KHWHURF\VWV - %DFWHULRO 0HHNV -& :\NRII ./ &KDSPDQ -6 DQG (QGHUOLQ &6 5HJXODWLRQ RI H[SUHVVLRQ RI QLWUDWH DQG GLQLWURJHQ DVVLPLODWLRQ E\ Anabaena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hlamydomonas reinhardtii DOWHUV SKRWRV\VWHPVWRLFKLRPHWU\DQGLPSURYHV WKHTXDQWXPHIILFLHQF\ RISKRWRV\QWKHVLV3KRWRV\Q5HV 0EULGD $ )ORUHV ( DQG )ORUHQFLR )- 5HJXODWLRQ RI JOXWDPLQH V\QWKHWDVH DFWLYLW\ LQ WKH XQLFHOOXODU F\DQREDFWHULXP Synechocystis VS VWUDLQ 3&& E\ WKH QLWURJHQ VRXUFH (IIHFWRIDPPRQLXP-%DFWHULRO 0WULGD $ )ORUHV ( DQG )ORUHQFLR )- 5HJXODWLRQ RI Anabaena VS VWUDLQ 3&& JOXWDPLQH V\QWKHWDVH DFWLYLW\ LQ D Synechocystis VS VWUDLQ 3&& GHULYDWLYH VWUDLQ EHDULQJ WKH Anabaena glnA JHQH DQG D PXWDWHG KRVW glnA JHQH - %DFWHULRO 0EULGD $ /HXUHQWRS / &DQGDX 3 DQG )ORUHQFLR )- 3XULILFDWLRQ DQG SURSHUWLHV RI JOXWDPLQH V\QWKHWDVHV IURP WKH F\DQREDFWHULD Synechocystis VS VWUDLQ 3&& DQG Culothrix VSVWUDLQ3&&-%DFWHULRO 0HUULFN 0- 5HJXODWLRQ RIQLWURJHQ IL[DWLRQ JHQHV LQ IUHH OLYLQJDQGV\PELRWLFEDFWHULD ,Q 6WDFH\*%XPV5+ (YDQV +- HGV %LRORJLFDOQLWURJHQIL[DWLRQSS &KDSPDQ +DOO1HZ
0LOOHU$*(VSLH*6 3KRWRV\QWKHWLFPHWDEROLVPRIF\DQDWH E\ WKH F\DQREDFWHULXP Synechococcus 87(; $UFK 0LFURELRO 0LOOHU $* (VSLH *6 DQG &DQYLQ '7 &KORURSK\OO a IOXRUHVFHQFH\LHOGDV DPRQLWRURIERWK DFWLYH &2 DQG +&2 WUDQVSRUW E\ WKH F\DQREDFWHULXP Synechococcus 87(; 3ODQW3K\VLRO 0LOOHU$*(VSLH*6DQG&DQYLQ'7 3K\VLRORJLFDODVSHFWV RI&2DQG+&2WUDQVSRUWE\F\DQREDFWHULD$UHYLHZ &DQ- %RW 0LWVXL$.XPD]DZD67DNDKDVKL$ONHPRWR+6&DQG$UDL7 6WUDWHJ\ E\ ZKLFK QLWURJHQIL[LQJ XQLFHOOXODU F\DQREDFWHULDJURZSKRWRDXWRWURSLFDOO\1DWXUH 0RQWHVLQRV 0/ +HUUHUR $ DQG )ORUHV ( $PLQR DFLG WUDQVSRUW V\VWHPV UHTXLUHG IRU GLD]RWURSLF JURZWK LQ WKH F\DQREDFWHULXP Anabaena VSVWUDLQ3&&-%DFWHULRO 0RQWHVLQRV 0/ +HUUHUR $ DQG )ORUHV ( $PLQR DFLG WUDQVSRUW LQ WD[RQRPLFDOO\ GLYHUVH F\DQREDFWHULD DQG LGHQWLILFDWLRQRIWZRJHQHVHQFRGLQJHOHPHQWVRIDQHXWUDODPLQR DFLG SHUPHDVH SXWDWLYHO\ LQYROYHG LQ UHFDSWXUH RI OHDNHG K\GURSKRELFDPLQRDFLGV-%DFWHULRO 0XOOLJDQ 0( DQG +DVHONRUQ :- 1LWURJHQ IL[DWLRQ (nif) JHQHV RI WKH F\DQREDFWHULXP Anabaena VSHFLHV VWUDLQ 3&& 7KHnifB-fdxN-nifS-nifU RSHURQ-%LRO&KHP 0XOOLQHDX[ &: ([FLWDWLRQ HQHUJ\ WUDQVIHU IURP SK\FRELOLVRPHV WR SKRWRV\VWHP , LQ F\DQREDFWHULD %LRFKLP %LRSK\V$FWD 0XOOLQHDX[&:DQG $OOHQ -) 6WDWHVWDWH WUDQVLWLRQV LQ WKH F\DQREDFWHULXP Synechococcus DUH FRQWUROOHG E\ WKH UHGR[ VWDWH RI HOHFWURQ FDUULHUV EHWZHHQ SKRWRV\VWHP , DQG SKRWRV\VWHP,,3KRWRV\Q5HV 0XOOLQHDX[ 30 *DOORQ -5 DQG &KDSOLQ $( $FHW\OHQH UHGXFWLRQ QLWURJHQ IL[DWLRQ E\ F\DQREDFWHULD JURZQ XQGHU DOWHUQDWLQJOLJKWGDUNF\FOHV)(060LFURELRO/HWW 0XUDNDPL $ DQG )XMLWD < 5HJXODWLRQ RI SKRWRV\VWHP VWRLFKLRPHWU\ LQ WKH SKRWRV\QWKHWLF V\VWHP RI WKH F\DQRSK\WH Synechocystis 3&& LQ UHVSRQVH WR OLJKW LQWHQVLW\ 3ODQW &HOO3K\VLRO 0\HUV - *UDKDP -5 DQG :DQJ 57 /LJKW KDUYHVWLQJ LQ Anacystis nidulans VWXGLHG LQ SLJPHQW PXWDQWV 3ODQW 3K\VLRO 1DJD\D0$LED+DQG0L]XQR7 7KHsphR SURGXFWDWZR FRPSRQHQW V\VWHP UHVSRQVH UHJXODWRU SURWHLQ UHJXODWHV SKRVSKDWH DVVLPLODWLRQ LQ Synechococcus VS VWUDLQ 3&& E\ ELQGLQJ WR WZR VLWHV XSVWUHDP IURP WKH phoA SURPRWHU - %DFWHULRO 1DYDUUR)&KDYH]6&DQGDX3DQG)ORUHQFLR)- ([LVWHQFH RI WZR IHUUHGR[LQJOXWDPDWH V\QWKDVHV LQ WKH F\DQREDFWHULXP Synechocystis VS 3&& ,VRODWLRQ DQG LQVHUWLRQDO LQDFWLYDWLRQRIgltB DQG glt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
'%KD\D 56FKZDU]DQG$5*URVVPDQ 7- HGV 7ZR&RPSRQHQW6LJQDO7UDQVGXFWLRQSS $60 3UHVV:DVKLQJWRQ'& 2FKRD GH $OGD - 7DSLD 0, )UDQFN ) /ODPD 0- DQG 6HUUD -/ D &KDQJHV LQ QLWURJHQ VRXUFH PRGLI\ GLVWULEXWLRQ RI H[FLWDWLRQ HQHUJ\ LQ WKH F\DQREDFWHULXP Phormidium laminosum. 3K\VLRO3ODQW 2FKRD GH $OGD -$* 7DSLD 0, /ODPD 0- DQG 6HUUD -/E &KDQJHVLQSKRWRV\QWKHWLF\LHOG DPLQRDFLGV DQGRUJDQLF DFLGV DUH LQGXFHG E\ DPPRQLXP DGGLWLRQ WR FHOOV RI Phormidium Laminosum F\DQRSK\FHDH - 3K\FRO 2JDZD 7 $ JHQH KRPRORJRXV WR WKH VXEXQLW JHQH RI 1$'+GHK\GURJHQDVHLVHVVHQWLDO WR LQRUJDQLF FDUERQ WUDQVSRUW RI Synechocystis 3&& 3URF1DWO$FDG6FL86$ 2JDZD 7 ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WKH ictA/ndhL JHQH SURGXFW HVVHQWLDO WR LQRUJDQLF FDUERQ WUDQVSRUW RI Synechocystis 3&& 3ODQW 3K\VLRO 2JDZD 7 FHP$ KRPRORJXH LQ F\DQREDFWHULD LWV IXQFWLRQ DQG ORFDOL]DWLRQ ,Q 86-DSDQ :RUNVKRS RQ $GYDQFHV LQ WKH 0ROHFXODU %LRORJ\ RI 3KRWRV\QWKHVLV *UDQG &DQ\RQ 1DWLRQDO3DUN86$ 2JDZD 7 $PLFKD\ ' DQG *XUHYLW] 0 D ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI WKH ccmM JHQH UHTXLUHG E\ WKH F\DQREDFWHULXP Synechocystis 3&& IRU LQRUJDQLF FDUERQ XWLOL]DWLRQ3KRWRV\Q5HV 2JDZD7 0DUFR(DQG2UXV 0,E $JHQHFFP$ UHTXLUHG IRUFDUER[\VRPHIRUPDWLRQLQWKHF\DQREDFWHULXP Synechocystis VSVWUDLQ3&&-%DFWHULRO 2JDZD 7 0L\DQR $ DQG ,QRXH < 3KRWRV\VWHP,GULYHQ LQRUJDQLF FDUERQ WUDQVSRUW LQ WKH F\DQREDFWHULXP Anacystis nidulans. %LRFKLP%LRSK\V$FWD 2KNL.DQG)XMLWD< $HURELFQLWURJHQDVHDFWLYLW\PHDVXUHG DV DFHW\OHQH UHGXFWLRQ LQ WKH PDULQH QRQKHWHURF\VWRXV F\DQREDFWHULXP Trichodesmium VS JURZQ XQGHU DUWLILFLDO FRQGLWLRQV0DULQH%LRORJ\ 2KPRUL 0 2KPRUL . DQG 6WURWPDQQ + ,QKLELWLRQ RI QLWUDWH XSWDNH E\ DPPRQLD LQ D EOXHJUHHQ DOJD Anabaena cylindrica. $UFK0LFURELRO 2PDWD7 &ORQLQJDQGFKDUDFWHUL]DWLRQRIWKHnrtA JHQHWKDW HQFRGHV D N'D SURWHLQ LQYROYHG LQ QLWUDWH WUDQVSRUW LQ WKH F\DQREDFWHULXP Synechococcus 3&& 3ODQW &HOO 3K\VLRO 2PDWD 7 1LWURJHQ UHJXODWLRQ RI FDUERQ DQG QLWURJHQ DVVLPLODWLRQ JHQHV LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& ,Q 86-DSDQ :RUNVKRS RQ $GYDQFHV LQ WKH 0ROHFXODU %LRORJ\ RI 3KRWRV\QWKHVLV *UDQG &DQ\RQ 1DWLRQDO3DUN86$ 2PDWD 7 $QGULHVVH ; DQG +LUDQR $ ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQRIDJHQH FOXVWHULQYROYHG LQQLWUDWH WUDQVSRUW RI WKH F\DQREDFWHULXP Synechococcus VS 3&& 0ROHF *HQ *HQHW 2PDWD 7 &DUOVRQ 7- 2JDZD 7 DQG 3LHUFH - 6HTXHQFLQJ DQGPRGLILFDWLRQRIWKHJHQHHQFRGLQJ WKH NLORGDOWRQ SURWHLQ LQ WKH F\WRSODVPLF PHPEUDQH RI Synechococcus 3&& 3ODQW3K\VLRO 2PDWD 7 DQG 2JDZD 7 %LRV\QWKHVLV RI D N' SRO\SHSWLGHLQWKHF\WRSODVPLFPHPEUDQHRIWKHF\DQREDFWHULXP Anacystis nidulans VWUDLQ 5 GXULQJ DGDSWDWLRQ WR ORZ &2 FRQFHQWUDWLRQ3ODQW3K\VLRO 2PDWD 7 2KPRUL 0 $UDL 1 DQG 2JDZD 7 *HQHWLFDOO\ HQJLQHHUHG PXWDQW RI WKH F\DQREDFWHULXP Synechococcus 3&& GHIHFWLYHLQQLWUDWHWUDQVSRUW 3URF1DWO$FDG6FL86$
&KDSWHU 0ROHFXODU5HVSRQVHVWR(QYLURQPHQWDO 6WUHVV gTXLVW* &KRZ :6 DQG $QGHUVRQ -0 3KRWRLQKLELWLRQ RI SKRWRV\QWKHVLV UHSUHVHQWV D PHFKDQLVP IRU WKH ORQJWHUP UHJXODWLRQRISKRWRV\VWHP,,3ODQWD 2ZQE\ -' 6KDQQDKDQ 0 DQG +RRG ( 3URWHLQ V\QWKHVLV DQG GHJUDGDWLRQ LQAnabaena GXULQJ QLWURJHQ VWDUYDWLRQ - *HQ 0LFURELRO 3DNUDVL +% *HQHWLF DQDO\VLV RIWKH IRUP DQG IXQFWLRQ RI SKRWRV\VWHP,DQGSKRWRV\VWHP,,$QQX5HY*HQHW 3DUNLQVRQ -6 DQG .RIRLG (& &RPPXQLFDWLRQ PRGXOHV LQ EDFWHULDOVLJQDOLQJSURWHLQV$QQX5HY*HQHW 3HLIIHU:( 'HVURVLHUV 0* DQG 0HQLFN '5 &ORQLQJ DQG H[SUHVVLRQ RI WKH XQLTXH &D$73DVH IURP Flavobacterium odoratum. -%LRO&KHP 3HUHJR 0 DQG +RFK -$ 3URWHLQ DVSDUWDWH SKRVSKDWDVHV FRQWURO WKH RXWSXW RI WZRFRPSRQHQW VLJQDO WUDQVGXFWLRQ V\VWHPV7UHQGV*HQHW 3HWHUVRQ 5% 'RODQ ( &DOYHUW +( DQG .H % (QHUJ\ WUDQVIHU IURP SK\FRELOLSURWHLQV WR SKRWRV\VWHP , LQ YHJHWDWLYH FHOOV DQG KHWHURF\VWV RIAnabaena variabilis. %LRFKLP %LRSK\V $FWD 3LHUFH-&DUOVRQ7-DQG:LOOLDPV-*. $QRPDORXVR[\JHQ VHQVLWLYLW\LQDF\DQREDFWHULDOPXWDQWUHTXLULQJWKHH[SUHVVLRQRI ULEXORVH ELVSKRVSKDWH FDUER[\ODVH IURP D SKRWRV\QWKHWLF DQDHUREH3URF1DWO$FDG6FL86$ 3RUWHU * 7UHGZHOO &- 6HDUOH *): DQG %DUEHU - 3LFRVHFRQG WLPHUHVROYHG HQHUJ\ WUDQVIHU LQ Porphyridium cruentum. 3DUW, ,Q WKHLQWDFWDOJD %LRFKLP%LRSK\V$FWD 3RVDV5:XUJOHU0XUSK\600DHGD7:LWWHQ($7KDL7&DQG 6DLWR +
RVPRVHQVRU&HOO 3RWWV06XQ+0RFNDLWLV..HQQHOO\3-5HHG'DQG7RQNV1. $ SURWHLQW\URVLQHVHULQH SKRVSKDWDVH HQFRGHG E\ WKH JHQRPH RIWKH F\DQREDFWHULXP Nostoc commune 87(; - %LRO&KHP 3ULFH *' DQG %DGJHU 0 D ([SUHVVLRQ RIKXPDQ FDUERQLF DQK\GUDVH LQ WKH F\DQREDFWHULXP Synechococcus 3&& 3ODQW3K\VLRO 3ULFH*'DQG%DGJHU05 E ([SUHVVLRQ RIKXPDQFDUERQLF DQK\GUDVH LQ WKH F\DQREDFWHULXP Synechococcus 3&& FUHDWHVDKLJK&2 UHTXLULQJSKHQRW\SH 3ODQW3K\VLRO 3ULFH *' DQG %DGJHU 05 (YLGHQFH IRU WKH UROH RI FDUER[\VRPHV RQ F\DQREDFWHULDO &2FRQFHQWUDWLQJ PHFKDQLVP &DQ-%RW 3ULFH *' &ROHPDQ -5 DQG %DGJHU 05 $VVRFLDWLRQ RI FDUERQLF DQK\GUDVH DFWLYLW\ ZLWK FDUER[\VRPHV IURP WKH F\DQREDFWHULXP Synechococcus 3&& 3ODQW 3K\VLRO 5DL $1 %RUWKDNXU 0 DQG %HUJPDQ % 1LWURJHQDVH GHSUHVVLRQLWVUHJXODWLRQDQGPHWDEROLFFKDQJHVDVVRFLDWHGZLWK GLD]RWURSK\ LQ WKH QRQKHWHURF\VWRXV F\DQREDFWHULXP Plectonema boryanum 3&& - *HQ 0LFURELRO 5DPDVXEUDPDQLDQ 76 :HL 7) DQG *ROGHQ -: 7ZR Anabaena VS VWUDLQ 3&& '1$ELQGLQJ IDFWRUV LQWHUDFW ZLWK YHJHWDWLYH FHOO DQG KHWHURF\VWVSHFLILF JHQHV - %DFWHULRO 5DPDVXEUDPDQLDQ 76 :HL 7) 2OGKDP $. DQG *ROGHQ -: 7UDQVFULSWLRQRIWKHAnabaena VS VWUDLQntcA JHQH 0XOWLSOHWUDQVFULSWVDQG1WF$ELQGLQJ-%DFWHULRO 5DYHQ-$ ,PSOLFDWLRQVRILQRUJDQLF& XWLOL]DWLRQ (FRORJ\ HYROXWLRQDQGJHRFKHPLVWU\&DQ-%RW
5DYLG 6 0DWVXPXUD 3 DQG (LVHQEDFK 0 5HVWRUDWLRQ RI IODJHOODUFORFNZLVHURWDWLRQLQEDFWHULDOHQYHORSHV E\LQVHUWLRQRI WKH FKHPRWD[LV SURWHLQ &KH< 3URF 1DWO $FDG 6FL 86$ 5D\-0 %KD\D'%ORFN0$DQG*URVVPDQ$5 ,VRODWLRQ WUDQVFULSWLRQ DQG LQDFWLYDWLRQ RI WKH JHQH IRU DQ DW\SLFDO DONDOLQH SKRVSKDWDVH RI Synechococcus VS VWUDLQ 3&& - %DFWHULRO 5HGG\ .- +DVNHOO -% 6KHUPDQ '0 DQG 6KHUPDQ /0 8QLFHOOXODU DHURELF QLWURJHQIL[LQJ F\DQREDFWHULD RI WKH JHQXV RI Cyanothece. -%DFWHULRO 5HGG\ .- 0DVDPRWR . 6KHUPDQ '0 DQG 6KHUPDQ /$ '1$ VHTXHQFH DQG UHJXODWLRQ RI WKH JHQH cbpA HQFRGLQJ WKH NLORGDOWRQ F\WRSODVPLF PHPEUDQH FDURWHQRSURWHLQ RI WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ3&& -%DFWHULRO 5HLQKROG/.RVORII5DQG.DSODQ$ $PRGHOIRULQRUJDQLF FDUERQ IOX[HV DQG SKRWRV\QWKHVLV LQ F\DQREDFWHULDO FDUER[\VRPHV&DQ-%RW 5HLQKROG / 9RORNLWD 0 =HQYLUWK ' DQG .DSODQ $ ,V +&2 WUDQVSRUW LQAnabaena D 1DV\PSRUW" 3ODQW 3K\VLRO 5HLQKROG / =YLPDQ 0 DQG .DSODQ $ 4XDQWLWDWLYH PRGHO IRULQRUJDQLFFDUERQIOX[HVDQGSKRWRV\QWKHVLVLQF\DQREDFWHULD 3ODQW3K\VLRO%LRFKHP 5HLWKPDQ + %XOOHMDKQ * 5HGG\ .- DQG 6KHUPDQ /$ 5HJXODWLRQ RI F\DQREDFWHULDO SLJPHQWSURWHLQ FRPSRVLWLRQ DQG RUJDQL]DWLRQ E\ HQYLURQPHQWDO IDFWRUV 3KRWRV\Q 5HV 5HXWHU: DQG 0XHOOHU & $GDSWDWLRQRIWKH SKRWRV\QWKHWLF DSSDUDWXV RI F\DQREDFWHULD WR OLJKW DQG FDUERQ GLR[LGH - 3KRWRFKHP 3KRWRELRO 5H\HV -& DQG )ORUHQFLR )- $ QHZ W\SH RI JOXWDPLQH V\QWKHWDVH LQ F\DQREDFWHULD 7KH SURWHLQ HQFRGHG E\ WKH glnN JHQH VXSSRUWV QLWURJHQ DVVLPLODWLRQ LQ Synechocystis VS VWUDLQ 3&&-%DFWHULRO 5H\HV -& DQG )ORUHQFLR )- D (OHFWURQ WUDQVSRUW FRQWUROV WUDQVFULSWLRQ RIWKH JOXWDPLQH V\QWKHWDVH JHQH glnA IURP WKH F\DQREDFWHULXP Synechocystis VS 3&& 3ODQW 0RO %LRO 5H\HV -& DQG )ORUHQFLR )- E $ QRYHO PHFKDQLVP RI JOXWDPLQH V\QWKHWDVH LQDFWLYDWLRQ E\ DPPRQLXP LQ WKH F\DQREDFWHULXP Synechocystis VS 3&& ,QYROYHPHQWRIDQ LQDFWLYDWLQJSURWHLQ)(%6/HWW 5RGULJXH] 5 *DUFLD*RQ]DOH] 0 *XHUUHUR 0* DQG /DUD & $PPRQLXPVHQVLWLYH SURWHLQ NLQDVH DFWLYLW\ LQ SODVPD PHPEUDQHV RI WKH F\DQREDFWHULXP Anacystis nidulans. )(%6 /HWW 5RGULJXH]5/DUD&DQG*XHUUHUR0* 1LWUDWHWUDQVSRUWLQ WKH F\DQREDFWHULXP Anacystis nidulans 5 .LQHWLF DQG HQHUJHWLFDVSHFWV%LRFKHP- 5RPDQV . &DUSHQWHU (- DQG %HUJPDQ % %RX\DQF\ UHJXODWLRQ LQ WKH FRORQLDO GLD]RWURSKLF F\DQREDFWHULXP Trichodesmium tenue: 8OWUDVWUXFWXUH DQG VWRUDJH RI FDUERK\GUDWH SRO\SKRVSKDWH DQG QLWURJHQ - 3K\FRO 5RQHQ7DUD]L0/LHPDQ+XQYLW]-*DED\&2UXV0DQG.DSODQ $ 7KH JHQRPLF UHJLRQ RI rbcLS RI Synechococcus VS VWUDLQ3&& FRQWDLQV JHQHV LQYROYHG LQ WKHDELOLW\WR JURZ XQGHU ORZ &2 FRQFHQWUDWLRQV DQG LQ FKORURSK\OO ELRV\QWKHVLV 3ODQW3K\VLRO 5RQHQ7DUD]L 0 6FKZDU] 5 %RXHYLWFK $ /LHPDQ+XQYLW] - (UH] - DQG .DSODQ $ E 5HVSRQVH RI SKRWRV\QWKHWLF PLFURRUJDQLVPV WR FKDQJLQJ UHVSRQVH RI DPELHQW FRQFHQWUDWLRQ
RI&2 ,Q -RLQW,HG 0ROHFXODU(FRORJ\RI$TXDWLF0LFUREHV 1$72$6,6HULHV9RO* 5XELR/0 +HUUHUR$DQG)ORUHV( $F\DQREDFWHULDO narB JHQH HQFRGHV D IHUUHGR[LQGHSHQGHQW QLWUDWH UHGXFWDVH 3ODQW 0RO %LRO 6DNDPRWR7 'HOJDL]R9% DQG%U\DQW'$ *URZWKRQXUHD FDQ WULJJHU GHDWK DQG SHUR[LGDWLRQ RI WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& $SSO (QYLURQ 0LFURELRO 6DOHKLDQ 2 DQG %UXFH ' 'LVWULEXWLRQ RIH[FLWDWLRQ HQHUJ\ LQ SKRWRV\QWKHVLV 4XDQWLILFDWLRQ RI IOXRUHVFHQFH \LHOGV IURP LQWDFWF\DQREDFWHULD-RXUQDO/XPLQ 6DPXHOVVRQ * /RQQHKRUJ $ *XVWDIVVRQ 3 DQG 2TXLVW * 7KH VXVFHSWLELOLW\ RISKRWRV\QWKHVLV WR SKRWRLQKLELWLRQ DQG WKH FDSDFLW\ RIUHFRYHU\ LQ KLJK DQGORZ OLJKWJURZQ F\DQREDFWHULD $QDF\VWLV nidulans. 3ODQW3K\VLRO 6DQGHUV&(0HOLV$DQG$OOHQ-) ,QYLYRSKRVSKRU\ODWLRQ RI SURWHLQV LQ WKH F\DQREDFWHULXP Synechococcus DIWHU FKURPDWLF DFFOLPDWLRQWRSKRWRV\VWHP,RUSKRWRV\VWHP ,, OLJKW %LRFKLP%LRSK\V$FWD 6DWRK.DQG)RUN'& $QHZPHFKDQLVPIRUDGDSWDWLRQ WR FKDQJHV LQ OLJKW LQWHQVLW\ DQGTXDOLW\ LQ WKHUHG DOJD Porphyra perforata. , 5HODWLRQ WR VWDWHOVWDWH WUDQVLWLRQV %LRFKLP %LRSK\V$FWD 6FDQODQ '- 0DQQ 1+ DQG &DUU 1* 7KH UHVSRQVH RIWKH SLFRSODQNWRQLF PDULQH F\DQREDFWHULXP Synechococcus sp. :+ WRSKRVSKDWHVWDUYDWLRQLQYROYHVDSURWHLQKRPRORJRXV WR WKH SHULSODVPLF SKRVSKDWHELQGLQJ SURWHLQ RI Escherichia coli. 0RO0LFURELRO 6FKDHIHU 05 DQG *ROGHQ 66 'LIIHUHQWLDO H[SUHVVLRQ RI PHPEHUV RI D F\DQREDFWHULDO SVE$ JHQH IDPLO\ LQ UHVSRQVH WR OLJKW-%DFWHULRO 6FKOXFKWHU : =KDR - DQG %U\DQW '$ ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI WKH ndhF JHQH RI Synechococcus VS VWUDLQ 3&&DQGLQLWLDOFKDUDFWHUL]DWLRQRIDQLQWHUSRVRQPXWDQW - %DFWHULRO 6FKPLGW $ DQG &KULVWHQ 8 $ IDFWRUGHSHQGHQW VXOIRWUDQVIHUDVH VSHFLILF IRU SKRVSKRDGHQRVLQH SKRVSKRVXOIDWH 3$36 LQ WKH F\DQREDFWHULXP Synechococcus 3ODQWD 6FKPLGW $ (UGOH , DQG .RVW +3 &KDQJHV RI F SK\FRF\DQLQ LQ Synechococcus LQ UHODWLRQ WR JURZWK RQ YDULRXVVXOIXUFRPSRXQGV=1DWXUIRUVFKF 6FKPLGW*RII&0 DQG )HGHUVSLHO 1$ ,QYLYR DQG LQ YLWUR IRRWSULQWLQJRIDOLJKWUHJXODWHGSURPRWHULQ WKHF\DQREDFWHULXP Fremyella diplosiphon. -%DFWHULRO 6FKZDU]5)ULHGEHUJ'5HLQKROG/DQG.DSODQ$ ,VWKHUH D UROH IRUWKHN'DSRO\SHSWLGH LQ LQRUJDQLF FDUERQ XSWDNHE\ F\DQREDFWHULD3ODQW3K\VLRO 6FKZDU] 5 DQG *URVVPDQ $5 $ UHVSRQVH UHJXODWRU RI F\DQREDFWHULD LQWHJUDWHV GLYHUVH HQYLURQPHQWDO VLJQDOV DQG LV FULWLFDO IRU VXUYLYDO XQGHU H[WUHPH FRQGLWLRQV 3URF 1DWO $FDG 6FL86$,Q SUHVV 6FKZDU]5 /LHPDQ+XUZLW]- +DVVLGLP0 DQG.DSODQ$ 3KHQRW\SLF FRPSOHPHQWDWLRQ RIKLJK&2UHTXLULQJ PXWDQWV RI WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& E\ LQRVLQH PRQRSKRVSKDWH3ODQW3K\VLRO 6FKZDU]55HLQKROG/DQG.DSODQ$ /RZDFWLYDWLRQVWDWH RI ULEXORVH ELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH LQ D FDUER[\VRPHGHIHFWLYH Synechococcus PXWDQW 3ODQW 3K\VLRO 6HDUOH *): %DUEHU - 3RUWHU * DQG 7UHGZHOO &- 3LFRVHFRQG WLPHUHVROYHG HQHUJ\ WUDQVIHU LQ Porphyridium
'%KD\D56FKZDU]DQG$5*URVVPDQ cruentum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doliolum DQG Anacystis nidulans. ,QGLDQ-([S%LRO 6REF]\N$%HO\$7DQGHDXGH0DUVDF1DQG+RXPDUG- $ SKRVSKRU\ODWHG '1$ ELQGLQJ SURWHLQ LV VSHFLILF IRU WKH UHG OLJKW VLJQDO GXULQJ FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ LQ F\DQREDFWHULD0RO0LFURELRO 6REF]\N $ 6FK\QV * 7DQGHDX GH 0DUVDF 1 DQG +RXPDUG - 7UDQVGXFWLRQ RI WKH OLJKW VLJQDO GXULQJ FRPSOHPHQWDU\ FKURPDWLF DGDSWDWLRQ LQ WKH F\DQREDFWHULXP Calothrix VS 3&& '1$ELQGLQJ SURWHLQV DQG PRGXODWLRQ E\ SKRVSKRU\ODWLRQ(0%2- 6WHZDUW:'3DQG/H[ 0 1LWURJHQDVH DFWLYLW\ LQWKHEOXH JUHHQDOJDPlectonema boryanum VWUDLQ $UFK 0LFURELRO 6XOWHPH\HU ' 3ULFH *' <X -: DQG %DGJHU 05 &KDUDFWHUL]DWLRQ RI FDUERQ GLR[LGH DQG ELFDUERQDWH WUDQVSRUW GXULQJVWHDG\VWDWHSKRWRV\QWKHVLVLQWKHPDULQHF\DQREDFWHULXP Synechococcus VWUDLQ3&&3ODQWD 6X]XNL ( )XNX]DZD + $EH 7 DQG 0L\DFKL 6 ,GHQWLILFDWLRQ RI WKH JHQRPLF UHJLRQ ZKLFK FRPSOHPHQWV D WHPSHUDWXUHVHQVLWLYH KLJK &2UHTXLULQJ PXWDQW RI WKH F\DQREDFWHULXP Synechococcus 3&& 0ROHF *HQ *HQHW 6X]XNL , +RULH 1 6XJL\DPD 7 DQG 2PDWD 7 D ,GHQWLILFDWLRQ DQG FKDUDFWHUL]DWLRQ RI WZR QLWURJHQUHJXODWHG JHQHV RI WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& UHTXLUHG IRUPD[LPXPHIILFLHQF\ RIQLWURJHQ DVVLPLODWLRQ -%DFWHULRO 6X]XNL , .LNXFKL + 1DNDQLVKL 6 )XMLWD < 6XJL\DPD 7 DQG 2PDWD 7 E $ QRYHO QLWUDWH UHGXFWDVH JHQH IURP WKH F\DQREDFWHULXP Plectonema boryanum. - %DFWHULRO 6X]XNL , 2PDWD7 DQG 6XJL\DPD7 *HQH H[SUHVVLRQ DQG UHJXODWLRQ RI QLWUDWH DVVLPLODWLQJ HQ]\PHV LQ Synechococcus 3&& ,Q 0XUDWD 1 HG 5HVHDUFK LQ 3KRWRV\QWKHVLV 3URFHHGLQJV RI WKH ,;WK ,QWHUQDWLRQDO &RQJUHVV RQ 3KRWRV\QWKHVLV9ROIV, SS .OXZHU'RUGUHFKW 6X]XNL , 6XJL\DPD7DQG 2PDWD 7 3ULPDU\VWUXFWXUH DQG WUDQVFULSWLRQDO UHJXODWLRQ RIWKH JHQH IRU QLWUDWH UHGXFWDVH IURP WKH F\DQREDFWHULXP Synechococcus 3&& 3ODQW &HOO 3K\VLRO 6X]XNL , 6XJL\DPD7DQG2PDWD7 5HJXODWLRQE\F\DQDWH RIWKHJHQHVLQYROYHGLQFDUERQ DQG QLWURJHQ DVVLPLODWLRQ LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ3&&- %DFWHULRO 6ZDQVRQ 59 =KRX - Leary -$ :LOOLDPV 7 GH /RULPLHU 5 %U\DQW '$ DQG *OD]HU $1 &KDUDFWHUL]DWLRQ RI SK\FRF\DQLQ SURGXFHG E\ cpcE DQG cpcF PXWDQWV DQG
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
9HJD3DODV 0$ )ORUHV ( DQG +HUUHUR $ 1WF$ D JOREDO QLWURJHQ UHJXODWRU IURP WKH F\DQREDFWHULXP Synechococcus WKDW EHORQJV WRWKH&US IDPLO\RIEDFWHULDOUHJXODWRUV 0RO0LFURELRO 9HJD3DODV 0$ 0DGXHxR ) +HUUHUR $ DQG )ORUHV ( ,GHQWLILFDWLRQ DQG FORQLQJ RI D UHJXODWRU\ JHQH IRU QLWURJHQ DVVLPLODWLRQ LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&&-%DFWHULRO 9HUPDDV: $SSURDFKHVWRDQDO\]HSKRWRV\VWHPVWUXFWXUH DQGIXQFWLRQ $QQX5HY RI3ODQW3K\VLRO3ODQW0ROHF %LRO 9RJHOPDQQ 7& DQG 6FKHLEH - $FWLRQ VSHFWUXP IRU FKURPDWLF DGDSWDWLRQ LQ WKH EOXHJUHHQ DOJD Fremyella diplosiphon. 3ODQWD 9RORNLWD0=HQYLUWK' .DSODQ$DQG 5HLQKROG/ 1DWXUH RI WKH LQRUJDQLF FDUERQ VSHFLHV DFWLYHO\ WDNHQ XS E\ WKH F\DQREDFWHULXP Anabaena variabilis. 3ODQW 3K\VLRO :DJQHU) 0DULR* $KRUQ +3HVFKHN *$ DQG)DONQHU* ,VRODWLRQ DQG IXQFWLRQDO UHFRQVWLWXWLRQ RI D SKRVSKDWH ELQGLQJ SURWHLQ RI WKH F\DQREDFWHULXP Anacystis nidulans LQGXFHG GXULQJSKRVSKDWHOLPLWHGJURZWK-%LRO&KHP :DJQHU .8 0DVHSRKO % DQG 3LVWRULXV (. 7KH F\DQREDFWHULXP Synechococcus VS VWUDLQ 3&& FRQWDLQV D VHFRQG DONDOLQH SKRVSKDWDVH HQFRGHG E\ phoV. 0LFURELRO :DJQHU6- 7KRPDV 63 .DXIPDQ 5, 1L[RQ %7 DQG 6WHYHQV 6( 7KH glnA JHQH RI WKH F\DQREDFWHULXP Agmenellum quadruplicatum 35 LV QRQHVVHQWLDO IRU DPPRQLXP DVVLPLODWLRQ-%DFWHULRO :DQQHU %/ *HQH UHJXODWLRQ E\ SKRVSKDWH LQ HQWHULF EDFWHULD-&HOO%LRFKHP :DQQHU%/D 0XOWLSOHFRQWUROV RIWKH Escherichia coli 3KR UHJXORQ E\ WKH 3L VHQVRU 3KR5 WKH FDWDEROLWH UHJXODWRU\ VHQVRU &UH&DQGDFHW\OSKRVSKDWH ,Q 7RUULDQL*RULQL $
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nabaena. %DFWHULRO :\DWW-7DQG6LOYH\-.* 1LWURJHQIL[DWLRQE\Gloeocapsa. 6FLHQFH :\PDQ 0 *UHJRU\ 53) DQG &DUU 1* 1RYHO UROH IRU SK\FRH,\WKULQ LQ D PDULQH F\DQREDFWHULXP Synechococcus VWUDLQ'&6FLHQFH :\PDQ 0 =HKU-3 DQG &DSRQH '* 7HPSRUDO YDULDELOLW\ LQ QLWURJHQDVH JHQH H[SUHVVLRQ LQ QDWXUDO SRSXODWLRQV RI WKH PDULQH F\DQREDFWHULXP Trichodesmium thiebautii. $SSO (QYLURQ0LFURELRO :\QQH' $OWHUDWLRQVLQDFWLYLW\RISKRVSKDWDVHVGXULQJWKH Pertdinium EORRPLQ/DNH.LQQHUHW3K\VLRO3ODQW ;LH:4 :KLWWRQ%$ 6LPRQ-: -DJHU.5HHG' DQG 3RWWV 0 Nostoc commune 87(; JHQH H[SUHVVLQJ LQGROH SKRVSKDWH K\GURODVH DFWLYLW\ LQ Escherichia coli. - %DFWHULRO
' %KD\D 56FKZDU]DQG$5*URVVPDQ Synechococcus VS 3&& ,Q 0XUDWD1 HG 5HVHDUFKLQ 3KRWRV\QWKHVLV9RO , SS .OXZHU'RUGUHFKW =KRX - *DVSDULFK *( 6WLUHZDOW 9 / GH /RULPLHU5 DQG %U\DQW '$ 7KHcpcE DQG cpcF JHQHV RISynechococcus VS 3&& &RQVWUXFWLRQ DQG SKHQRW\SLF FKDUDFWHUL]DWLRQ RI LQWHUSRVRQPXWDQWV-%LRO&KHP =LPPHUPDQQ + QXFOHRWLGDVH PROHFXODU VWUXFWXUH DQG IXQFWLRQDODVSHFWV%LRFKHP-
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hitton and M. Potts (eds). The Ecology of Cyanobacteria. . Kluwer Academic Publishers . Printed in the Netherlands
1- 5RELQVRQ-& 5XWKHUIRUG05 3RFRFNDQG-6&DYHW
'1$RIPDQ\F\DQREDFWHULD ,WLVSUHGLFWHGWKDWWKHVWXG\RIF\DQREDFWHULDZLOODOVRJLYHXQLTXHLQVLJKWVLQWRWKH HYROXWLRQRIUHSHWLWLYH'1$
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~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ynechococcus 3&&KDVSUHFLVH]LQFKRPRHRVWDVLVPHGLDWHGE\ PHWDOORWKLRQHLQDQGLQWKLVRUJDQLVP]LQFLVH[SORLWHG LQ]LQFILQJHUVIRUH[DPSOHLQ'1$SULPDVH6HFWLRQ 'O %\ FRQWUDVW '1$ SULPDVH LQ Synechocystis 3&& GRHV QRWFRQWDLQ DFDUER[\WHUPLQDO ]LQF ILQJHU DQG QR PHWDOORWKLRQHLQ JHQH KDV EHHQ LGHQWLILHG ZLWKLQ WKH Synechocystis 3&& JHQRPH DW OHDVW EDVHG XSRQ VHTXHQFH VLPLODULW\ :KHWKHU RU QRW Synechocystis 3&& H[SORLWV ]LQF DQG ]LQFILQJHUV WR DQ\ OHVVHU H[WHQW WKDQ Synechococcus 3&& LV SUHVHQWO\ XQNQRZQ 6HFWLRQ)
& 6SDWLDO 'LVWULEXWLRQ RI 0HWDOV DQG 0HWDO 5HVLVWDQFHLQ &\DQREDFWHULD 0HWDO3ROOXWLRQDQG 9DULDWLRQLQ0HWDO /HYHOV 3ROOXWLRQ RIDTXDWLF HQYLURQPHQWV ZLWK PHWDOV RIWKH FRSSHUDQG]LQFWULDGVEHFDPHFRPPRQNQRZOHGJHLQ WKH V IROORZLQJ WZR LQFLGHQWV ERWK LQ -DSDQ FLWHG LQ 0DVRQ &ODUN 7KH ILUVW UHVXOWHG IURPWKH XVH RIPHUFXULF R[LGH DV D FDWDO\VW LQ WKH SURGXFWLRQ RI DFHWDOGHK\GH E\ D IDFWRU\ DW 0LQLPDWD%D\ 0HUFXU\ GLVFKDUJHG LQWR WKH ED\ OHG WRWKHVXEVHTXHQWDFFXPXODWLRQRIWKHVHPHWDOLRQV LQ
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¶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halassiosira weissflogii JURZLQJ LQ KLJKO\ ]LQF GHSOHWHG ZDWHUV 3ULFH DQG 0RUHO +RZHYHU DV SUHYLRXVO\ QRWHGFDGPLXPLVJHQHUDOO\WKRXJKWWREHH[FOXVLYHO\ WR[LFDQGQRWXVHGLQELRORJ\DQGLQGHHGWKHUHDUHQR GRFXPHQWHG FDVHV RI WKH XVH RI FDGPLXP LQ F\DQREDFWHULD ([SRVXUH WR HOHYDWHG FDGPLXP LV JHQHUDOO\ WKRXJKWWR KDYH µRQO\¶ RFFXUUHG DV DUHVXOW RI PDQ¶V LQIOXHQFHV RQ HQYLURQPHQWV 7KXV ZKHUH FDGPLXPVHQVLWLYHSKHQRW\SHVKDYHEHHQ REVHUYHGLQ JHQH GHOHWLRQ PXWDQWV IRU H[DPSOH 7XUQHU HW DO WKLV KDV LQYDULDEO\ EHHQ DWWULEXWHG WR WKH µJUDWXLWRXV¶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
1- 5RELQVRQ-& 5XWKHUIRUG05 3RFRFNDQG-6&DYHW
WHPSRUDOO\ DQG WKHVH FKDQJHV FDQ RSHUDWH RYHU UHODWLYHO\VKRUWGLVWDQFHVDGMDFHQWWRRUGLVWDQWIURP SDUWLFOHV DQG WLPHVFDOHV IRU H[DPSOH ZLWK UDSLG FKDQJHVLQVRLOPRLVWXUHFRQWHQW ,WLVRIFRXUVHWKH SRWHQWLDO IRU PRYHPHQWWR PLFUR QLFKHV ZKHUH PHWDO DYDLODELOLW\ FKDQJHV DQG WKH SRWHQWLDO IRU VZLIW IOXFWXDWLRQV LQPHWDO DYDLODELOLW\ WKDWVHOHFW IRUPHWDO SHUFHSWLRQV\VWHPV6HFWLRQ(
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ynechococcus 3&& DQG FHOO HORQJDWLRQ &KLQWDPDQL DQG0RKDQW\ :KLWWRQ DQG 6KHKDWD 6KHKDWD DQG :KLWWRQ ,Q FRQWUDVW HOHYDWHG FRSSHU KDV EHHQ VKRZQ WR FDXVH SURGXFWLRQ RI VXEVSKHULFDO UDWKHU WKDQ FRFFRLG Synechococcus 3&& :KLWWRQ DQG 6KHKDWD (OHYDWHG FRQFHQWUDWLRQV RI PHWDOV LQIOXHQFH SLJPHQW FRPSRVLWLRQ IRU H[DPSOH ZLWK LQFUHDVHG FDURWHQRLG WR FKORURSK\OO UDWLRV DQG RYHUDOO GHFUHDVHV LQ SK\FRF\DQLQ FKORURSK\OO a DQG WRWDO SURWHLQ LQ Synechococcus 3&& H[SRVHG WR HOHYDWHG ]LQF &KLQWDPDQLDQG0RKDQW\ $QDQDO\VLVRIWKH HIIHFWV RI HOHYDWHG FRQFHQWUDWLRQV RI FRSSHU RQ Nostoc calcicola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
FRQFHQWUDWLRQV6LQJKDQG3DQGH\ *XSWDHWDO (QKDQFHGUHVLVWDQFHLQVXFKDFRSSHUUHVLVWDQW OLQHRINostoc calcicola ZDVFRUUHODWHGZLWKHQKDQFHG FRSSHU HIIOX[ ZKLOH HQKDQFHG FDGPLXP UHVLVWDQFH LQ Synechococcus 3&& ZDV FRUUHODWHG ZLWK HQKDQFHG HQGRJHQRXV PHWDO FKHODWLRQ PHGLDWHG E\ LQFUHDVHGH[SUHVVLRQRIPHWDOORWKLRQHLQ6HFWLRQ'
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nacystis nidulans (Synechococcus 3&& RU H[SRVHG WR FDGPLXP 0DFOHDQ HW DO 6XEVHTXHQWO\ 07V ZHUH SXULILHG IURP D PDULQH F\DQREDFWHULXP Synechococcus 55,03 1, 2ODIVRQ HW DO DQG IUHVKZDWHU VWUDLQV GHVLJQDWHG Synechococcus 87(; DQG Synechococcus 7; DJDLQ ERWK VWUDLQV DUH WKRXJKW WR EH V\QRQ\PRXV ZLWK RU DW OHDVW YHU\ FORVHO\ UHODWHG WR Synechococcus 3&& DQG 7KH SURWHLQ IURP Synechococcus 7; ZDV VXEVHTXHQWO\ VHTXHQFHG DQG VSHFWURVFRSLF VWXGLHV LQGLFDWHG WKH SUHVHQFH RI D VLQJOH PHWDO FOXVWHU LQYROYLQJ FRRUGLQDWLRQ RIPHWDO LRQV WR WKLRO JURXSV RIF\VWHLQH UHVLGXHV 2ODIVRQ HW DO )UDJPHQWV RI WKH FRUUHVSRQGLQJ JHQHV ZHUH REWDLQHG IURP '1$ SXULILHG IURP Synechococcus 3&& DQG Synechococcus 3&& YLD3&5 XVLQJ GHJHQHUDWH SULPHUV 5RELQVRQ HW DO 7KHVH 3&5 IUDJPHQWV ZHUH XVHG DV SUREHV WR LVRODWH WKH FRUUHVSRQGLQJ JHQRPLF UHJLRQ DQG WKH JHQH GHVLJQDWHG smtA +XFNOH HW DO $ UHODWHG
&KDSWHU 0HWDOV5HSHWLWLYH'1$ JHQH GHVLJQDWHG mtnA, KDV DOVR EHHQ LGHQWLILHG LQ '1$ VHTXHQFHV IURP Synechococcus vulcanus 6KLPL]X HW DO 7R GDWH QR KRPRORJXH RI smtA KDV EHHQ GHWHFWHG LQ WKH IXOO\ VHTXHQFHG JHQRPH RI Synechocystis 3&& +RZHYHU WKH VPDOO VL]H RI WKH DQWLFLSDWHG RSHQ UHDGLQJ IUDPH 25) DQG WKH UHODWLYH ODFN RI VHTXHQFH KRPRORJ\ DPRQJFODVV,,07VPHDQWKDWVXFKDQ25)FRXOGEH HDVLO\RYHUORRNHG 2ODIVRQ DQG FRZRUNHUV QRWHG WKDW 07 DFFXPXODWHG LQ Synechococcus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in vivo DQG PXWDQWVGHILFLHQWLQWKHFRUUHVSRQGLQJ JHQHVIRU H[DPSOH CUP1 LQ Saccharomyces cerevisiae) DUH K\SHUVHQVLWLYH WR H[FHVV FRSSHU EXW QRW ]LQF 7KH smtA JHQH KDV EHHQ RYHUH[SUHVVHG LQ Escherichia coli WR JHQHUDWH D UHFRPELQDQW IXVLRQ SURWHLQ 6KL HW DO 6PW$ DVVRFLDWHG ZLWK DOO RIWKH DQDO\VHG PHWDO LRQV ]LQF FDGPLXP FRSSHU DQG PHUFXU\ IROORZLQJ SXULILFDWLRQ IURP E. coli JURZQ LQ PHWDO VXSSOHPHQWHGPHGLD 5HODWLYH WR HTXLQHUHQDO 07 DIILQLWLHV IRU WKHVH PHWDOV ZHUH HVWLPDWHG E\ GLVSODFLQJ PHWDO LRQV ZLWKSURWRQV DWORZS+ 6KL HW DO 0RUH SURWRQV ZHUH UHTXLUHG WR GLVSODFH ]LQF IURP F\DQREDFWHULDO 07 WKDQ IURP HTXLQH UHQDO 07VXJJHVWLQJWKDWWKHIRUPHUKDV DSDUWLFXODUO\KLJK DIILQLW\ IRU ]LQF (YLGHQFH WKDW F\DQREDFWHULD FRQWDLQLQJ smtA DFFXPXODWH PRUH ]LQF WKDQ PXWDQWV GHILFLHQW LQ smtA DOVR VXSSRUWV WKH SURSRVDO WKDW 6PW$ ELQGV ]LQF ZLWKLQ F\DQREDFWHULDO FHOOV 7XUQHU HWDO 0XWDQWV RI Synechococcus 3&& ZKLFK DUH GHILFLHQW LQ D IXQFWLRQDO 07 JHQH KDYH EHHQ FUHDWHG E\ KRPRORJRXV UHFRPELQDWLRQ PHGLDWHG JHQH LQVHUWLRQ 7XUQHU HW DO 7KHVH FHOOV DUH DSSUR[LPDWHO\ IROG PRUH VHQVLWLYH WR H[FHVV ]LQF VKRZ VRPH UHGXFHG WROHUDQFH WR FDGPLXP EXW KDYH
QRUPDO OHYHOV RI WROHUDQFH WR H[FHVV FRSSHU 7KH smtA JHQH KDV EHHQ XVHG DV D VHOHFWDEOH PDUNHU WR VHOHFW IRU WUDQVIRUPDQWV RI07 GHILFLHQW FHOOV RQ WKH EDVLVRIWKHLUUHVWRUHGUHVLVWDQFHWR]LQF7XUQHUHWDO
7ZR GLIIHUHQW W\SHV RI PXWDWLRQV ZKLFK DIIHFW WKH smtA JHQH KDYH EHHQ FKDUDFWHULVHG LQ FDGPLXP WROHUDQW F\DQREDFWHULD ,Q ERWK FDVHV WKHVH ZHUH GHWHFWHG LQ FXOWXUHG FHOOV VHOHFWHG WR JURZ LQ PHGLD VHTXHQWLDOO\ HQULFKHG ZLWK LQFUHDVLQJ FRQFHQWUDWLRQV RI FDGPLXP )LUVW DQ LQFUHDVH LQ WKH QXPEHU RI FRSLHV RI WKH smtA JHQH *XSWD HW DO DQG VHFRQG WKH GHOHWLRQ RI D UHSUHVVRU VHH IROORZLQJ VHFWLRQV *XSWD HW DO %RWK PXWDWLRQV DUH DVVXPHG WR FRQIHU DQ LQFUHDVH LQ WKH H[SUHVVLRQ RI smtA DQG KHQFH WKH DPRXQW RIPHWDO VHTXHVWHUHG ,W ZLOO EH RI LQWHUHVW WR H[DPLQH ZKHWKHU VLPLODU PXWDWLRQVKDYHRFFXUUHGLQF\DQREDFWHULDVHOHFWHGIRU UHVLVWDQFH WR FDGPLXP RU ]LQF LQ PHWDO SROOXWHG
QDWXUDO HQYLURQPHQWV 8SVWUHDP RIsmtA LV WKH '1$ SULPDVH JHQH dnaG %LUGHWDO '1$SULPDVH LV UHTXLUHG IRU WKH V\QWKHVLV RIVKRUW 51$ SULPHUV WKDW DUH HVVHQWLDO IRU ERWK OHDGLQJ VWUDQG V\QWKHVLV DW WKH RULJLQ RI UHSOLFDWLRQ DQG IRU UHSHDWHG UHLQLWLDWLRQ RI ODJJLQJ VWUDQG '1$ V\QWKHVLV DW UHSOLFDWLRQ IRUNV ,QPDQ\ RWKHU EDFWHULD dnaG LV DW WKH FHQWUH RI D PDFURPROHFXODU V\QWKHVLV RSHURQ ZKLFK LV JHQHUDOO\ FRPSRVHG RI rpsU-dnaG-rpoD HQFRGLQJ SURWHLQV UHTXLUHG IRU WKH LQLWLDWLRQ RI V\QWKHVLV RI SURWHLQ '1$ DQG 51$ UHVSHFWLYHO\ 7KLV JHQH DUFKLWHFWXUH LVZLGHO\FRQVHUYHGDQGLWKDVEHHQVXJJHVWHGWKDWWKH FRRUGLQDWHG H[SUHVVLRQ ZLWKLQ DQ RSHURQ RI JHQHV UHTXLUHG IRU WKH V\QWKHVLV RI WKH NH\ LQIRUPDWLRQDO PDFURPROHFXOHVPXVWEHDGYDQWDJHRXV 9HUVDORYLFHW DO 7KH DEVHQFH RI VXFK DQ RSHURQ LQ Synechococcus 3&& FRXOG VXJJHVW WKDW DOWHUQDWLYH PHFKDQLVPV DUH LQYROYHG LQ WKH FRRUGLQDWLRQ RI PDFURPROHFXODU V\QWKHVLV LQ WKLV F\DQREDFWHULXPFRPSDUHGWRRWKHUDQDO\VHGEDFWHULD $W WKH DPLQRWHUPLQDO HQG RI DOO '1$ SULPDVHV LQFOXGLQJ WKH RQH IURP Synechococcus 3&& LV D FRQVHUYHG ]LQFELQGLQJ UHJLRQ 6WDPIRUG HW DO DQGWKHSXULILHGSULPDVHIURPE. coli KDV EHHQ VKRZQ WR FRQWDLQ WLJKWO\ ERXQG ]LQF *ULHS DQG /RNH\ +RZHYHU DWWKHFDUER[\WHUPLQDOHQG RI WKH Synechococcus 3&& SULPDVH LV DQ DGGLWLRQDOHXNDU\RWLFVW\OH]LQFILQJHUZKLFKLVDEVHQW IURP DOO RWKHU UHSRUWHG SULPDVHV %LUG HW DO ,Q E. coli, WKH H[WUHPH FDUER[\WHUPLQXV RISULPDVH WKH ODVW DPLQR DFLGV LQWHUDFWV ZLWK WKHUHSOLFDWLYH KHOLFDVHDQGLWLVWKHVWUHQJWKRIWKLVLQWHUDFWLRQZKLFK
VHWV WKH UHSOLFDWLRQ IRUN FORFN 7RXJX DQG 0DULDQV '1$ UHSOLFDWLRQ DQG FHOO GLYLVLRQ LQ Synechococcus 3&& GR QRW ILW WKH SURNDU\RWLF SDUDGLJP %LQGHU DQG &KLVKROP ZLWK DV\QFKURQRXV LQLWLDWLRQ RI FKURPRVRPH UHSOLFDWLRQ
1- 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
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in vitro =HQJ HW DO D E &DQR*DXFL DQG 6DUNHU $ JOREDO UHJXODWRU\ UROH IRU DQLPDO 07 PRGLI\LQJ WKH DYDLODELOLW\ RI ]LQF KDV EHHQVXJJHVWHG &RXOGVXFKDUROHIRU07DQGLQGHHG D UHJXODWRU\ UROH IRU ]LQF KDYH ILUVW HPHUJHG LQ F\DQREDFWHULD" %\ DQDORJ\ WR SURSRVHG UROHV IRU 07V LQKLJKHUHXNDU\RWHV FKDQJHV LQWKH H[SUHVVLRQ RI07LQSynechococcus 3&&FRXOGPRGLI\WKH DYDLODELOLW\RI]LQFDQGKHQFHUHJXODWH]LQFRFFXSDQF\ RIWKH]LQFILQJHULQ'1$SULPDVH ,W LV QRWDEOH WKDW D JHQH HQFRGLQJ D VXEXQLW RI SKHQ\ODQ\O W51$ V\QWKHWDVH pheT, LV FRWUDQVFULEHG ZLWK dnaG LQ Synechococcus 3&& %LUG HW DO 3KHQ\ODODQ\O W51$ V\QWKHWDVH LQ FRPPRQ ZLWK PDQ\ RWKHU DPLQRDF\O W51$ V\QWKHWDVHV QRW RQO\ FDWDO\VHV WKH DVVRFLDWLRQ RIDQ DPLQR DFLG ZLWK LWV FRJQDWH W51$ PROHFXOH EXW DOVR FDWDO\VHV WKH SURGXFWLRQRIGLDGHQRVLQHWHWUDSKRVSKDWH $S$ LQD ]LQF GHSHQGHQW PDQQHU 3ODWHDX HW DO &KDQJHV LQ ]LQF DYDLODELOLW\ PHGLDWHG E\ 6PW$ DUH WKHUHIRUH OLNHO\ WR PRGXODWH SURGXFWLRQ RI $S$ D NQRZQUHJXODWRURIWKH FHOO F\FOH 1LVKLPXUD DQGKHQFHUDWHVRIFHOOSUROLIHUDWLRQ
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
DUH NQRZQ WR VSHFLILFDOO\ WUDQVSRUW WUDQVLWLRQ PHWDOV (QHUJ\GHSHQGHQWFRSSHUHIIOX[ ZDV SURSRVHG DV D PHFKDQLVPRIFRSSHUUHVLVWDQFHLQ Synechococcus VS 2ODIVRQ DQG LQNostoc calcicola 9HUPDDQG 6LQJK 6XEVHTXHQWO\ JHQHV HQFRGLQJFRSSHU WUDQVSRUWLQJ 3W\SH $73DVHV GHVLJQDWHG pacS .DQDPXUD HW DO DQG ctaA 3KXQJ HW DO KDYH EHHQ LVRODWHG VXJJHVWLQJ UROHV IRU WKLV FODVV RI PHPEUDQH SURWHLQV LQ FRSSHU DFTXLVLWLRQ GLVWULEXWLRQ DQG KRPRHRVWDVLV LQ DW OHDVW VRPH F\DQREDFWHULD .DQDPXUD DQG FRZRUNHUV FORQHGWZR JHQHV HQFRGLQJGHGXFHG 3W\SH$73DVHV QDPHG pacS DQG pacL. 3DF/ ZDV VKRZQ WR EH D FDOFLXPWUDQVSRUWLQJ $73DVH %HUNHOPDQ HW DO ZKLOH 3DF6 ZDV VKRZQ WR WUDQVSRUW FRSSHU .DQDPXUD HW DO 7KH pacS WUDQVFULSWV LQFUHDVH LQ DEXQGDQFH LQ UHVSRQVH WR HOHYDWHG FRQFHQWUDWLRQV RIFRSSHU DQG VLOYHULRQV DQG :HVWHUQ EORWV FRQILUP WKDW WKH 3DF6 SURWHLQ VLPLODUO\ DFFXPXODWHV LQ FRSSHU WUHDWHG Synechococcus 3&& $QLQVHUWLRQDOPXWDQWRIpac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ynechococcus 3&&&WD$3KXQJHWDO 0XWDQWV GHILFLHQW LQ IXQFWLRQDO ctaA VKRZ LQFUHDVHG WROHUDQFH WR FRSSHU FRPSDUHG WR ZLOG W\SH FHOOV VXJJHVWLQJ D UROH IRU &WD$ LQ WKH LPSRUW RI FRSSHU LRQV
=LQF([SRUWHU=LD$ DQGRWKHU +RPRORJXHV IURP6\QHFKRF\VWLV 3&& :LWKLQ WKH JHQRPH RI Synechocystis 3&& .DQHNR HW DO LV D FOXVWHU RI25)V HQFRGLQJ GHGXFHG SURWHLQV ZLWK VLPLODULW\ WR NQRZQ SURWHLQV LQYROYHG LQ FRSSHU DQG ]LQF KRPRHRVWDVLV LQ RWKHU EDFWHULD )LJ 7KLV LQFOXGHV VHTXHQFHV ZLWK VLPLODULW\ WR czcA DQG czcB. 7KHVH WZR JHQHV ZHUH RULJLQDOO\LVRODWHG IURPAlcaligenes eutrophus &+ 7KLVRUJDQLVPZDVUHFRYHUHG IURPD]LQFGHFDQWDWLRQ WDQN0HUJHD\HWDO DQG WKHSODVPLGHQFRGHG czc IRUFREDOW ]LQF DQGFDGPLXP FRQIHUVUHVLVWDQFH
&KDSWHU 0HWDOV5HSHWLWLYH '1$ WR WKHVH WKUHH PHWDO LRQV 7KH czc GHWHUPLQDQW LV FRPSRVHG RIWKUHH czcCBA 1LHV DQG 6LOYHU 'LHOV HW DO VWUXFWXUDO JHQHV 7KH &]F&%$ SURWHLQVFRPELQHLQDUDWLRWRIRUPDFDWLRQ
Fig. I. $ UHJLRQ RI WKH 6\QHFKRF\VWLV 3&& JHQRPH FRQWDLQLQJ D FOXVWHURIJHQHV HQFRGLQJ GHGXFHG SURWHLQV LQYROYHG LQ WKH WUDQVSRUW DQG SHUFHSWLRQ RI PHWDO LRQV ,Q DGGLWLRQ WR GHVLJQDWHG 25)V GHVFULEHG LQ WKH WH[W WKLV UHJLRQ FRQWDLQV D VHFRQG SDLU RIJHQHV HQFRGLQJ D GHGXFHG WZR FRPSRQHQW VHQVRU orfs DQG orfR ZKLFKFRXOGEHLQYROYHGLQPHWDOSHUFHSWLRQ
HIIOX[FRPSOH[1LHV DQG 6LOYHU 'HOHWLRQ RI czcC OHDGV WR ORVV RIFDGPLXP DQG FREDOW UHVLVWDQFH EXW WKH &]F%$ FRPSOH[ DORQH VWLOO FRQIHUV KLJK HIILFLHQF\ ]LQF HIIOX[ 1LHV HW DO 7KH SUHVHQFH RI KRPRORJXHV RI czcBA EXW QRW czcC LQ Synechocystis 3&& FRXOG WKHUHIRUH VXJJHVW DQ H[FOXVLYHUROHLQ]LQFHIIOX[ DOWKRXJKWKLVUHPDLQVWR EH WHVWHG ,Q DGGLWLRQ WKHUH DUH WZR GLYHUJHQWO\ WUDQVFULEHG IURPWKH czcOLNH JHQHVGHVLJQDWHGczcR DQG czcS )LJ ZKLFK VKRZ VLPLODULW\ WR WZR FRPSRQHQWDGDSWLYHUHVSRQVHUHJXODWRUV UHYLHZHG LQ 3DUNLQVRQ DQG .RIRLG 6HTXHQFHV HQFRGLQJ D KLVWLGLQH NLQDVH DQG D UHVSRQVH UHJXODWRU DUH DOVR SUHVHQW LQ VHTXHQFHV IODQNLQJ czc IURP Alcaligenes eutrophus DQG WKHVH DUH NQRZQ WR UHJXODWH WKH H[SUHVVLRQ RI WKH czc JHQHV YDQ GHU /HOLH HW DO (OVHZKHUH LQ WKH Synechocystis 3&& JHQRPHLVDQ25)HQFRGLQJDSURWHLQZLWKVLPLODULW\ WR&]F' &]F' LV LQYROYHG LQ WKH PHWDO UHVSRQVLYH H[SUHVVLRQ RI WKH czc GHWHUPLQDQW 1HLV DQG KDV VLPLODULW\ WR =5& DQG &27 ZKLFK FRQIHU ]LQFFDGPLXP DQG FREDOW UHVLVWDQFH UHVSHFWLYHO\ LQ Saccharomyces cerevisiae .DPL]RPR HW DO &RQNOLQHWDO 7ZR25)VZLWKLQWKLV FOXVWHUHQFRGH SURWHLQV ZLWK VHTXHQFH IHDWXUHV RI WUDQVLWLRQ PHWDOWUDQVSRUWLQJ &3[W\SH $73DVHV 2QH RI WKHVH ZDV LQLWLDOO\ GHVLJQDWHG 3DF6 GXH WR LWV VLPLODULW\ WR WKH FRSSHU WUDQVSRUWHU EXW DQ DQDO\VLV RI GHOHWLRQ PXWDQWV KDV UHYHDOHGVHQVLWLYLW\WR]LQFUHGXFHG]LQFH[SRUWWRWKH SHULSODVP DQG WKH JHQH KDV EHHQ UHGHVLJQDWHG ziaA 7KHOZHOOHW DO 7KH JHQH FRQIHUV HQKDQFHG UHVLVWDQFH WR ]LQF DQG UHGXFHG ]LQF DFFXPXODWLRQ ZKHQ LQWURGXFHG LQWR smt GHILFLHQW Synechococcus 3&& 7KHOZHOO HW DO 'LYHUJHQWO\
WUDQVFULEHG IURP ziaA LV DQ 25) HQFRGLQJ D SURWHLQ ZLWKVLPLODULW\WR6PW%VHHIROORZLQJVHFWLRQV 7KLV SURWHLQKDVEHHQVKRZQWREHDVHQVRURILQWHUQDO ]LQF FRQFHQWUDWLRQVDQGUHGHVLJQDWHG=LD57KHOZHOOHWDO )LJ 7KLV VXJJHVWV WKDW WKH FOXVWHU RI JHQHV FRXOG LQFOXGH WZR GLVWLQFW ]LQF WUDQVSRUWHUV ZLWK czc UHVSRQGLQJ WR SHULSODVPLF ]LQF OHYHOV PHGLDWHGE\ DWZRFRPSRQHQWVHQVRUDQGWKH$73DVH UHVSRQGLQJ WR LQWHUQDO ]LQF OHYHOV PHGLDWHG E\ DQ 6PW%OLNHPHWDOVHQVRU=LD5 (OVHZKHUH LQ WKH Synechocystis 3&& JHQRPH DUH RWKHU GHGXFHG PHWDO WUDQVSRUWHUV LQFOXGLQJ WKH czc'OLNH VHTXHQFH ,W LV QRZ QHFHVVDU\ WR GHOHWH WKHVH JHQHV DQG WKRVH LQ )LJ WR HVWDEOLVK WKH HIIHFWV RQ PHWDO WROHUDQFH DQG DFFXPXODWLRQ DQG UHVROYHZKLFKPHWDOVDUHWUDQVSRUWHG
7KH8VHRI=LQF 'HVSLWHWKHSOHWKRUDRINQRZQ ]LQFUHTXLULQJSURWHLQV LQ RWKHU RUJDQLVPV UHPDUNDEO\ IHZ F\DQREDFWHULDO SURWHLQV KDYH WR GDWH EHHQ XQHTXLYRFDOO\ GHPRQVWUDWHGWRELQGWRDQGWRUHTXLUHWKLVPHWDOLRQ $ONDOLQHSKRVSKDWDVHLVNQRZQWREHD]LQF UHTXLULQJ HQ]\PH LQ RWKHU SURNDU\RWHV 6LPSVRQ DQG 9DOOHH 7KH Synechococcus 3&& HQ]\PH LV ODUJHU WKDQ DQ\ RWKHU FKDUDFWHULVHG DONDOLQH SKRVSKDWDVH5D\HWDO 0RVWLPSRUWDQWO\WKH SURWHLQIURPSynechococcus 3&& LV LUUHYHUVLEO\
LQKLELWHG E\ ]LQF %ORFN DQG *URVVPDQ +RZHYHU GDWD DUH HFOHFWLF WKHUH KDYLQJ EHHQ QR H[WHQVLYH DQG V\VWHPDWLF VXUYH\ RI]LQFUHTXLUHPHQWV LQF\DQREDFWHULD ,W ZRXOG EH SUHPDWXUH WR FRQFOXGH WKDW WKH XVH RI ]LQF LV DQ\ OHVV FRPPRQ LQ F\DQREDFWHULDWKDQLQRWKHURUJDQLVPV =LQF FHUWDLQO\ LV QHHGHG IRU RSWLPDO JURZWK RI DW OHDVW VRPH VSHFLHV DQG LV UHTXLUHG IRU VRPH ELRFKHPLFDO SDWKZD\V )RU H[DPSOH ]LQF LV QHHGHG IRUWR[LQSURGXFWLRQ LQ Microcystis aeruginosa ZKLOH RWKHUPHWDOVWHVWHG LQFOXGLQJFRSSHU PDQJDQHVHDQG QLFNHO DUH QRWUHTXLUHG /XNDF DQG $HJHUWHU 2IFRXUVH 07 IURP Synechococcus 3&& ELQGV DQG GHWR[LILHV H[FHVV ]LQF EXW LW UHPDLQV WR EH HVWDEOLVKHGZKHWKHURUQRWLWDOVRKDVDUROHLQVWRULQJ RU RWKHUZLVH PRGLI\LQJ WKH DYDLODELOLW\ RIWKLV PHWDO LRQIRU]LQFUHTXLULQJSURWHLQV 6HTXHQFHV RI VHYHUDO F\DQREDFWHULDO SURWHLQV DUH NQRZQ ZKLFK FRQWDLQ GHGXFHG ]LQFELQGLQJ PRWLIV 2QH ZKLFKKDV EHHQ PHQWLRQHG LQ DSUHYLRXV VHFWLRQ LV '1$ SULPDVH ,Q DGGLWLRQ WR WKH SUHGLFWHG ]LQF ELQGLQJ VLWH FRPPRQ WR DOO SULPDVHV WKH Synechococcus 3&& SURWHLQ DOVR FRQWDLQV WKH
1 - 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
DGGLWLRQDO]LQFILQJHUDQGWKLVVWUXFWXUHLVUHTXLUHGIRU QRUPDO OHYHOV RI '1$ V\QWKHVLV LQ Synechococcus 3&& %LUG HW DO =LQFILQJHU PRWLIV KDYH DOVR EHHQ GHVFULEHG LQ RWKHU GHGXFHG F\DQREDFWHULDO SURWHLQV IRU H[DPSOH XQNQRZQ 25)V IURP Synechocystis 3&& 2JXUD HW DO EXWELQGLQJ RI]LQF WR WKHVH RWKHU VLWHVUHPDLQV WREH WHVWHG 51$ SRO\PHUDVH LV NQRZQ WR EH D ]LQF FRQWDLQLQJHQ]\PHDQGWKH VXEXQLWRIE. coli 51$ SRO\PHUDVH FRQWDLQV D ]LQFELQGLQJ SRFNHW ZKLFK LV DOVRSUHVHQWLQWKHUHODWHGVHTXHQFHRIWKHrpoC1 JHQH SURGXFW RI Anabaena 3&& %HUJVODQG DQG +DVHONRUQ ,QGHHG WKLV IRXU F\VWHLQH FRQWDLQLQJ PRWLI LV SUHVHUYHG WKURXJKRXW 5SR& VHTXHQFHV IURP HLJKW GLIIHUHQW F\DQREDFWHULD LQFOXGLQJ WKH Cyanophora F\DQHOOH Prochlorococcus VS DQG Prochlorothrix hollandica) 3DOHQLNDQG+DVHONRUQ VXJJHVWLQJWKDW]LQFLV OLNHO\WR EHDQHVVHQWLDOUHTXLUHPHQW IRUWUDQVFULSWLRQ WKURXJKRXW F\DQREDFWHULDO GLYHUVLW\ +RZHYHU WKH DVVRFLDWLRQ RI]LQF ZLWK WKLV GHGXFHG ELQGLQJ SRFNHW UHPDLQV WR EH WHVWHG 7KH GHGXFHG VHTXHQFH RI FDUERQLF DQK\GUDVH IURP Synechococcus 3&& HQFRGHG E\ WKH icfA JHQH FRQWDLQV ILYH DPLQR DFLGV &\V *OX +LV &\V DQG *OX ZKLFK DUH FDQGLGDWHV IRU ]LQFELQGLQJ DQG DUH FRQVHUYHG LQ DOO SURNDU\RWLF FDUERQLF DQK\GUDVHV )XNX]DZD HW DO +RZHYHU RQFH DJDLQ DVVRFLDWLRQ RI ]LQF ZLWK WKH F\DQREDFWHULDO FDUERQLF DQK\GUDVH KDV QRWIRUPDOO\ EHHQ GHPRQVWUDWHG 7KH GHGXFHG VHTXHQFH RI DPLQROHYXOLQLF DFLG GHK\GUDWDVH $/$' IURPSynechococcus 3&& FRQWDLQV D FRQVHUYHG ]LQFELQGLQJ GRPDLQ IRXQG LQ $/$' IURP PDPPDOV \HDVW DQG RWKHU HXEDFWHULD -RQHV HW DO 8QOLNH $/$' IURP KLJKHU SODQWV WKH F\DQREDFWHULDO SURWHLQ ZDV VKRZQ QRW WR UHTXLUH PDJQHVLXP RU PDQJDQHVH LRQV DQG LW ZDV SURSRVHG WR FRQWDLQ WLJKWO\ ERXQG ]LQF DQDORJRXV WR WKH PDPPDOLDQ \HDVW DQG RWKHU NQRZQ HXEDFWHULDO HQ]\PHV
=LQF ,PSRUW $Q$%& ,PSRUWHU ,QWKHSynechocystis 3&& JHQRPHWKHUHDUHWZR JHQHV VOU DQGVOU HQFRGLQJFRPSRQHQWV RI DGHGXFHG$73ELQGLQJFDVVHWWHW\SHLPSRUWHUZKLFK IURP WKH ORFDO JHQH DUFKLWHFWXUH DSSHDU OLNHO\ WR EH FRWUDQVFULEHG ZLWK D JHQH VOU HQFRGLQJ D GHGXFHGSHULSODVPLF]LQFELQGLQJSURWHLQ 7KHODWWHU KHUHLQ GHVLJQDWHG ,W]$ IRU LQZDUG WUDQVSRUW RI=LQF KDV VHTXHQFH VLPLODULW\ WR H[WUDSODVPDPHPEUDQH ]LQFELQGLQJ SURWHLQV 3]S IURP Haemophilus
influenzae /X HW DO $GF$ IURP Streptococcus pneumoniae 'LQWLOKDF HW DO DQG =QX$ IURP E. coli 3DW]HU DQG +DQWNH 7KHVH SURWHLQV DUH DOO SURSRVHG WR DFW LQ WKH SHULSODVPLF RUDWOHDVWWKHH[WUDSODVPDPHPEUDQHLQ WKH *UDPSRVLWLYH Streptococcus pneumonia) ELQGLQJ RI]LQF DQG VXEVHTXHQW GRQDWLRQ RIWKH PHWDO LRQV WR WKH$%&LPSRUWHU 3UHVXPDEO\ ]LQF LV H[SORLWHG E\ DOORIWKHF\DQREDFWHULDZKLFKFRQWDLQVXFK DQLPSRUW V\VWHP $ VXUYH\ RI WKH VSHFLHV DQG KDELWDW GLVWULEXWLRQ RIWKHVH JHQHV ZLWKLQ 7KH &\DQREDFWHULD ZRXOGEHRILQWHUHVW
Fig. 2. $ GLYHUJRQ ZKLFK HQFRGHV D GHGXFHG ]LQF VHQVRU QRZ GHVLJQDWHG=XU6HFWLRQ( ZKLFKUHJXODWHVDGHGXFHG$%&]LQF LPSRUW V\VWHP QRZ GHVLJQDWHG ,W] 7KH UHODWHG JHQH SURGXFWV IURPRWKHUEDFWHULDDUHVKRZQ
7KH&RSSHULURQ &RQQHFWLRQLQ 3KRWRV\QWKHWLF (OHFWURQ 7UDQVSRUW 7KHUH DUH UHODWLYHO\ IHZ NQRZQ XVHV IRU FRSSHU LQ F\DQREDFWHULD EXW RQH ZKLFK KDV EHHQ VWXGLHG LQ VRPH GHWDLO LV LQ SODVWRF\DQLQ 7ZR SURWHLQV FDQ IXQFWLRQ DV HOHFWURQ FDUULHUV EHWZHHQ SKRWRV\VWHP ,, DQGSKRWRV\VWHP, WKHLURQKDHPSURWHLQF\WRFKURPH c553 DQG FRSSHU FRQWDLQLQJ SODVWRF\DQLQ 6RPH F\DQREDFWHULD DQG LQGHHG VRPH HXNDU\RWLF DOJDH FRQWDLQ JHQHV IRU ERWK FDUULHUV ZKLOH RWKHUV FRQWDLQ RQO\ F\WRFKURPH c553 %RKQHU DQG %RJHU :RRG 0HUFKDQW DQG %RJRUDG 6DQGPDQQ ,Q Anabaena variabilis, ZKLFK FRQWDLQV ERWK JHQHV WKHLU H[SUHVVLRQ LV FRPSOHPHQWDU\ 3ODVWRF\DQLQ HQFRGHG E\ WKH petE JHQH H[SUHVVLRQ LQFUHDVHV ZLWK DGGHG FRSSHU DQG F\WRFKURPH c553 VKRZV D FRLQFLGHQW GHFUHDVH LQ H[SUHVVLRQ %RY\ HW DO 7KLV FKDQJH LQ H[SUHVVLRQLVUHJXODWHG DWWKH OHYHO RIWUDQVFULSWLRQ LQ
&KDSWHU 0HWDOV5HSHWLWLYH '1$ Anabaena variabilis. Synechocystis 3&& DOVR FRQWDLQV ERWK JHQHV EXW LQ WKLV F\DQREDFWHULXP H[SUHVVLRQRIpetE LQUHVSRQVHWRFRSSHULV FRQWUROOHG DW WKH WUDQVODWLRQDO RU SRVWWUDQVODWLRQDO OHYHO Synechococcus 3&& ODFNV SODVWRF\DQLQ DQG H[SUHVVLRQ RI F\WRFKURPH c 553 LV QRW UHJXODWHG E\ FRSSHU $QDELOLW\WRVZLWFKEHWZHHQWKHXVHRIWKHVH DOWHUQDWLYHUHGR[DFWLYHPHWDOVFRSSHUDQGLURQ LQDW OHDVW VRPH F\DQREDFWHULD SUHVXPDEO\ UHIOHFWV FULWLFDO IOXFWXDWLRQVLQDYDLODELOLW\GXULQJHYROXWLRQDWOHDVWLQ VRPHKDELWDWV 0RUDQG HW DO KDYH QRWHG WKDW WKHDEVHQFHRISODVWRF\DQLQLQVRPH VWUDLQVFRUUHODWHV ZLWK NQRZQ HQYLURQPHQWDO IDFWRUV IRU H[DPSOH Spirulina maxima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
( 3HUFHSWLRQ RI0HWDOVRIWKH &RSSHUDQG =LQF 7ULDGVLQ WKH(QYLURQPHQW 6PW% ,QUHVSRQVHWRHOHYDWHG FRQFHQWUDWLRQVRIDQXPEHURI WUDFHPHWDO LRQV LQFOXGLQJFDGPLXPDQG ]LQF WKHUH LV DQ LQFUHDVH LQ WKH DEXQGDQFH RI smtA WUDQVFULSWV +HDW VKRFN GRHV QRW HOLFLW DQ HTXLYDOHQW UHVSRQVH +XFNOHHWDO ,QKLELWRU VWXGLHV KDYH VKRZQ WKDWWKHUH LV QRHIIHFW RIFDGPLXPRQ smtA WUDQVFULSW VWDELOLW\ LQGLFDWLQJ FRQWURO DW WKH OHYHO RI WUDQVFULSWLRQDO LQGXFWLRQ ([SUHVVLRQ RI D UHSRUWHU JHQH lacZ GULYHQ E\ WKH smtA RSHUDWRUSURPRWHU ZDV VLPLODUO\ VKRZQ WR LQFUHDVH LQ UHVSRQVH WR D QXPEHU RIWUDFH PHWDO LRQV +RZHYHU DW PD[LPXP SHUPLVVLYH FRQFHQWUDWLRQV RQO\ ]LQF DQG WR D OHVVHU H[WHQW FDGPLXP DQG FRSSHU LRQV JDYH VXEVWDQWLDO LQFUHDVHV LQ UHSRUWHU JHQH H[SUHVVLRQ +XFNOH HW DO
,PPHGLDWHO\XSVWUHDPES IURPWKHsmtA JHQH LV D GLYHUJHQWO\ WUDQVFULEHG JHQH HQFRGLQJ D SURWHLQ GHVLJQDWHG 6PW% +XFNOH HW DO 7KH 6PW% SRO\SHSWLGH FRQWDLQV D GHGXFHG KHOL[WXUQKHOL[ ,W DOVR VKRZV VHTXHQFH '1$ELQGLQJ PRWLI VLPLODULW\WR QXPHURXV EDFWHULDO SURWHLQV D QXPEHU RI ZKLFK DUH NQRZQ WUDQVFULSWLRQDO UHSUHVVRUV LQYROYHG LQ WKH SHUFHSWLRQ RI PHWDOV +XFNOH HW DO 7XUQHUHWDO 7KHVH LQFOXGH SODVPLG HQFRGHG $UV5 IURP E. coli 6DQ )UDQFLVFR HW DO Staphylococcus xylosus 5RVHQVWHLQ HW DO Staphylococcus aureus -L DQG 6LOYHU DQG FKURPRVRPDOO\HQFRGHG$UV5IURPE. coli ;XHWDO DOO RI ZKLFK DUH PHWDOR[\DQLRQ UHVSRQVLYH UHSUHVVRUV RIars DUVHQLF UHVLVWDQFH RSHURQV 2WKHU KRPRORJXHV DUH&DG&SURWHLQV IURPS. aureus (QGR DQG 6LOYHU DQG Listeria monocytogenes /HEUXQHWDO ZKLFK DUH FDGPLXPUHVSRQVLYH UHSUHVVRUV RI JHQHV HQFRGLQJ FDGPLXP HIIOX[ $73DVHV DQG=QW5IURP6 aureus ZKLFK DSSHDUV WR UHJXODWH H[SUHVVLRQ RI D FREDOW]LQF H[SRUWHU ;LRQJ DQG -D\DVZDO )LQDOO\ 0HU5 IURP Streptomyces lividans ZKLFK LV WKRXJKW WR EH D PHUFXU\ UHVSRQVLYH UHSUHVVRU RI JHQHV HQFRGLQJ PHUFXU\ GHWR[LILFDWLRQ V\VWHPV LQFOXGLQJ PHUFXU\ UHGXFWDVH KRZHYHULWLVXQOLNHWKHPRUHZHOONQRZQ 0HU5 WUDQVFULSWLRQDO VZLWFKHV ZKLFK KDYH EHHQ ZHOO FKDUDFWHULVHGLQRWKHUEDFWHULD ,QPXWDQWVRISynechococcus 3&&GHILFLHQWLQ WKH smt GLYHUJRQ smtA DQG smtB WKHUH LV KLJKO\ HOHYDWHG EDVDO H[SUHVVLRQ RI lacZ) DVVRFLDWHG ZLWK WKH smt RSHUDWRUSURPRWHU +XFNOH HW DO 5HSUHVVLRQ DQG PHWDOGHSHQGHQW H[SUHVVLRQ ZDV UHVWRUHG LQ FHOOV FRQWDLQLQJ SODVPLG ERUQH DQGRU FKURPRVRPDO smtB +XFNOH HW DO LPSO\LQJ WKDW 6PW% LV D PHWDO UHVSRQVLYH UHSUHVVRU RI smtA H[SUHVVLRQ (OHFWURSKRUHWLF PRELOLW\ VKLIW DVVD\V GHWHFWHG PXOWLSOH FRPSOH[HV IRUPLQJ ZLWK WKH smtA RSHUDWRUSURPRWHU DW OHDVW RQH RI ZKLFK ZDV DWWULEXWHG WR 6PW% VLQFH WKLV FRPSOH[ZDV DEVHQWLQ FHOOV GHYRLG RI smtB. )XUWKHUPRUH WKLV FRPSOH[ GLVVRFLDWHGIURP'1$LQUHVSRQVHWR]LQF DQGERXQG DJDLQ IROORZLQJ WUHDWPHQW RI H[WUDFWV ZLWK FKHODWLQJ DJHQWV WR UHPRYH ]LQF 0RUE\ HW DO 7KHVH REVHUYDWLRQV KDYHEHHQ FRQILUPHG XVLQJ UHFRPELQDQW 6PW% (UEH HW DO ,Q DGGLWLRQ VLWH GLUHFWHG PXWDWLRQV ZLWKLQ smtB KDYHHVWDEOLVKHG WKDW DSDLU RI KLVWLGLQH UHVLGXHV + DQG + DUH HVVHQWLDO IRU WKH SHUFHSWLRQ RI ]LQF E\ 6PW% ZKLOH F\VWHLQH UHVLGXHV DUH QRW 7XUQHU HW DO 7KLV LV RI LQWHUHVW VLQFH F\VWHLQH UHVLGXHV KDYH EHHQ SURSRVHG DQG LQ WZR FDVHV GHPRQVWUDWHG WR EH LPSRUWDQW IRU
1- 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
WKHSHUFHSWLRQRIPHWDOLRQVE\RWKHUPHPEHUVRIWKLV IDPLO\RIUHSUHVVRUV 6KLHWDO 7KHOZHOOHWDO 7KLV LQGLFDWHV WKH LQWULJXLQJ SRVVLELOLW\ RI D IDPLO\ RI PHWDOVHQVRUV SRVVHVVLQJ GLVWLQFW PHWDO ELQGLQJ VLWHV ZKLFK JLYH GLYHUVLW\ LQ WKH VSHFWUD RI PHWDOVVHQVHG D SUHIHUHQFH IRU FDGPLXP E\ &DG& DUVHQLFE\$UV5]LQFE\6PW%DQGPHUFXU\E\0HU5 7KH SRVVLELOLW\ UHPDLQV WKDW RWKHU PHPEHUV RI WKLV IDPLO\ RI PHWDO VHQVRUV ZLWK GLIIHUHQW PHWDO VSHFLILFLWLHV PD\ DOVR EH SUHVHQW LQ DW OHDVW VRPH F\DQREDFWHULDO JHQRPHV VHH IROORZLQJ VHFWLRQV $ SDUWLDO VWUXFWXUH RI6PW% LV QRZNQRZQ &RRNHWDO DQGWKLVKDVDOORZHGWKH]LQFVHQVLQJVLWHVWREH PRGHOOHG ,WZDVVXJJHVWHGIURPWKHVHVWUXFWXUDOGDWD WKDWDSR6PW% PD\ DVVRFLDWHZLWK'1$WKDWLV EHQW 0HWDOORUHJXODWLRQ E\ 6PW% FRXOG LQYROYH PHWDO GHSHQGHQW ORVV RI '1$ EHQGLQJ SHUKDSV DW VRPH LQWHUPHGLDWH OHYHO RI]LQF LQ DGGLWLRQ WR GLVVRFLDWLRQ DWKLJKO\HOHYDWHGFRQFHQWUDWLRQVRILQGXFHU
=LD5DQGRWKHU+RPRORJXHVRI6PW%$ )DPLO\RI0HWDO6HQVRUV 8SVWUHDP RI mtnA IURP Synechococcus vulcanus 6KLPL]X HW DO LV D SDUWO\ VHTXHQFHG GLYHUJHQWO\WUDQVFULEHG25)ZKLFKHQFRGHVDSURWHLQ ZLWK VLPLODULW\ WR 6PW% GHVLJQDWHG 0WQ% 7XUQHU DQG 5RELQVRQ 7KLV LV SUHVXPHG WR EH DQ DQDORJRXV]LQFVHQVLQJUHSUHVVRU ,QDGGLWLRQZLWKLQ WKH Synechocystis 3&& JHQRPH DUH WZR IXUWKHU 25)V HQFRGLQJ GHGXFHG SURWHLQV ZLWK VLPLODULW\ WR 6PW% 2QH RIWKHVH ziaR, KDV DOUHDG\ EHHQ QRWHG 5HSRUWHU JHQH FRQWUXFWV VKRZ WKDW =LD5 DFWV DV D UHSUHVVRU UHTXLUHG IRU ]LQFGHSHQGHQW H[SUHVVLRQ IURP WKH ziaA RSHUDWRUSURPRWHU 7KHOZHOO HW DO *HO UHWDUGDWLRQ DVVD\V GHWHFWHG =LD5 ELQGLQJ WRWKHziaA RSHUDWRUSURPRWHUDQG=LD5'1$ELQGLQJ ZDV HQKDQFHG E\ WUHDWPHQW ZLWK WKH PHWDO FKHODWRU SKHQDQWKUROLQH 7KH DQDO\VLV RI VLWH GLUHFWHG PXWDQWV RI =LD5 HVWDEOLVKHG WKDW D SDLU RI F\VWHLQH UHVLGXHVDGMDFHQWWRWKHKHOL[WXUQKHOL['1$ELQGLQJ VLWH DUH HVVHQWLDO IRU ]LQF SHUFHSWLRQ DV LV DKLVWLGLQH UHVLGXH 7KHOZHOO HW DO ZKLFK DOLJQV ZLWK D KLVWLGLQH UHVLGXH + LQ 6PW% ZKLFK IRUPV D VHFRQG SDLU RI VLWHV ORFDWHG DW WKH 6PW% GLPHU LQWHUIDFH&RRNHWDO $VHFRQG25)ZLWKLQWKHSynechocystis 3&& JHQRPH ZLWK VLPLODULW\ WR smtB KDV EHHQ GHVLJQDWHG arsA DQGLVPRVWVLPLODUWRWKHR[\DQLRQ VHQVRUJHQH arsR, DOWKRXJK DJDLQ LW UHPDLQV WR EH HVWDEOLVKHG ZKLFK LI DQ\ PHWDOV DUH VHQVHG 7KH ORFDWLRQ RI arsA SURYLGHV IHZFOXHV DERXWWKHOLNHO\DFWLRQRILWV
SURGXFW ,W LV QHDU WR D JHQH HQFRGLQJ SXWDWLYH DTXDSRULQ 7KH LGHQWLILFDWLRQ RI D IDPLO\ RIUHODWHG PHWDO VHQVRUV ZKLFK GLIIHU LQ WKH VSHFWUD RI PHWDOV WKH\ SHUFHLYH PD\ EH H[SORLWHG WR LGHQWLI\ WKH GHWHUPLQDQWVRIPHWDOVSHFLILFLW\
$ 3XWDWLYH /RZ=LQF 6HQVRU =XU 'LYHUJHQWO\ WUDQVFULEHG IURP WKH itz JHQHV VHH VHFWLRQ ' LV D JHQH VOO HQFRGLQJ D )XUOLNH SURWHLQ)LJ $QRWKHUJHQHLQ Synechocystis 3&& HQFRGHV D )XUOLNH SURWHLQ ZKLFK LV WKH PRUH VLPLODU LQ VHTXHQFH WR )XU IURP Synechococcus 3&& WKH ODWWHU KDYLQJ EHHQ GHPRQVWUDWHG WR DFW LQ UHVSRQVH WR HOHYDWHG FRQFHQWUDWLRQV RILURQ WR UHSUHVV H[SUHVVLRQ RI JHQHV RQO\ UHTXLUHG XQGHU ORZ LURQ FRQGLWLRQV *KDVVHPLDQ DQG 6WUDXV 7KH IXQFWLRQ RI WKH SURGXFW RI VOO LV WKHUHIRUH XQNQRZQ +RZHYHU LQ E. coli LW KDV UHFHQWO\ EHHQ GHPRQVWUDWHGWKDWD)XUKRPRORJXHUHGHVLJQDWHG=XU LVLQIDFWD]LQFVHQVRUZKLFKUHSUHVVHV JHQHVWKDWDUH RQO\ UHTXLUHG XQGHU ORZ ]LQF FRQGLWLRQV 3DW]HU DQG +DQWNH :HWKHUHIRUHVXJJHVWWKDWVOOLVD =XUOLNH VHQVRU ZKLFK UHJXODWHV WKH GLYHUJHQWO\ WUDQVFULEHG GHGXFHG ]LQF LPSRUW V\VWHP :H ORRN IRUZDUGWRWKHFKDUDFWHULVDWLRQRIWKHIXOOFRPSOHPHQW RIVHQVRUVWKDWUHVSRQGWRIOXFWXDWLRQVLQ]LQF DQGWKH IXOO FRPSOHPHQW RI KRPRHRVWDWLF V\VWHPV WKDW WKH\ UHJXODWHLQSynechocystis 3&&)LJ
)XUWKHU6HTXHQFHVIURP WKH 6\QHFKRF\VWLV 3&& *HQRPH 6HYHUDO SXWDWLYH PHWDO VHQVRUV KDYH DOUHDG\ EHHQ GHVFULEHG LQ WKH FRQWH[W RI GLVFXVVLRQV RI WKH DVVRFLDWHG VWUXFWXUDO JHQHV DQG GHVFULSWLRQV RI6PW% =LD5 DQG =XU LQWKHSUHFHGLQJ VHFWLRQV ,QDGGLWLRQ WKHUHDUHWZR 25)V HQFRGLQJSURWHLQV ZLWK VLPLODULW\ WR WKH PHUFXU\ LRQ VHQVRU 0HU5 QRW 6PW%OLNH 2QHRIWKHVHLVORFDWHGZLWKWKHUHJLRQ VKRZQLQ)LJ DQG SDUWO\ DOLJQV ZLWK 0HU5 DQG SDUWO\ ZLWK SUHFRUULQ LVRPHUDVH HQFRGHG E\ cobH). 3UHFRUULQ LVRPHUDVH LV LQYROYHG LQ WKH ELRV\QWKHVLV RI YLWDPLQ % WKH VROH FREDOW FRQWDLQLQJ FRIDFWRU 7KH UHFUXLWPHQW RI '1$ IUDJPHQWV HQFRGLQJ GRPDLQV RI VXFK DQ HQ]\PH VXJJHVWV WKDW WKLV UHJXODWRU PD\ EH LQYROYHG LQ WKH SHUFHSWLRQ RI FREDOW DQG LQGHHG RXU GDWD VXSSRUW WKLV K\SRWKHVLV DQG WKH VHQVRUKDV EHHQ SURYLVLRQDOO\ GHVLJQDWHG &RD5 5XWKHUIRUG HW DO XQSXEOLVKHG 7KHUH DUH XQFKDUDFWHULVHG WZR FRPSRQHQW VHQVRUV LQ WKH FRPSOHWH Synechocystis 3&&JHQRPHDQGLWLVRIFRXUVHSRVVLEOHWKDW
&KDSWHU 0HWDOV5HSHWLWLYH'1$
LOW Zn2+
High Zn2+
Fig 3. $PRGHOVKRZLQJWZR SURSRVHG µKLJK DQGORZ¶ ]LQF VHQVRUV LQ Synechocystis 3&& DQG WKH SURSRVHG PRGHV RIDFWLRQ RIWKH SURGXFWVRIWKHJHQHVWKH\UHJXODWH
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µFURVVWDON¶ EHWZHHQ WKH KRPRHRVWDWLF V\VWHPV IRUGLIIHUHQWPHWDOV
) &RQWUDVWLQJ5HVSRQVHV RI7ZR &\DQREDFWHULD WR HOHYDWHG=LQF ,I WKH GHWR[LILFDWLRQ DQG VXEVHTXHQWO\ WKH PRUH ZLGHVSUHDG XVH RI ]LQF ILUVW HPHUJHG GXULQJ WKH DGDSWLYH UDGLDWLRQ RI WKH F\DQREDFWHULD WKHQ D
GLYHUVLW\ RI UHVSRQVHV WR HOHYDWHG ]LQF PLJKW EH DQWLFLSDWHG ,QGHHG Synechocystis 3&& DQG Synechococcus 3&& SRVVHVV FORVHO\ UHODWHG ]LQF VHQVRUV =LD5 DQG 6PW% EXWWKH\ UHJXODWH YHU\ GLIIHUHQW VWUXFWXUDO SURWHLQV ZLWK YHU\ GLIIHUHQW FRQVHTXHQFHVIRUWKHFHOOELRORJ\RI]LQF 7KHIRUPHU WULJJHUV H[SXOVLRQ RI]LQF YLD =LD$ PHGLDWHG HIIOX[ ZKLOH WKH ODWWHU WULJJHUV LQWHUQDO VHTXHVWUDWLRQ E\ 6PW$ +XFNOH HW DO 7KHOZHOO HW DO )LJ ,V ]LQF KRPRHRVWDVLV LQ Synechocystis 3&& SXUHO\ UHJXODWHG E\ PHWDO LRQ LQIOX[HIIOX[ ZKLOH LQ Synechococcus 3&& LW LV UHJXODWHG E\ VHTXHVWUDWLRQLQWKHF\WRSODVP" ,VWKHSHULSODVPXVHG DV DQ DOWHUQDWLYH ]LQF VWRUH LQ Synechocystis 3&& RUGRHVWKLV RUJDQLVPKDYH DOHVVHUUHTXLUHPHQW IRU ]LQF VWRUDJH" 'RHV WKLV UHIOHFW PRUH ZLGHVSUHDG XVHRI]LQFLQSynechococcus 3&&DQGPHUHO\D UHTXLUHPHQWIRUWKHGHWR[LILFDWLRQRIXQQHFHVVDU\]LQF LQ Synechocystis 3&&RUGLGWKHIRUPHUHYROYH
1 - 5RELQVRQ -& 5XWKHUIRUG 05 3RFRFN DQG -6 &DYHW
Synechococcus 3&&
Synechocystis 3&&
Fig. 4. 7KHVHWZRF\DQREDFWHULDKDYHFORVHO\UHODWHGVHQVRUVRIHOHYDWHGF\WRSODVPLF]LQFEXWWKH\UHJXODWHYHU\GLIIHUHQWVWUXFWXUDOSURWHLQV ZLWKGLIIHUHQWFRQVHTXHQFHVIRUWKHFHOOELRORJ\RI]LQF
LQ PRUH ]LQF GHILFLHQW HQYLURQPHQWV ZKLFK KDYH VHOHFWHG IRU PRUH HIILFLHQW VHTXHVWUDWLRQ DQG VWRUDJH RIHQGRJHQRXV]LQF"
,, 5HSHWLWLYH'1$LQ&\DQREDFWHULD $ 7KH.QRZQ5HSHDWV $ QXPEHU RI UHSHDWHG '1$ VHTXHQFHV KDYH EHHQ GHVFULEHG LQ WKH JHQRPHV RIGLIIHUHQW F\DQREDFWHULD 7KUHH W\SHV RI VKRUW WDQGHPO\ UHSHDWHG UHSHWLWLYH 6755 VHTXHQFHV ZHUH ILUVW REVHUYHG LQ Calothrix 3&& $ UHSHDWHG KHSWDQXFOHRWLGH IRUPV WKH FRUH 6755 VHTXHQFH DQG WKLV LV NQRZQ WR DOVR EH SUHVHQW LQ Nostoc 3&& DQG Anabaena 3&& 0D]HO HW DO 6755 HOHPHQWV DUH HVWLPDWHG WR RFFXU DSSUR[LPDWHO\ WLPHV ZLWKLQ WKH 0E JHQRPH RI Calothrix 3&& 6755 HOHPHQWV DUH HYHQ PRUH DEXQGDQW LQ Anabaena azollae WKDQ Anabaena 3&& EXW DQ H[DPLQDWLRQ RI VLWH ORFDWLRQ VXJJHVWHG WKDW WKH
HOHPHQWV KDYH D GLIIHUHQW GLVWULEXWLRQ HYHQ LQ WKHVH FORVHO\UHODWHGVWUDLQV-DFNPDQDQG0XOOLJDQ 6L[ QHZ 6755 VHTXHQFHV ZHUH DOVR LGHQWLILHG LQ Anabaena azollae. ,Q DGGLWLRQ WR 6755 VHTXHQFHV Anabaena 3&& DQG VHYHUDO RWKHU F\DQREDFWHULD DOVR FRQWDLQ ORQJHU WDQGHPO\ UHSHDWHG UHSHWLWLYH /755 VHTXHQFHV ZKLFK DUH ES LQ OHQJWK IRXQG LQ FOXVWHUV DQG DOVR FRQVHUYHG LQ VHTXHQFH LQ VHYHUDO VWUDLQV 0DVHSRKO HW DO 7ZR UHSHDWV GHVLJQDWHG 5(3$ DQG 5(3% KDYH EHHQGHVFULEHG LQMicrocystis VS ZKLFK DSSHDUWREH DEVHQW IURP Synechococcus 3&& DQG Synechocystis 3&&$VD\DPDHWDO 7KH RFWDPHULF VHTXHQFH *&*$7&*& GHVLJQDWHG+,3 IRUKLJKO\ LWHUDWHGSDOLQGURPH ZDV ILUVW LGHQWLILHG DW WKH MXQFWLRQV RI D JHQH WKH smtB UHSUHVVRU GHOHWLRQHYHQWLQSynechococcus 3&& *XSWD HW DO 7KH VHTXHQFH ZDV QRWHG WR RFFXU PDQ\ WLPHV ZLWKLQ WKH smt UHJLRQ DQG VXEVHTXHQWDQDO\VHV RIGDWDEDVHHQWULHV LQGLFDWHGWKDW +,3 ZDV RYHUUHSUHVHQWHG LQ PDQ\ F\DQREDFWHULDO
&KDSWHU 0HWDOV5HSHWLWLYH'1$
Fig. 5. 6LQJXODU RYHUUHSUHVHQWDWLRQ RI+,3 LQ FXUUHQWO\NQRZQ VHTXHQFH QXFOHRWLGHV IURP Synechococcus 3&& XSSHU SDQHO DQGLQWKHFRPSOHWHJHQRPHRISynechocystis 3&& ERWWRPSDQHO 7KHILUVWDQGVHFRQGQXFOHRWLGHSDLUV DUHOLVWHGRQWKHD[HV IURPOHIWWRULJKWDQG IURPIURQWWREDFN LQWKHRUGHUVKRZQ RQ WKHNH\ 7KHSDWWHUQV RQWKHNH\FRUUHVSRQGWRWKHILUVWQXFOHRWLGHSDLU
1 - 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
Fig. 6. $QDO\VLVRIQXFOHRWLGHVIODQNLQJRFWDPHULF+,3O VLWHVLQSynechocystis 3&&UHYHDOVWKDWWKHPDMRULW\RIVLWHVDUHIODQNHGE\* DQG & WKXV LGHQWLI\LQJ D QRYHO RYHUUHSUHVHQWHG GHFDPHULF VHTXHQFH LQ WKLV RUJDQLVP GHVLJQDWHG +,3Q 7KH IUHTXHQF\ RI HDFK QXFOHRWLGH QHJDWLYHQXFOHRWLGHSRVLWLRQV DQG SRVLWLYHQXFOHRWLGHSRVLWLRQV DUHUHSUHVHQWHGLQWKHRUGHU$*&7UHVSHFWLYHO\
JHQRPHV5RELQVRQHWDO $Q DQDO\VLV RIWKH IUHTXHQF\RIRFFXUUHQFHRIDOOSRVVLEOHRFWDPHULF SDOLQGURPHV UHYHDOHG WKDW WKHUH ZDV VLQJXODU RYHU UHSUHVHQWDWLRQ RI +,3O LQ VHYHUDO VWUDLQV )LJ 7KLV VHTXHQFH FRQVWLWXWHG RI QRYHO VHTXHQFHV REWDLQHG IURP FORQHV VHOHFWHG DW UDQGRP IURP D OLEUDU\ SUHSDUHG IURP Synechococcus 3&& 7KLVLVHTXLYDOHQWWRDVLWHRFFXUULQJRQDYHUDJHRQFH HYHU\ QXFOHRWLGHV ZLWKLQ WKLV RUJDQLVP +RZHYHU DQDO\VLV RIHTXLYDOHQW UDQGRP VHTXHQFHV IURP RQH VWUDLQ Calothrix ' LQGLFDWHG WKDW+,3O LV QRW DEXQGDQW LQ DOO F\DQREDFWHULDO JHQRPHV 5RELQVRQHWDO $Q ROLJRQXFOHRWLGH LQFOXGLQJ +,3O JDYH PXOWLSOH SURGXFWV ZLWK WHPSODWH '1$ IURP F\DQREDFWHULD NQRZQWRFRQWDLQDEXQGDQW+,3O EXWQRWIURPVWUDLQV LQ ZKLFK +,3O LV UDUH RU DEVHQW 5RELQVRQ HW DO )XUWKHUPRUH FKDUDFWHULVWLF +,33&5 EDQGLQJ SDWWHUQV ZHUH REWDLQHG IURP VRPH VWUDLQV VXJJHVWLQJ SRVVLEOH DSSOLFDWLRQV LQ VWUDLQ LGHQWLILFDWLRQ DQGRU WKH GLDJQRVWLF WHVWLQJ RI
HQYLURQPHQWDO VDPSOHV IRU WKH SUHVHQFH RI F\DQREDFWHULD ([DPLQDWLRQ RI WKH VHTXHQFHV IODQNLQJ DOO RI WKH +,3O VLWHV LQ WKH FRPSOHWH JHQRPH RISynechocystis 3&& UHYHDOV D UHPDUNDEOH FRQVHQVXV )LJ ZLWK DQ H[WHQGHG +,3O VLWH **&*$7&*&& EHLQJ SUHVHQW LQ WKLV RUJDQLVP 7KLV GHFDPHULF YHUVLRQ RI +,3O KHUH GHVLJQDWHG +,3Q LV GLVWULEXWHGWKURXJKRXWWKHJHQRPH)LJ DQG LV QRW FOXVWHUHG LQWR RQH UHJLRQ IRU H[DPSOH SUR[LPDO RU GLVWDO WR WKH RULJLQ RIUHSOLFDWLRQ +,3Q LV IRXQG ERWK RXWVLGH DQG ZLWKLQ SURWHLQ FRGLQJ UHJLRQV DQG RFFXUVLQDOOWKUHHUHDGLQJIUDPHV +RZHYHU WKH H[WHQGHG +,3O GHFDPHU GRHV QRW DFFRXQW IRU WKH PDMRULW\ RI WKH +,3O VLWHV LQ Synechococcus 3&& ZKHUH RQO\ WKH RFWDPHU DSSHDUV WR EH RYHUUHSUHVHQWHG ,QNQRZQVHTXHQFHV IURP Anabaena 3&& D GLIIHUHQW GHFDPHULF YDULDQWRI+,3 $*&*$7&*&7 DSSHDUV WR EH RYHUUHSUHVHQWHG XQSXEOLVKHGREVHUYDWLRQV
&KDSWHU 0HWDOV5HSHWLWLYH'1$
QW
QW Fig. 7. 7KH GHFDPHULF YDULDQW RI+,3O +,3Q LV GLVWULEXWHG WKURXJKRXW WKH FRPSOHWH JHQRPH RISynechocystis 3&& DQG RFFXUV ERWKLQVLGHDQGRXWVLGHRSHQUHDGLQJIUDPHV 7KHXSSHUOLQHVUHSUHVHQWWKHORFDWLRQVRIWKHGHFDPHUVDQGWKHORZHUEDUVVKRZWKHSRVLWLRQV RIRSHQUHDGLQJIUDPHV /HVVWKDQWHQFRSLHVLQWRWDORIDQ\GHFDPHULF VHTXHQFHZRXOGEHH[SHFWHGWRRFFXUE\UDQGRPFKDQFHZLWKLQWKLV HQWLUHJHQRPH
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coli), UHYHDO WKDW+,3 GRHV QRWLQWURGXFH JDSV LQWR VXFK DOLJQPHQWV +,3 DOLJQV ZLWK SDUWLDO +,3 VLWHV LQ KRPRORJRXV VHTXHQFHV IURPQRQF\DQREDFWHULD FRQVLVWHQW ZLWK LWV SURSDJDWLRQ YLD QXFOHRWLGH VXEVWLWXWLRQV UDWKHU WKDQ LQVHUWLRQ5RELQVRQHWDO ,QPRVWFDVHV RQO\
1 - 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
3RVLWLRQQW Fig. 8. 7KH QXPEHU RI WLPHV WKDW HDFK QHDU+,3Q VLWH RFFXUV ZLWKLQ WKH FRPSOHWH JHQRPH RI Synechocystis 3&& 7KH QHDU +,3Q VLWHVDUHGHILQHGDVWKRVHLQZKLFKRXWRIQXFOHRWLGHVPDWFKWKHFRQVHQVXVVHTXHQFH ,WLVPRVWFRPPRQIRUWKHVHVLWHVWRKDYH DQXFOHRWLGH LQFRUUHFWZLWKLQWKHRXWHUIRXUQXFOHRWLGHV 7KH GDVKHG OLQH VKRZV WKH SUHGLFWHGQXPEHURI WLPHV DQ\ JLYHQ GHFDPHU VKRXOG RFFXUE\FKDQFH
D VPDOO QXPEHU RIQXFOHRWLGH VXEVWLWXWLRQV ZRXOG EH UHTXLUHG WR FUHDWH +,3O VLWHV DW HTXLYDOHQW ORFDWLRQV LQ WKH KRPRORJRXV QRQF\DQREDFWHULDO VHTXHQFHV 0DQ\ RI WKH QXFOHRWLGH GLIIHUHQFHV DUH HLWKHU V\QRQ\PRXV RU UHVXOW LQ RQO\ FRQVHUYDWLYH DPLQR DFLGFKDQJHV 7KHVH REVHUYDWLRQV VXJJHVW WKDW HLWKHU D FRPPRQ DQFHVWRU FRQWDLQHG SDUWLDO +,3O VLWHV ZKLFK KDYH EHHQ ORVW LQ RWKHU EDFWHULD VXFK DV E. coli RU WKDW D FRPPRQ DQFHVWRU FRQWDLQHG SDUWLDO +,3O VLWHV ZKLFK VXEVHTXHQWO\ DFFXPXODWHG QXFOHRWLGHVXEVWLWXWLRQV ZLWKLQVRPHF\DQREDFWHULDWR JHQHUDWH SHUIHFW +,3O VLWHV %RWK H[SODQDWLRQV DUH FRQVLVWHQW ZLWK WKH SUHVHQFH RI VHOHFWLRQ ZLWKLQ F\DQREDFWHULD WR UHWDLQ +,3O VLWHV WKDW GR QRW RWKHUZLVHLPSDLUJHQHIXQFWLRQ )LJ UHYHDOV WKH QXPEHU RI HDFK GLIIHUHQW QHDU +,3Q VLWH LQ ZKLFK RXWRI QXFOHRWLGHV PDWFK WKH FRQVHQVXV VHTXHQFH ZLWKLQ Synechocystis 3&& 2I FRXUVH LW PLJKW EH DQWLFLSDWHG WKDW DQ\ VHTXHQFH FORVHO\ UHODWHG WR +,3Q FRXOG EH RYHU
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
&KDSWHU 0HWDOV5HSHWLWLYH'1$ GDWH LGHQWLILHG DQ\ FRPSOH[HV ZKLFK IRUP VSHFLILFDOO\ DQG H[FOXVLYHO\ ZLWK +,3O 5RELQVRQ HW DO 6LQFH +,3O ZDV ILUVW LGHQWLILHG DW WKH ERUGHUV RI D JHQH GHOHWLRQ LW KDV EHHQ K\SRWKHVLVHG WKDW LW FRXOG SURPRWHZLGHVSUHDGJHQHUHDUUDQJHPHQWJHQHUDWLQJD SRRO RI JHQHWLF GLYHUVLW\ DQG WKHUHE\ HQKDQFLQJ DGDSWDWLRQ WR HQYLURQPHQWDO FKDQJH +RZHYHU IUHTXHQFLHV RI GHOHWLRQ EHWZHHQ D GHILQHG SDLU RI +,3O VLWHVZHUHVKRZQWREH VLPLODULQE. coli DQGLQ Synechococcus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einococcus radiodurans )XMLWDQL HW DO ,Q FRPPRQ ZLWK PDQ\ F\DQREDFWHULD 0DQQ DQG &DUU D. radiodurans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ulothrix desertica ZDVLVRODWHGIURPILQHGHVHUWVDQG LQ &KLOH 5LSSND HW DO ZKLOH WKH FORVHO\ UHODWHG Culothrix 'ZDVLVRODWHGIURPDPDQJURYH LQ&XED ,WZLOOEHRILQWHUHVWWRHVWDEOLVKZKHWKHUWKH IRUPHU XQOLNH WKH ODWWHU KDV PDVVLYH RYHU UHSUHVHQWDWLRQ RI +,3O ZLWKLQ LWV QXFOHLF DFLGV $Q H[WHQVLYH DQDO\VLV RI WKH +,3O FRQWHQW RI PXOWLSOH VWUDLQV RI F\DQREDFWHULD IURP KDELWDWV ZKLFK DUH SURQHWRGHVLFFDWLRQ RULQ ZKLFKWKLV LV XQOLNHO\ IRU H[DPSOH PDULQH KDELWDWV FRXOG SURYH WR EH LQIRUPDWLYH )XUWKHUPRUHDGHWDLOHGLQVSHFWLRQRIWKH JHQHV LQ Synechocystis 3&& ZKLFK GR RU GR QRW FRQWDLQ +,3Q PD\ DLG WKH LGHQWLILFDWLRQ RI IDFWRUVZKLFKKDYHHQFRXUDJHGLWVSUROLIHUDWLRQ
,,, &RQFOXGLQJ 5HPDUNV :LWK WKH GHYHORSPHQW RI 3&5EDVHG PHWKRGV IRU UDSLGO\ DQDO\VLQJ '1$ VHTXHQFHV IURP RUJDQLVPV LQ GLIIHUHQW HQYLURQPHQWV WKH WHFKQRORJ\ RI PROHFXODU JHQHWLFV LV LQFUHDVLQJO\ EHLQJ XVHG WR DGGUHVV HFRORJLFDOHQYLURQPHQWDODQGHYROXWLRQDU\TXHVWLRQV
1 - 5RELQVRQ-& 5XWKHUIRUG 05 3RFRFNDQG-6&DYHW
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
$FNQRZOHGJHPHQWV 5HVHDUFK RQ PHWDOJHQHLQWHUDFWLRQV DQG +,3 LV VXSSRUWHG E\ WKH %LRWHFKQRORJ\ DQG %LRORJLFDO 6FLHQFHV 5HVHDUFK &RXQFLO DQG WKH 1DWXUDO (QYLURQPHQWDO5HVHDUFK&RXQFLO
5HIHUHQFHV $VD\DPD 0 .DEDVDZD 0 7DNDKDVKL , $LGD 7 DQG 6KLUDL 0 +LJKO\ UHSHWLWLYH VHTXHQFHV DQG FKDUDFWHULVWLFV RI JHQRPLF '1$ LQ XQLFHOOXODU F\DQREDFWHULDO VWUDLQV )(06 0LFURELRO/HWW %HUJ -0 DQG 6KL < 7KH JDOYDQL]DWLRQ RI ELRORJ\ $ JURZLQJDSSUHFLDWLRQIRUWKHUROHV RI]LQF 6FLHQFH %HUJVODQG .- DQG +DVHONRUQ 5 (YROXWLRQDU\ UHODWLRQVKLSV DPRQJ HXEDFWHULD F\DQREDFWHULD DQG FKORURSODVWV HYLGHQFH IURP WKH rpoCL JHQH RI Anabaena VS VWUDLQ 3&& - %DFWHULRO %HUNHOPDQ7 *DUUHW(QJHOH3DQG +RIIPDQ 1( 7KHpacL JHQH RI Synechococcus VS VWUDLQ 3&& HQFRGHV D &D WUDQVSRUWLQJ$73DVH-%DFWHULRO %LQGHU %- DQG &KLVKROP 6: 5HODWLRQVKLSEHWZHHQ '1$ F\FOH DQG JURZWKUDWHLQ Synechococcus VS VWUDLQ3&& - %DFWHULRO %LQGHU %- DQG &KLVKROP 6: &HOOF\FOH UHJXODWLRQ LQ PDULQH Synechococcus VS VWUDLQV $SSO (YLURQ 0LFURELRO %LUG $- 7XUQHU&DYHW -6 /DNH\ -+ DQG 5RELQVRQ 1- $ FDUER[\OWHUPLQDO &\V+LVW\SH ]LQFILQJHU PRWLI LQ '1$ SULPDVHLQIOXHQFHV'1$FRQWHQWLQSynechococcus 3&& - %LRO&KHP
%ORFN 0$ DQG *URVVPDQ $5 ,GHQWLILFDWLRQ DQG SXULILFDWLRQ RI D GHUHSUHVVLEOH DONDOLQH SKRVSKDWDVH IURP Anacystis nidulans 5 3ODQW3K\VLRO %RKQHU+DQG%RJHU3 5HFLSURFDOIRUPDWLRQRIF\WRFKURPH c-553 DQG SODVWRF\DQLQ LQ Scenedesmus. )(%6 /HWW %RY\ $ GH 9ULH]H * %RUULDV 0 DQG :HLVEHHN 3 7UDQVFULSWLRQDO UHJXODWLRQ RI WKH SODVWRF\DQLQ DQG F\WRFKURPH c553 JHQHV IURP WKH F\DQREDFWHULXP Anabaena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ynechococcus 3&& =HQWUDOEODWW IXU 0LNURELRORJLH &ODUN 5% 0DULQH 3ROOXWLRQ 2[IRUG 8QLYHUVLW\ 3UHVV 2[IRUG &RQNOLQ '6 0F0DVWHU -$ &XOEHUWVRQ 05 DQG .XQJ & COTI, D JHQH LQYROYHG LQ FREDOW DFFXPXODWLRQ LQ Saccharomyces cerevisiae. 0RO&HOO%LRO &RRN :- .DU 65 7D\ORU .% DQG +DOO /0 &U\VWDO VWUXFWXUH RIWKH F\DQREDFWHULDO PHWDOORWKLRQHLQ UHSUHVVRU 6PW% $ PRGHO IRU PHWDOORUHJXODWRU\ SURWHLQV - 0RO %LRO 'DQLHOV 0- 7XUQHU&DYHW -6 6HONLUN 5 6XQ + 3DUNLQVRQ -$ 6DGOHU 3- DQG 5RELQVRQ 1- &RRUGLQDWLRQ RI =Q DQG &G E\ SURNDU\RWLF PHWDOORWKLRQHLQ ,QYROYHPHQW RI +LV LPLGD]ROH-%LRO&KHP 'LHOV / 'RQJ 4 YDQ GHU /HOLH ' %DH\HQV : DQG 0HUJHD\ 0 7KH czc RSHURQ RI Alcaligenes eutrophus &+ IURP UHVLVWDQFH PHFKDQLVP WR WKH UHPRYDO RI KHDY\ PHWDOV - ,QG 0LFURELRO 'LQWLOKDF $ $OORLQJ * *UDQDGHO & DQG &ODYHU\V -3 &RPSHWHQFH DQG YLUXOHQFH RI Streptococcus pneumoniae: $GF DQG3VD$PXWDQWVH[KLELWDUHTXLUHPHQWIRU=QDQG0QUHVXOWLQJ IURP LQDFWLYDWLRQ RI SXWDWLYH $%& PHWDO SHUPHDVHV 0RO 0LFURELRO (QGR * DQG 6LOYHU 6 &DG& WKH WUDQVFULSWLRQDO UHJXODWRU\ SURWHLQ RI WKH FDGPLXP UHVLVWDQFH V\VWHP RI Staphylococcus aureus SODVPLGS, -%DFWHULRO (UEH-/ 7D\ORU.% DQG+DOO/0 0HWDOORUHJXODWLRQ RIWKH F\DQREDFWHULDO VPWORFXV LGHQWLILFDWLRQ RI 6PW%ELQGLQJ VLWHV DQG GLUHFW LQWHUDFWLRQ ZLWK PHWDOV 1XFO $FLGV 5HV )RUVWQHU 8 %LQGLQJ IRUPV RI KHDY\PHWDOV LQ VHGLPHQWV DQG VOXGJHV VRUSWLRQ PRELOLVDWLRQ FKHPLFDOH[WUDFWLRQ DQG ELRDYDLODELOLW\ )UHVHQVLXV =HLWVFKULIW )XU $QDO\WLVFKH &KHPLH )UD~VWR GD 6LOYD --5 DQG :LOOLDPV 5-3 =LQF /HZLV DFLG FDWDO\VLV DQG UHJXODWLRQ ,Q 7KH %LRORJLFDO &KHPLVWU\ RI WKH (OHPHQWVSS &ODUHQGRQ3UHVV 2[IRUG )XMLWDQL 6 .RPDQR 7 DQG ,QRX\H 6 $ XQLTXH UHSHWLWLYH '1$ VHTXHQFH LQ WKHMyxococcus xanthus JHQRPH - %DFWHULRO )XNX]DZD+6X]XNL(.RPXNDL<DQG0L\DFKL6 $JHQH KRPRORJRXVWRFKORURSODVWFDUERQLFDQK\GUDVHicfA LVHVVHQWLDO
&KDSWHU 0HWDOV5HSHWLWLYH'1$ WR SKRWRV\QWKHWLF FDUERQ GLR[LGH IL[DWLRQ E\ Synechococcus 3&&3URF1DWO$FDG6FL86$ *KDVVHPLDQ0DQG6WUDXV1$ )XUUHJXODWHV WKHH[SUHVVLRQ RI LURQVWUHVV JHQHV LQ WKH F\DQREDFWHULXP Synechococcus VS VWUDLQ3&&0LFURELRORJ\ *LOVRQ ( &OpPHQW -0 %UXWODJ ' DQG +RIQXQJ 0 $ IDPLO\ RIGLVSHUVHG UHSHWLWLYH H[WUDJHQLF SDOLQGURPLF VHTXHQFHV LQ E. coli (0%2- *LOVRQ ( &OpPHQW -0 3HPQ ' DQG +RIQXQJ 0 3DOLQGURPLFXQLWV DFDVHRIKLJKO\UHSHWLWLYH'1$VHTXHQFHV LQ EDFWHULD 7UHQGV*HQHW *LOVRQ ( 5RXVVHW -3 &OpPHQW -0 DQG +RIQXQJ 0 $ VXEIDPLO\ RI E. coli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ynechococcus 3&& LQYROYLQJ D KLJKO\ LWHUDWHG SDOLQGURPH+,3 0RO 0LFURELRO *XSWD $ :KLWWRQ %$ 0RUE\ $3 +XFNOH -: DQG 5RELQVRQ 1- $PSOLILFDWLRQ DQG UHDUUDQJHPHQW RI D SURNDU\RWLF PHWDOORWKLRQHLQ ORFXV smt LQ Synechococcus 3&& VHOHFWHG IRUWROHUDQFHWRFDGPLXP 3URF5R\6RF /RQGRQ % +XFNOH -: 0RUE\ $3 7XUQHU -6 DQG 5RELQVRQ 1- ,VRODWLRQ RIDSURNDU\RWLF PHWDOORWKLRQHLQ ORFXV DQG DQDO\VLV RI WUDQVFULSWLRQDO FRQWURO E\ WUDFH PHWDO LRQV 0RO 0LFURELRO -DFNPDQ '0 DQG 0XOOLJDQ 0( &KDUDFWHUL]DWLRQ RI D QLWURJHQIL[DWLRQ (nif) JHQHFOXVWHU IURP Anabaena azollae OD VKRZVWKDWFORVHO\UHODWHGF\DQREDFWHULDKDYHKLJKO\YDULDEOHEXW VWUXFWXUHGLQWHUJHQLFUHJLRQV0LFURELRO -HQVHQ 7( %D[WHU0 5DFKOLQ -: DQG -DQL 9 8SWDNH RI KHDY\ PHWDOV E\ Plectonema boryanum &\DQRSK\FHDH LQWR FHOOXODUFRPSRQHQWV HVSHFLDOO\SRO\SKRVSKDWHERGLHV $Q [UD\ HQHUJ\GLVSHUVLYHVWXG\(YLURQ3ROOXW6HULHV$ -L* DQG6LOYHU6 5HJXODWLRQDQGH[SUHVVLRQRIWKHDUVHQLF UHVLVWDQFHRSHURQ IURP Staphylococcus aureus SODVPLG S, - %DFWHULRO -RQHV 0& -HQNLQV -0 6PLWK $* DQG +RZH &- &ORQLQJ DQG FKDUDFWHULVDWLRQ RIJHQHV IRU WHWUDS\UUROH ELRV\QWKHVLV IURP WKH F\DQREDFWHULXPAnacystis nidulans 5 3ODQW 0RO %LRO .DJL-+5DQG6FKDIIHU$ %LRFKHPLVWU\RIPHWDOORWKLRQHLQ %LRFKHPLVWU\ .DPL]RPR$1LVKL]DZD07HUDQLVKL<0XUDWD.DQG.LPXUD$ ,GHQWLILFDWLRQRIDJHQHFRQIHUULQJUHVLVWDQFHWR]LQF DQG FDGPLXP LRQV LQWKH \HDVW Saccharomyces cerevisiae. 0RO *HQ *HQHW .DQDPXUD . .DVKLZDJL 6 DQG 0L]XQR 7 7KH F\DQREDFWHULXP Synechococcus VSHFLHV 3&& SRVVHVVHV WZRGLVWLQFWJHQHVHQFRGLQJFDWLRQWUDQVSRUWLQJ3W\SH$73DVHV )(%6/HWW
.DQDPXUD . .DVKLZDJL 6 DQG 0L]XQR 7 $ FRSSHU WUDQVSRUWLQJ3W\SH$73DVHIRXQGLQWKHWK\ODNRLGPHPEUDQHRI WKH F\DQREDFWHULXP Synechococcus VSHFLHV 3&& 0RO 0LFURELRO .DQHNR7 6DWR6.RWDQL+7DQDND$ $VLPL]X(1DNDPXUD1 0L\DMLPD1 +LURVDZD0 6XJLXUD0 6DVDPRWR 0 .LPXUD7 +RVRXFKL 7 0DWVXQR $ 0XUDNL $ 1DND]DNL 1 1DUXR . 2NXPXUD 6 6KLPSR 6 7DNHXFKL & :DGD 7 :DWDQDEH $
1- 5RELQVRQ-& 5XWKHUIRUG05 3RFRFNDQG-6&DYHW
1LHV '+ &]F5 DQG &]F' JHQH SURGXFWV DIIHFWLQJ UHJXODWLRQ RI FREDOW ]LQF DQG FDGPLXP UHVLVWDQFH F]F LQ $OFDOLJHQHVHXWURSKXV -%DFWHULRO 1LHV '+ DQG 6LOYHU 6 ,RQ HIIOX[ V\VWHPV LQYROYHG LQ EDFWHULDOPHWDOUHVLVWDQFHV-,QG0LFURELRO 1LHV '+ 1LHV $ &KX / DQG 6LOYHU 6 ([SUHVVLRQ DQG QXFOHRWLGH VHTXHQFH RI SODVPLG GHWHUPLQHG GLYDOHQW FDWLRQ HIIOX[ V\VWHP IURP$OFDOLJHQHV HXWURSKXV 3URF 1DWO $FDG 6FL 86$ 1LVKLPXUD$ 7KHWLPLQJRIFHOOGLYLVLRQ$S$DVDVLJQDO 7UHQGV%LRFKHP6FL 2JXUD <
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dnaG SULPDVH LV D ]LQF PHWDOORSURWHLQ %LRFKLP%LRSK\V$FWD 6WHP 0- 3URVVQLW] ( DQG $PHV *)/ 5ROH RI WKH LQWHUFLVWURQLF UHJLRQ LQ SRVWWUDQVFULSWLRQDO FRQWURO RI WKH JHQH
&KDSWHU 0HWDOV5HSHWLWLYH'1$ H[SUHVVLRQ LQ WKH KLVWLGLQH WUDQVSRUW RSHURQ RI 6DOPRQHOOD typhimurium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smt. -,QG0LFURELRO 9DOOHH %/ ,QWURGXFWLRQ WRPHWDOORWKLRQHLQ 0HWK (Q]\PRO 9DOOHH%/DQG)DOFKXN .+ 7KHELRFKHPLFDO EDVLV RI]LQF SK\VLRORJ\ 3K\VLRO 5HY 9DQ GHU /HOLH ' 6FKZXFKRZ 7 6FKZLGHW]N\ 8 :XHUW] 6 %DH\HQV : 0HUJHD\ 0 DQG 1LHV + 7ZRFRPSRQHQW UHJXODWRU\ V\VWHP LQYROYHG LQ WUDQVFULSWLRQDO FRQWURO RIKHDY\ PHWDO KRPRHRVWDVLV LQ $OFDOLJHQHV eutrophus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
:KLWWRQ %$ DQG 6KHKDWD)+$ ,QIOXHQFH RIFREDOW QLFNHO FRSSHUDQG FDGPLXP RQ WKHEOXHJUHHQ DOJD$QDF\VWLVQLGXODQV (QYLURQ3ROOXW6HULHV$ :RRG 50 ,QWHUFKDQJHDEOH FRSSHU DQG LURQ SURWHLQV LQ DOJDOSKRWRV\QWKHVLV(XU-%LRFKHP ;LRQJ$DQG -D\DVZDO 5. 0ROHFXODUFKDUDFWHUL]DWLRQ RID FKURPRVRPDO GHWHUPLQDQW FRQIHPQJ UHVLVWDQFH WR ]LQF DQG FREDOW LRQV LQ 6WDSK\ORFRFFXV aureus. - %DFWHULRO ;X&6KL:DQG5RVHQ%3 7KHFKURPRVRPDOarsR JHQHRI E. FROL HQFRGHV D WUDQVDFWLQJ PHWDOORUHJXODWRU\ SURWHLQ - %LRO &KHP =HQJ- +HXFKHO 5 6FKDIIQHU:DQG .DJL-+5 D 7KLRQHLQ DSRPHWDOORWKLRQHLQ FDQ PRGXODWH '1$ ELQGLQJ DQG WUDQVFULSWLRQ DFWLYDWLRQ E\ ]LQF ILQJHU FRQWDLQLQJ IDFWRU 6S )(%6/HWW =HQJ - 9DOOHH %/ DQG .DJL -+5 E =LQF WUDQVIHU IURP WUDQVFULSWLRQ IDFWRU,,,$ ILQJHUV WR WKLRQHLQ FOXVWHUV 3URF1DWO $FDG6FL86$
&KDSWHU Nostoc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hitton and M Potts (eds), The Ecology of Cyanobacteria, © Kluwer Academic Publishers Pririted in the Netherlands
0DOFROP3RWWV
3DWHQWVDQG$SSOLFDWLRQV $&KURQRORJ\RI 1RVWRF $ 7KH +DLU9HJHWDEOH % 7KH 3KLORVRSKHU¶V6WRQH & )URP%OXH*UHHQ$OJDWR&\DQREDFWHULXP 9,, (SLORJXH
9 9,
6XPPDU\ 6SHFLHV RINostoc DUH IRXQG RQ HYHU\ FRQWLQHQW RQ (DUWK LQ D ZLGH UDQJH RIWHUUHVWULDO DQG DTXDWLF HFRV\VWHPV ZKHUHWKHLUJURZWKVEHFRPHERWKFRQVSLFXRXVDQGDEXQGDQW (QYLURQPHQWVZKHUHZDWHULV VFDUFH RUDEVHQWIRU SURWUDFWHG SHULRGV UHPDLQ VRPH RIWKH PRVWH[WUHPH RQ WKH (DUWK \HW WKH\ W\SLFDOO\ VXSSRUWSRSXODWLRQV RIWKH WHUUHVWULDOGHVLFFDWLRQWROHUDQWIRUPN. commune. 7KHQLWURJHQIL[LQJDFWLYLWLHV RINostoc VSS FRQWULEXWHWR WKH TXDOLW\ RIQXWULHQWSRRUVRLOV HVSHFLDOO\LQNDUVWUHJLRQV DQG VHYHUDOIRUPV FRQWULEXWHWRWKHSURGXFWLYLW\RIULFH SDGGLHVWKURXJKRXWDQH[SDQVLYHUHJLRQRI$VLD&KDSWHU 6RPHNostoc VSSHQWHULQWRDUDQJHRIDVVRFLDWLRQV ZLWK KLJKHU DQG ORZHUSODQWV DQG WKHVH LQFOXGH WKH RQO\ NQRZQ V\PELRVLV ZLWK DIORZHULQJSODQW &KDSWHU 7KHRQO\NQRZQHQGRF\DQRVLV RIDIXQJXV Geosiphon pyriforme, LQYROYHV N. punctijorme DQG PD\UHSUHVHQWD W\SH RISKRWRDXWRWURSKLF DVVRFLDWLRQ ZKLFK ZDV RQH VWHS LQ WKH HYROXWLRQ RIDOO ODQG SODQWV *URZWKV RINostoc HODERUDWH D UDQJH RIQDWXUDO SURGXFWV WKDW UDQJH IURP WKH LQQRFXRXV ZKLFK FRQWULEXWH QR PRUH WKDQ D PLOGO\ RIIHQVLYHRGRUWRSRWDEOHZDWHUVXSSOLHVWRSRWHQWWR[LQVZKLFKFDXVHVLFNQHVVFHOOWUDQVIRUPDWLRQDQGWKHGHDWK RIZLOGOLIHDQGKXPDQV&KDSWHU 'HVSLWHWKHIDFWWKDWNostoc VSSFROOHFWLYHO\UHSUHVHQWDXQLTXHELRORJLFDO UHVRXUFH WKHUH LV QR DGHTXDWH VSHFLHV FRQFHSW IRU WKH JURXS WKH RULJLQV RINostoc DQG RWKHU KHWHURF\VWRXV F\DQREDFWHULDDUHXQFOHDUDQGWKHSK\VLRORJLFDODQGPROHFXODUWUDLWVZKLFKFRXOGSURYLGHGLDJQRVWLFIHDWXUHVRID Nostoc UHPDLQ HOXVLYH &KDSWHU Nostoc LV FROOHFWHG DQG FXOWLYDWHG DV D VRXUFH RIIRRG E\ WKH LQGLJHQRXV SRSXODWLRQV RIPDQ\ FRXQWULHV DQG LQ DWOHDVW RQH RIWKHVH &KLQD Nostoc RFFXSLHV DSRVLWLRQ LQ KXPDQ VRFLDO EHKDYLRULWKDVGRQHVRIRUDWOHDVW \HDUV 7KLVFKDSWHUFRQVLGHUVLQDEURDGVHQVHZKDWLVNQRZQDQGZKDW QHHGVWREHNQRZQDERXWNostoc VSS
, ,QWURGXFWLRQ 7KHUH LV DFRQVLGHUDEOH DFFXPXODWLRQ RIOLWHUDWXUH RQ Nostoc VSSDQGDQH[WHQVLYHQXPEHURIVWXGLHVDUHLQ WKH &KLQHVH OLWHUDWXUH 7KH VWXGLHV RI 0ROOHQKDXHU D E D E UHFHQWDFFRXQWV E\ 'RGGV HW DO DQG *DR WKH UHYLVLRQ RI .RPiUHN DQG $QDJQRVWLGLV DQG WZR UHYLHZV 3RWWV IRFXVHGRQDVSHFWVRIWKH HFRSK\VLRORJ\ ELRFKHPLVWU\ DQG FHOO VWUXFWXUH RI Nostoc. $ FRPSUHKHQVLYH FRQVLGHUDWLRQ RI WKH PHWDEROLVP DQG GHYHORSPHQW RI KHWHURF\VWV DQG D GHVFULSWLRQ RIWKH XVH RINostoc VSS DV JHQHWLF WRROV ZLWK ZKLFK WR XQGHUVWDQG WKH PROHFXODU EDVLV IRU KHWHURF\VWIRUPDWLRQ DSSHDUHG UHFHQWO\ :RONHWDO :RON 7KHSUHVHQWDQG OLPLWHG DFFRXQW Abbreviations: $%$ DEVFLVLF DFLG '6%¶V GRXEOHVWUDQG EUHDNV (36 H[WUDSRO\VDFFKDULGH +,3 KLJKO\LWHUDWHG SDOLQGURPLF UHSHDW VHTXHQFH 3$6 SHULRGLF DFLG6FKLII UHDJHQW 6755 VKRUWWDQGHPUHSHDWUHSHWLWLYHVHTXHQFH 895 XOWUDYLROHW UDGLDWLRQ:VSZDWHUVWUHVVSURWHLQV
LV QRW D UHYLHZ ,W LV D VHOHFWLYH FRQVLGHUDWLRQ WKDW HPSKDVL]HV DVSHFWV RIPROHFXODU HFRORJ\ LQ RUGHU WR SODFH FRQFHSWV UDLVHG E\ RWKHU FRQWULEXWRUV WR WKLV YROXPHLQSHUVSHFWLYH
,, 1DWXUDO &RPPXQLWLHV $ 1LFKHV .DUVWDQG7LQWHQVWULFK 6SHFLHVRINostoc HQMR\DUDQJHWKDWLVSUHGRPLQDQWO\ WHUUHVWULDO DQG GHVFULSWLRQV IURP DTXDWLF V\VWHPV DUH DOPRVW H[FOXVLYHO\ WR GR ZLWK IUHVKZDWHU IRUPV ,Q IDFW .RPiUHN DQG $QDJQRVWLGLV FRQFOXGHG WKDW DOO RI WKH GDWD DERXW Nostoc VSHFLHV IURP VXEPHUVHGDQGPDULQHELRW\SHVFRQFHUQRWKHUWD[D 2QHHQYLURQPHQWWKDWFOHDUO\GRHV QRWDSSHDUWREH FRORQL]HG E\ Nostoc LV WKH LQWHULRU RI URFNV 7KLV PXVWEHGXHWRVRPHSK\VLRORJLFDOFRQVWUDLQWEHFDXVH RWKHU F\DQREDFWHULD LQFOXGLQJ KHWHURF\VWRXV IRUPV
&KDSWHU 1RVWRF
Fig. 1. Nostoc commune. D .DUVW VFHQHU\ ³6XJDU´ OLPHVWRQH LQ WKH DUHD DURXQG 0DOKDP 7DP ,QJOHERURXJK (QJODQG &RPPXQLWLHV RI Nostoc commune DFFXPXODWH LQ WKH VKDOORZ GHSUHVVLRQV RI WKHURFNWKDWSHULRGLFDOO\ILOOZLWKUDLQZDWHUE )XOO\K\GUDWHG FRORQLHVWKDWKDYHWKHFRQVLVWHQF\RIDVHPLULJLGJHO
VXFK DV Mastigocoleus testarum, Kyrtuthrix dulmatica DQG Scytonema endolithicum, GR LQIHVWWKH URFNV DQG VHGLPHQWV RI IUHVKZDWHU WHUUHVWULDO DQG PDULQHORFDOLWLHV&KDSWHUV $ FXUVRU\ LQVSHFWLRQ RI QXWULHQWSRRU VRLOV HVSHFLDOO\WKRVHLQDUHDVZKHUHOLPHVWRQHLVSUHYDOHQW RIWHQ UHYHDOV WKH SDUFKPHQWOLNH FRORQLHV RI N. commune 6FKHUHU :KHQ GU\ WKH FRORQLHV IRUP GDUNHQHG FUXVWV ZLWK WKH FRQVLVWHQF\ DQG DSSHDUDQFH RIEXUQHG EDFRQ ULQGV 7KHVH FUXVWV DUH
UHDGLO\GHWDFKHG DQG FDQ DFFXPXODWH LQ URFNKROORZV WKDWSHULRGLFDOO\ILOOZLWKUDLQZDWHU3RWWV 7KH FKDUDFWHULVWLF VWUDQGV RI N. commune YDU flagelliforme FRYHU H[SRVHG URFNV LQ VHPL DULG DUHDV VXFK DV WKH *REL 'HVHUW LQ ,QQHU 0RQJROLD 6FKHUHU DQG=KRQJ Nostoc commune LV HVSHFLDOO\ SUHYDOHQW LQ OLPHVWRQHRUNDUVWUHJLRQVVXFKDVWKRVHRIWKH%XUUHQ &R &ODUH (LUH 0DOKDP 7DUQ (QJODQG DQG WKH $GULDWLFFRDVWRIWKHIRUPHU-XJRVODYLD)LJ OD 7KH
&DKHU 5LYHU GUDLQV WKH OLPHVWRQHULFK %XUUHQ DQG SURYLGHV QXPHURXV KDELWDWV ZKHUH PDFURVFRSLF FRORQLHVRINostoc DQG Rivularia FDQEHIRXQG 3RWWV XQSXEOLVKHG GDWD )LJ E /LPHVWRQH SHEEOHV WKDW DUH XVHG DV D EDOODVW WR FRYHU URRIV RI XQLYHUVLW\ EXLOGLQJV VFKRROKRXVHV DQG WKH OLNH LQ WKH 8QLWHG 6WDWHV DV ZHOO DV JUDYHO SDWKV ³.LHVZHJ´ LQ *HUPDQ DQGIODWFRUDOOLPHVWRQHSDYHPHQW³SODWLQ´ LQ)UHQFK&UHROH LQWKH7URSLFV3ODWHF SURYLGH WKHSHUIHFWFRQGLWLRQV RIZDWHUUHWHQWLRQDQG GUDLQDJH IRU Nostoc JURZWK Tintenstrich FRPPXQLWLHV *HUPDQ IRU ³LQN VWULS´ UHFHLYH WKHLU QDPH EHFDXVH RI WKH VWUHDNV VXFK F\DQREDFWHULDO JURZWKV GLVSOD\ SDUWLFXODUO\ LQ PRVW VSHFWDFXODU IRUP RQ D EDFNJURXQG RI YHUWLFDO ZKLWH OLPHVWRQHV DQG GRORPLWHV /LLWWJH 3ODWH E Nostoc PD\ RFFXULQ VXFK FRPPXQLWLHV EXW GRHV VR W\SLFDOO\DVPRUHPDWXUHFUXVWVWKDWFDQDOVRIRUPRQ JUDQLWLFDQGVDQGVWRQHVXEVWUDWHV/XWWJH 7KH S+ UHTXLUHPHQWV RI Nostoc VSS IURP OLPHVWRQH UHJLRQV VXJJHVW D SUHIHUHQFH IRU DONDOLQH FRQGLWLRQV%LLGHOHWDO DOWKRXJKDVWUDLQRIN. microscopicum LVRODWHG IURP D ULFH ILHOG LQ WKH 3KLOLSSLQHVVKRZHGRSWLPXPJURZWKLQWDSZDWHUDWD S+EHWZHHQDQG(PUDOLQRDQG5RGXOIR
6RLOVDQG:DWHU 7KHUH LV QR VKRUWDJH RI PLFURELDO LQRFXOD DQ\ZKHUH RQ WKH SODQHW DQG DQ\ DSSDUHQWUHVWULFWHG GLVWULEXWLRQ RIDJLYHQIRUPPXVWEHHLWKHUDUHDORQH ZLWKVRPH GLVWLQFW HFRORJLFDO EDVLV RU WKH UHIOHFWLRQ RI LQVXIILFLHQW GRFXPHQWDWLRQ RI VSRUDGLF JURZWKV )RU H[DPSOHWKHVRLOV LQFHUWDLQSDUWVRIWKH'U\9DOOH\V $QWDUFWLFDDUHWUXO\VWHULOHDQGWKLVLVQRWIRUZDQWRI LQRFXOD 0ROOHQKDXHU SURYLGHG DQ H[KDXVWLYH OLVWLQJ RI VLWHV ZKHUH N. punctiforme ZDV UHFRUGHG DQG FRQFOXGHG WKDW WKH DSSDUHQW ODFN RIWKLV IRUP LQ $XVWUDOLD DQG $QWDUFWLFD ZDV SUREDEO\ UHDO +RZHYHU 6HDEXUJHWDO OLVWHGVHYHQGLIIHUHQW Nostoc VSHFLHV LQFOXGLQJ N. punctiforme, IURP D OLWWRUDO EHQWKLF PDW LQ /DNH %RQQH\ 7D\ORU 9DOOH\ 6RXWKHUQ 9LFWRULD /DQG $QWDUFWLFD &XULRXVO\ WKH VHYHQVSHFLHVZHUHQRWUHFRUGHGIURPWKHRWKHU/DNHV LQWKHYDOOH\/DNHV)U\[HOO+RDUH&KDGDQG&KRLFH GHVSLWH WKHLU UHODWLYH SUR[LPLW\ WR RQH DQRWKHU ,Q FRQWUDVW N. commune ZDVSUHVHQWWKURXJKRXWWKHDUHD LQ D UDQJH RI KDELWDWV LQFOXGLQJ WKH SODQNWRQ VRLOV DQG JODFLDO PHOWVWUHDPV &KDSWHU 7KHVH PDUNHG GLIIHUHQFHV LQ GLVWULEXWLRQ SDWWHUQV RIGLIIHUHQW IRUPV DSSHDU FRQYLQFLQJ VR ZK\ VKRXOG VXFK GLIIHUHQFHV KDYH DULVHQ" )RU WKH PRVW SDUW WKH GLVWULEXWLRQ RIN.
0DOFROP3RWWV commune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ostoc VSS DQGRWKHUF\DQREDFWHULDNostoc, DOWKRXJKVFDUFHZDV SUHVHQW LQ WKH UHPDUNDEOH FRQVRUWLD ZKLFK GHYHORS ZLWKLQVRODUJHQHUDWHGZDWHUSRFNHWVLQWKHSHUPDQHQW LFHFRYHUVRI$QWDUFWLFODNHV3ULVFXHWDO $Q DEXQGDQW JURZWK RI N. microscopicum ZDV GHWHFWHG DW DGHSWK RIP ZLWKLQ D FDYH LQ &ODUHQV VDQGVWRQH RI WKH 'UDNHQVEHUJ 0RXQWDLQV 6RXWK $IULFDZKHUHVHHSDJHZDWHUS+WR ZDVWKHRQO\ VRXUFH RI PRLVWXUH %XGHO HW DO %URZQ JUDQXOHVRINostoc ZHUHUHSRUWHG IURPZLWKLQDZDWHU VXSSO\WDQNZKHUHLOOXPLQDWLRQZDV VFDUFH+XHWDO ZKLOH PRUH H[WHQVLYH JURZWKV RFFXU LQ WKH K\SHUWURSKLF FRDVWDO ODJRRQ LQ $OEXIHUD 9DOHQFLD ( 6SDLQ0RUHQRHWDO 6WXGLHVRIWKHJURZWKRI Nostoc parmelioides LQ VWUHDPV LQGLFDWHG D FOHDU SUHIHUHQFH IRUURXJKDV RSSRVHG WR VPRRWK VXEVWUDWHV 'XGOH\DQG'¶$QWRQLR N. parmelioides ZDV OLWWOH DIIHFWHG E\ JUD]LQJ RI WKH FDGGLVIO\ Agapetus celatus EXW JUHDWO\ DIIHFWHG E\ GLVWXUEDQFH DQG RQO\ EDVDOFUXVWVZHUHDEOHWRZLWKVWDQGGLVWXUEDQFHVLQWKH DEVHQFHRIVHYHUHVFRXU 7KH ZLGHVSUHDG XVH RISKRVSKDWH DQG QLWUDWHEDVHG IHUWLOL]HUV KDV DOO EXW HUDGLFDWHG Nostoc IURP PDQ\ VRLOV LQFOXGLQJ UHVLGHQWLDO JUDVV ODZQV ZKHUH LW ZRXOG RWKHUZLVHSUROLIHUDWH ,Q &KLQD WKHFROOHFWLQJ RI N. commune YDU flagelliforme IRU IRRG DQG SDVWXULQJRIFDWWOHDUHGLPLQLVKLQJWKLVUHVRXUFH *DX 5HSRUWV RINostoc IURP VRPH OP GHSWK LQ WKH ,QGLDQ2FHDQDQGWKH0HGLWHUUDQHDQDUHTXHVWLRQDEOH XQFRQILUPHGDQGWKHUHIRUHUHPDLQLQWULJXLQJ%HUQDUG %HUQDUGDQG/HFDO
$OGDEUD$WROO $OGDEUD$WROO LQWKHH[WUHPH6RXWKZHVWFRUQHURIWKH ,QGLDQ 2FHDQ RIIHUV DQ XQVXUSDVVHG VHWWLQJ LQ ZKLFK WR VWXG\ WKH HFRORJ\ RIERWK IUHVKZDWHU 'RQDOGVRQ :KLWWRQ 'RQDOGVRQ DQG 3RWWV DQG PDULQH 3RWWV 3RWWV DQG :KLWWRQ
&KDSWHU 1RVWRF F\DQREDFWHULD Nostoc commune LV FRPPRQ DPRQJ WKH JUDVV Sporobolus virginicus, HVSHFLDOO\ LQ JURYHV RI Cocos nucifera FRFRQXW RQWKHDWROO ZKHUHGDLO\ WHPSHUDWXUHV PD\ UHDFK WKH PLG V FHQWLJUDGH 6KDOORZ ZDWHUILOOHG GHSUHVVLRQV LQ OLPHVWRQH ³SODWLQ´ VXSSRUW GHQVH JURZWKV RIWKH FKDUDFWHULVWLF VSKHULFDOFRORQLHVRIN. sphaericum; LQGULHUDUHDVWKH FRORQLHVWHQGWREHOHDWKHU\LUUHJXODUO\OREHG RUHYHQ KDLUOLNHLQDSSHDUDQFH ,WVHHPVFHUWDLQWKDWSK\VLFDO FRQGLWLRQV SOD\ D PDMRU UROH LQ GHWHUPLQLQJ WKH DSSHDUDQFH RI WKHVH FRPPXQLWLHV EXW FDQ RQH EH FHUWDLQ WKDWWKHVH IRUPV DUH JHQRW\SLFDOO\ HTXLYDOHQW RU LV WKHUH D FRQWLQXXP RIJHQHWLF GLYHUVLW\ &KDSWHU " 7KH UDWHV RIQLWURJHQ IL[DWLRQ SHU XQLW FKORURSK\OO UHFRUGHG IRU VRPH WHUUHVWULDO Nostoc SRSXODWLRQV RQ $OGDEUD DUH DPRQJ WKH KLJKHVW UHFRUGHG LQ WKH OLWHUDWXUHIRULQVLWXVWXGLHVRIF\DQREDFWHULD:KLWWRQ HWDO &KDSWHU 'XULQJWKHGU\VHDVRQZKHQ WKH VRXWKHDVW WUDGH ZLQGV EORZ 0D\ WKURXJK 1RYHPEHU FRORQLHV DUH GHVLFFDWHG IRU ORQJ SHULRGV 'XULQJ WKH ZHW VHDVRQ FDXVHG E\ WKH ZHVWQRUWKZHVW PRQVRRQ'HFHPEHUWKURXJK$SULO WKHFRPPXQLWLHV RQ DGDLO\ EDVLV XQGHUJR DFODVVLFSDWWHUQ RIZHWWLQJ DQG GU\LQJ 'HVLFFDWHG FRORQLHV RI N. commune IURP $OGDEUD ZHUH IRXQG WR WDNH XS ZDWHU VLJQLILFDQWO\IDVWHUWKDQWKRVHRIHLWKHUN. sphaericum RU D Nostoc-Phormidium FRPPXQLW\ UDWHV RIXSWDNH ZHUH DOVR IDVWHU WKDQ E\ N. commune IURP D WHPSHUDWH ORFDOLW\ 7DUQ 0RRU (QJODQG ,Q IDFW DERXW KDOIWKH PD[LPDO UDWH RIQLWURJHQ IL[DWLRQ ZDV DWWDLQHG DIWHU UHZHWWLQJ RIWKH FRORQLHV IRU RQO\ RQH KRXU:KLWWRQHWDO )UHHOLYLQJ Nostoc VSS DV ZHOO DV Tolypothrix byssoidea DQG VRPH OLFKHQL]HG FRPPXQLWLHV RQ $OGDEUD FRQWULEXWH QLWURJHQ WR WKH VFDUFH VKDOORZ DQGLPSRYHULVKHGVRLOV7KHLPSDFWRIQLWURJHQIL[LQJ FRPPXQLWLHV LQFOXGLQJ F\DQREDFWHULD LV DSSDUHQW 7KHVHVRLOVVXSSRUWDJUDVVDQG VKUXEYHJHWDWLRQWKDW LQ WXUQ VXVWDLQV D SRSXODWLRQ RI PRUH WKDQ JLDQW ODQG WRUWRLVHV (Geochelone gigantea); PDQ\ PHPEHUVRIZKLFKKDYHZHLJKWVLQH[FHVVRINJ
% 7KH&RORQ\ 3K\VLFDO3URSHUWLHV Nostoc VSS GHYHORS FRORQLHV WKDW KDYH D UDQJH RI FKDUDFWHULVWLF VKDSHV VL]HV VPHOOV WH[WXUHV DQG FRORUV DQG PDQ\ EHFRPH ERWK DEXQGDQW DQG YLVXDOO\ FRQVSLFXRXV ,I WKHUH LV D SHUVLVWHQW VRXUFH RI SUHFLSLWDWLRQ WKHVH FRORQLHV PD\ DVVXPH GUDPDWLF
SURSRUWLRQV &RORQLHV RIN. pruniforme, DV ODUJH DV 25 FP LQ GLDPHWHU JURZ LQ 0DUH¶V (JJ 6SULQJ 2UHJRQ 86$ 'RGGV DQG &DVWHQKRO] 3ODWH D 'HSHQGLQJ RQ WKH IUHTXHQF\ RI ZHWWLQJ N. commune PD\ IRUP VSKHUHV ³SHDUOV´ IURP WHQV RI PP WR FP LQ GLDPHWHU RU GLVFRLG DQG IODWWHQHG FUXVWV0DUWLQH]DQG4XHULMHUR 7KHIRUPDWLRQ RIVSKHULFDO PDFURVFRSLF FRORQLHV GHVHUYHV PHQWLRQ 2WKHUF\DQREDFWHULDDV ZHOO DVVRPHHXNDU\RWLFDOJDH WKDW GR QRW DWWDFK WR VXEVWUDWD FDQ IRUP VSKHULFDO FRORQLHV DHJDJURSLOHV DQG ³6HHNQLLGHO´ RI DSSUHFLDEOH VL]H ZKHQ WKH\ JURZLQ VKHOWHUHG ZDWHUV ZKHUH WKHUH LV RQO\ SHULRGLF DJLWDWLRQ 7KLV LV D SDVVLYH SK\VLFDO SKHQRPHQRQ HJ Scytonema; 3RWWV DQG:KLWWRQ 6SKHULFDO FRORQLHV RINostoc GR DULVH LQ ERWK VXEPHUJHG DQG H[SRVHG VLWXDWLRQV EXW WKHLUIRUPDWLRQLVFHUWDLQO\QRWSDVVLYH LWUHSUHVHQWVD GLVWLQFW SDUW RI WKHLU OLIH F\FOH 7KH FKDUDF WHULVWLF³KDLUOLNH´ FRORQLHV RI N. commune YDU flagelliforme VKRZ GLIIHUHQW FKDUDFWHULVWLFV RI ZDWHU ORVV DQG UHWHQWLRQ WKDQ N. commune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commune FROOHFWHG in situ, DQG RI ³SHDUOV´ 3ODWH G ZKLFKEXGIURPWKHWKDOOXV 3RWWV LVWKDW WKH LQWHULRU RI WKH FRORQ\ LV GHYRLG RI DQ\ RWKHU PLFURRUJDQLVPV VDYH IRU N. commune.
0DOFROP3RWWV
Fig. 2. 'LVWULEXWLRQVRIFHOOOHQJWKVDQGFHOOZLGWKVLQYHJHWDWLYHFHOOV9 KHWHURF\VWV+ DQGDNLQHWHV$ RIDOONostoc VSHFLHVGHVFULEHG PD[LPXP E\ *HLWOHU D &HOO OHQJWK GLVWULEXWLRQ E &HOO ZLGWK GLVWULEXWLLRQ PLQLPXP PHDVXUHPHQW IRU D JLYHQ VSHFLHV PHDVXUHPHQW IRU DJLYHQVSHFLHV
&KDSWHU 1RVWRF GLVDSSHDUDQFH RISHULSKHUDO JUDQXOHV IURP YHJHWDWLYH FHOOV RI WKLV VWUDLQ GXULQJ H[SRQHQWLDO JURZWK FRLQFLGHG ZLWK WKH UHOHDVH RI D EURZQ SLJPHQW &XULRXVO\WKHEURZQSLJPHQW DQGDQXPEHURI895 DEVRUELQJ SLJPHQWV ZHUH QRW REVHUYHG XQGHU ORZ R[\JHQ FRQGLWLRQV QRU ZHUH WKH SLJPHQWV SURGXFHG ZKHQ FHOOV ZHUH JURZQXQGHUDK KOLJKW GDUN F\FOH
0DFURPROHFXODU&RPSRQHQWV ³7KH SXULILFDWLRQ RI WKH PXFLODJH ZDV GLIILFXOW DQG WHGLRXVDQGWKHK\GURO\VLVSURGXFWVVRFRPSOH[DVWR UHQGHU DQDO\VLV GLIILFXOW´ WKHVH FRPPHQWV ZHUH PDGH E\ +RXJK HW DO IROORZLQJ DWWHPSWV WR SXULI\ WKH (36 RI JHODWLQRXV QRGXOHV RI D Nostoc VS FROOHFWHG IURP DURXQG WKH )UHVKZDWHU %LRORJLFDO $VVRFLDWLRQ $PEOHVLGH LQ (36 ZDV H[WUDFWHG IURP WKH FXOWXUH OLTXLG RI Nostoc VS VWUDLQ ' DQG IRXQG WR FRQWDLQ DUDELQRVH DQG JDODFWXURQLF DFLG ULERVH [\ORVH JOXFRVH DQG WZR XQLGHQWLILHG FRPSRXQGV 6(0UHYHDOHGWKHPXFLODJHKDG YHOXP OLNH DQG QHWOLNH VWUXFWXUHVWKDWZHUH DOVR VHHQLQWKH LVRODWHG SRO\VDFFKDULGH &XSDF DQG *DQWDU 7KHUH DUH UHSRUWV WKDW WKH VKHDWK PDWHULDO RI N. commune FRQWDLQV D JUDQXODU FRPSRQHQW DVVRFLDWHG ZLWK WKH ILEULOODU QHWZRUN SRVVHVVLQJ VRPH PRUSKRORJLFDO DQG SK\VLRORJLFDO VLPLODULWLHV WR WKH PDWUL[ JUDQXOH RI DQLPDO FDUWLODJH FROODJHQ )MHUGLQJVWDG HW DO $EGHODKDG HW DO .RPiUHN DQG $QDJQRVWLGLV 6\QWKHVLV RI WKH VWUXFWXUDO SURWHLQ FROODJHQ WKDW LV XQLYHUVDOO\ GLVWULEXWHG DPRQJ PHWD]RDQV RQO\ EHFDPH SRVVLEOH ZKHQ WKH R[\JHQ FRQFHQWUDWLRQ LQ SULPLWLYH (DUWK URVH IURP OHVV WKDQ WR RI SUHVHQW DWPRVSKHULF OHYHOV &DQILHOG DQG 7HVNH $EXQGDQWSURWHLQVVHHEHORZ DUHSUHVHQWLQWKH(36 RI N. commune DQG PD\ FRQWULEXWHWRWKH FDUWLODJH OLNH SURSHUWLHV RI WKH (36 PDWUL[ 7KH ILEURXV VWUXFWXUH W\SLFDOO\ VHHQ LQ WKLQ VHFWLRQV RI WKH (36 3RWWV LVJHQHUDOO\LQGLFDWLYHRIJHOV .XPDU HW DO GHPRQVWUDWHG WKH HIILFLHQW 895% VKLHOGLQJ UROH RI )H&O DQG SLJPHQWHG SURGXFWV IURP N. spongiaeforme ZKHQ WKH\ ZHUH LQFRUSRUDWHGLQ PPWKLFNDJDURVHJHOV ,W V D SLW\ WKHVH H[SHULPHQWV ZHUH QRW DOVR FRQGXFWHG ZLWK WKH VDPH JHOV WKDW KDG EHHQ GULHG WR D WKLQ ILOP WR VLPXODWH WKH IRUP RI D GULHG Nostoc FRORQ\ 2WKHU H[WUDFHOOXODU SLJPHQWV LQFOXGHG D YLROHW SLJPHQW QRVWRFLQH $ PHWK\O SHQWD D]DELF\FOR>@QRQDQ WULHQRQH ZLWK D
EURDG VSHFWUXP RI JURZWK LQKLELWRU\ DFWLYLW\ +LUDWD HWDO $OO RI WKH Nostoc commune SURWHLQV LGHQWLILHG WR GDWH LQ WKH (36 ZHUH FDUERK\GUDWHPRGLI\LQJ SURWHLQV DQG WKHLU ELRFKHPLFDO SURSHUWLHV ZHUH VWXGLHGLQGHWDLO 6FKHUHUDQG3RWWV +LOOHWDO D 3RWWV 7KH (36 RI Nostoc commune KDV XQLTXHUKHRORJLFDOSURSHUWLHV WKDWPD\ EH GHWHUPLQHG LQ SDUW E\ WKHVH VHFUHWHG SURWHLQV 3RWWV +HOPDQG3RWWVXQSXEOLVKHGGDWD
& &HOOV 2UJDQL]DWLRQ &HOOV RINostoc VSS DUH VSKHULFDO EDUUHOVKDSHG RU RYDO DQG WKHLU FRQWLJXRXV DUUDQJHPHQW OHDGV WR WKH IRUPDWLRQ RI XQEUDQFKHG ILODPHQWV ZKLFK LIVWUDLJKW VPDOO ZULWKH ODUJH WZLVW PD\ H[FHHG PP LQ OHQJWK 7KHZLGWKRIFHOOVLQWKHRUVRGHVFULSWLRQV DQQRWDWHG E\ *HLWOHU YDU\ RYHU DSSUR[LPDWHO\ DRQH RUGHURIPDJQLWXGHIURPDPLQLPXP RIDURXQG PP WR D PD[LPXP RI DURXQG PP )LJ D +HWHURF\VWV DQG DNLQHWHV 6HFWLRQ ,,,& WHQG WR EH EURDGHUDQGORQJHUWKDQYHJHWDWLYHFHOOV)LJE 7KH VPDOOHVWNostoc UHFRUGHGKDV DFHOO ZLGWKWKDW LV RQH ILIWLHWK WKDW RI Chroococcus turgidus 87(; DUHSUHVHQWDWLYHRIDJURXS ZKLFKPD\UHSUHVHQW WKH ODUJHVW RIWKH F\DQREDFWHULD 3RWWV HW DO 7KHVH DUH SXUHO\ HPSLULFDO REVHUYDWLRQV DQG ZLWKRXW DQ\ELRSK\VLFDOPHDVXUHPHQWVLWLV KDUG WR PDNH DQ\ JHQHUDOL]DWLRQVDVWRWKHLUVLJQLILFDQFH.RFK 7KH ILODPHQWV RI N. commune DUH HPEHGGHG ZLWKLQDQGGLVWULEXWHGWKURXJKRXW DGHQVH(36 7KH RYHUDOO DUUDQJHPHQWRIILODPHQWV LQWKH(36 DQG WKHLU VWUXFWXUDO DQG ELRFKHPLFDO SURSHUWLHV ZHUH GHVFULEHG LQGHWDLO+LOOHWDO E3RWWV 3RWWV 2QHVWULNLQJIHDWXUHRIWKHGLVWULEXWLRQRIILODPHQWV LQ ³SHDUOV´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
0DOFROP3RWWV
0DJQHVLXP FDOFLXP VLOLFRQ SKRVSKRUXV DQG VXOIXU SUHGRPLQDWHG WKH ('; VSHFWUXPRIDOO PDWHULDOV WKDW ZHUH VWXGLHG 5HODWLYH WR WKH JO\FDQ FHOOV ZHUH HQULFKHGLQ VXOIXUDQGSKRVSKRUXV 7KHPRVWREYLRXV UHVXOW RI WKHVH DQDO\VHV ZDV WKH LGHQWLILFDWLRQ RI D FRQVSLFXRXV DFFXPXODWLRQ RI VLOLFRQ DQG FDOFLXP LQ WKHH[WHUQDOOD\HU /LJKWPLFURVFRS\DQG7(0UHVROYHGWKHLPPHGLDWH HQYLURQPHQW RI HDFK FHOO LQ GHVLFFDWHG FRORQLHV RI ILHOG PDWHULDO RI N. commune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commune &+(1 SHDUOV RU N. commune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³WXQQHO´OLNH VWUXFWXUHV GHVFULEHG DERYH ZHUH DEVHQW ILODPHQWV ZHUH HQFDVHG LQ D OD\HU ZKLFK KDG GLIIHUHQW VWUXFWXUDO SURSHUWLHV WR WKH EXON (36 $QDO\VLV WKURXJK OLJKW PLFURVFRS\ DOVR LQGLFDWHG GLIIHUHQW VWDLQLQJ SURSHUWLHV RI WKLV OD\HU )LODPHQWV WHQGHG WR RFFXS\ WKH RXWHU SRUWLRQV RI WKH FRORQ\ ZKHUH WKH\ IRUPHG VPDOOHU ³SHDUOV´ DQG SDFNHWV ZKLOH QRQHQFDVHG ILODPHQWV ZHUH SUHVHQW LQ WKH FHQWUDOSRUWLRQVRIHDFK³SHDUO´ 'XULQJ JURZWK PLQXWH ³SHDUOV´ EXG RII IURP WKH SDUHQWSHDUO DWWKHSHULSKHU\ RIWKH ODWWHU 3ODWH G 7KHVH EXGV RULJLQDWH DV HQFDVHG ILODPHQWV LPPHGLDWHO\EHORZWKHVXUIDFHRIWKHSHDUODQGDWWKLV VWDJH DQ H[WHUQDO OD\HU LV DOUHDG\ DSSDUHQW DW WKH
SHULSKHU\ RI WKHVH VPDOO SDFNHWV RI ILODPHQWV 7KH VWDLQLQJ FKDUDFWHULVWLFV RI WKH PDWHULDO LQ DQG VXUURXQGLQJ WKHVH SDFNHWV GLIIHUHG IURP WKRVH RIWKH SDUHQW JO\FDQ VKHDWK PDWUL[ )RU H[DPSOH ZLWK $OFLDQ EOXH DW S+ WKH VPDOO SDFNHWV VWDLQHG DQ LQWHQVH GDUN EOXH LQ FRPSDULVRQ WR WKH OLJKW EOXHJUHHQFRORURIWKHVXUURXQGLQJJO\FDQ+LOOHWDO E
8OWUDVWUXFWXUH 8OWUDVWUXFWXUDO VWXGLHV RI Nostoc VSS OLNH WKRVH RI F\DQREDFWHULDLQJHQHUDO WHQGHGWRIRFXVRQWKHIRUP DQGRFFXUUHQFHRIFHOOLQFOXVLRQV$OOHQ -HQVHQ 6LPRQ 7ZHQW\ QLQH Nostoc LVRODWHV FRQWDLQHG GLIIHUHQWW\SHV RILQFOXVLRQV 2WKHUWKDQ D VHHPLQJ XELTXLWRXV RFFXUUHQFH RI PHPEUDQH OLPLWHG FU\VWDOOLQH ERGLHV WKHUH ZDV QR REYLRXV WUHQG LQ DVVRFLDWLRQ RI FHUWDLQ VWUXFWXUHV ZLWK SDUWLFXODU IRUPVRINostoc. &\WRFKHPLFDO DQDO\VLV UHYHDOHG WKDW D KLJK VDOW HQYLURQPHQW DFWLYDWHG DQ RXEDLQVHQVLWLYH 1D . $73DVH SUHVXPDEO\ LQYROYHG LQ HIIOX[ RI 1D LRQV ,ZDQR ([SRVXUH RI Nostoc VS 8$0 DQG RWKHU F\DQREDFWHULD WR WKH LQVHFWLFLGH WULFKORUIRQ OHDG WR GHVWDELOL]DWLRQ RI WKH KHWHURF\VW HQYHORSH 7KLV PD\ EH VHOHFWLYH IRU FRPPXQLWLHV LQ VLWX DV WKH HIIHFWVRIWKHLQVHFWLFLGHZHUHODUJHO\SUHYHQWHGXQGHU DQDHURELFFRQGLWLRQV 2UXVDQG0DUFR 0DQ\ VWXGLHVIRFXVHGRQWKHXOWUDVWUXFWXUHRINostoc VSS LQ LQWHUDFWLRQV ZLWK SODQWV *DQWDU HW DO -RKDQVVRQ DQG %HUJPDQ DQGHIIHFWV RIKHDY\ PHWDOV *XSWD DQG 6LQJKDO )HUQDQGH]SLQDV HW DO
' 6WUXFWXUHDQG)XQFWLRQ ([RSRO\VDFFKDULGH $ QXPEHU RIGLIIHUHQW DWWULEXWHV ZHUH DVFULEHG WR WKH H[WUDFHOOXODU (36 RIEDFWHULD3RWWV +LOO HW DO 7KH DOJLQDWH RI Pseudomonas aeruginosa UHVWULFWVGLIIXVLRQRIR[\JHQDIHDWXUHZKLFKLVVKDUHG E\F\DQREDFWHULDO(36V&KDQJ ([SHULPHQWV XVLQJ OLJKW PLFURVFRS\ LGHQWLILHG D FRQVSLFXRXV UHDFWLRQ RI KHWHURF\VWV LQ GHVLFFDWHG PDWHULDO ZLWK 3$6 D VWDLQ XVHG LQ JO\FRSURWHLQ GHWHFWLRQ +LOO HW DO E ,Q IXOO\ K\GUDWHG SHDUOV D PRUH JHQHUDO DQG FRQVSLFXRXV UHDFWLRQ ZLWK 3$6 ZDV REVHUYHG 7ZR SRO\SHSWLGHV WKDW VKRZHG D SRVLWLYH JO\FDQ UHDFWLRQ ZHUH GHWHFWHG LQ DTXHRXV H[WUDFWV RI
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commune '5+O LV WKDW D VSKHUH UHSUHVHQWV WKH PLQLPXP VXUIDFH DUHD IRU D JLYHQ YROXPHZKLFKZRXOGFOHDUO\SURYLGHDUHGXFWLRQ DQG XQLIRUPLW\ LQWKHQHWUDWHRIHYDSRUDWLRQRIZDWHU 2I FRXUVH IRU WKH VDPH FRQVLGHUDWLRQV WKH VSKHULFDO JO\FDQVXUIDFHKDV DUHGXFHGFDSDFLW\IRUJDV XSWDNH &\DQREDFWHULDO VKHDWK PDWHULDOV ZHUHIRXQG WRUHWDUG JDV H[FKDQJH &KDQJ 6SKHULFDO FRORQLHV DUH IRUPHG E\ PDQ\ Nostoc VSS JURZLQJ LQ VLWX LQFOXGLQJ WKRVH WKDW JURZ VXEPHUJHG VXJJHVWLQJ IXQFWLRQV LQ DGGLWLRQ WR D UHWDUGDWLRQ RI ZDWHU ORVV 0DUWLQH] DQG 4XHULMHUR 'RGGV DQG &DVWHQKRO] 3ODWHD 3UHYLRXV VWXGLHV RI F\DQREDFWHULDO HQYHORSHV DQG VKHDWK VWUXFWXUHV VXJJHVWHG WKDW WKHVH PD\ VHUYH WR FRQFHQWUDWHPHWDOVDQGWKXVPD\ DLGLQH[FOXGLQJWKH FRORQLHV IURP SUHGDWLRQ E\ JDVWURSRGV LQVHFWV HWF 7HDVHDQG:DONHU $OWKRXJKPLFURRUJDQLVPV DUH FHUWDLQO\ SUHVHQW DW WKH VXUIDFHV RI WKH N. commune FRORQLHV WKH RXWHU VLOLFRQULFK OD\HU UHSUHVHQWV DQ LPSHQHWUDEOH EDUULHU IRU WKHP 7KH VLOLFRQULFK OD\HU PXVW EH PDGH WKURXJK SK\VLFRFKHPLFDOSUHFLSLWDWLRQDVLWLVKDUGWRDFFRXQW IRUDFRQFHUWHGV\QWKHVLVRIWKLVOD\HURQEHKDOIRIWKH FHOOV 0RUH OLNHO\ WKH OD\HU LV WKH SURGXFW RI VRPH R[\JHQGU\LQJGHSHQGHQW HIIHFW RQ WKH SHULSKHUDO VKHDWK DOWKRXJK LQ OLTXLG FXOWXUHV RI N. commune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
(KOLQJ6KXO] HW DO ,W UHPDLQV WR EH GHWHUPLQHG ZKHWKHU N. commune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
895$EVRUELQJ 3LJPHQWV $W OHDVW WZR GLVWLQFW 895DEVRUELQJ SLJPHQWV DUH SUHVHQWLQWKH(36 RIN. commune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
( ,QWHUDFWLRQV ZLWK2WKHU2UJDQLVPV Nostoc HQWHUVLQWRDUDQJHRIDVVRFLDWLRQVZLWKKLJKHU DQG ORZHU SODQWV DV ZHOO DV ZLWK WKH IXQJXV Geosiphon pyriforme &KDSWHU 6FKXEOHU HW DO 3ODWH ,QDGGLWLRQWKHUHDUHDQXPEHU RI LQWHUHVWLQJ REVHUYDWLRQV RQ LQGLUHFW LQWHUDFWLRQV EHWZHHQ Nostoc, IXQJLLQVHFWVSODQWVDQGEDFWHULD $ Nostoc VS LQDVVRFLDWLRQZLWK Oscillatoria VSS DQG Desulfovibrio VS IRUPV EODFN VXEVXUIDFH JOREXOHV DQG OD\HUV RIWHQ LQGLFDWLYH RIGHDWK RIWXUI RQ KLJK VDQG JROIJUHHQV +RGJHV D +RZHYHU WKH SUHVHQFH RI WKHVH F\DQREDFWHULD LQ EODFNOD\HUHG VDQGVGHFUHDVHGWKHURRWDQGVKRRWGU\ZHLJKWORVV RI WKH JUDVV Agrostis palustris ZKHQ LW ZDV LQIHFWHG E\ WKH IXQJDO SDWKRJHQ Pythium torulosum +RGJHV E 7KHLVRODWHGNostoc VS LPSHGHGZDWHUIORZ LQ DUWLILFLDO VDQG FROXPQV EXW GLG QRW OHDG WR EODFNHQLQJRIWKHVXEVWUDWH%ODFNHQLQJRFFXUUHGRQO\ ZKHQ WKH FROXPQV ZHUH LQRFXODWHG ZLWK Nostoc, RU WKH Oscillatoria VWUDLQV DQG FXOWXUHVRIDesulfovibrio VS 7KH XVH RID IUHVK LQRFXOXP RI F\DQREDFWHULD WKDW LQFOXGHG Nostoc muscorum GHFUHDVHG OHDI EODVW EXW QRW QHFN EODVW RI -DSRQLFD ULFH OLQH *L]D FDXVHG E\ Pyricularia oryzae
,,, (YROXWLRQ $ 7KH5LVHRI1RVWRF 2ULJLQ 7R FRQVLGHU WKH HYROXWLRQ RI Nostoc VSS WZR WLPH SHULRGV DUHRIUHOHYDQFH WKHSHULRG OHDGLQJXS WR WKH WLPHZKHQDQFHVWUDONostoc DFTXLUHGLWVFKDUDFWHULVWLF PRUSKRORJ\ DQG WKH SHULRG ZKLFK KDV HODSVHG VLQFH WKHQ
0DOFROP3RWWV /LIH PD\ KDYH HPHUJHG ELOOLRQ \HDUV DJR 0RM]VLV HW DO $Q DYDLODELOLW\ RI FRPELQHG QLWURJHQ LV QRW WKRXJKW WR KDYH SRVHG D VHULRXV OLPLWDWLRQ IRU WKH HYROXWLRQ RI HDUO\ OLIH EHFDXVH DWPRVSKHULFFKHPLVWU\LVVXFKWKDWOLJKWQLQJPD\KDYH SURYLGHGDVPXFKDV WRQV RIQLWURJHQSHU\HDUWR WKHSULPLWLYHRFHDQ
&KDSWHU 1RVWRF
Fig. 3. 5HFRUGVRIPLFURIRVVLOV WKRXJKWWREHRIF\DQREDFWHULDORULJLQ 5HFRUGV DUHWKRVHOLVWHGLQ6FKRSI 7D[RQRPLFJURXSV FRUUHVSRQGWRWKRVHLQWKHVFKHPHRI5LSSNDHWDO
,W ZDV FRQFOXGHG WKDW Nostoc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rpoB, SHUIRUPWKH VDPH³KRXVHNHHSLQJ´GXWLHVLQHYHU\WD[RQWKH\PD\
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ostoc VSS DQG N. commune LQ SDUWLFXODU DUH HVSHFLDOO\ ZHOO VXLWHG IRU WKH UHVROXWLRQ RI VRPH RI WKHVH SUREOHPV E\ YLUWXH RI WKHLU SUHYDOHQFH DV FRQVSLFXRXV PRUSKRORJLFDOO\VLPLODUFRPPXQLWLHV LQ
D EURDG UDQJH RI KDELWDWV IURP WKH 7URSLFV WR WKH SRODUUHJLRQV 'HVSLWH WKH FDXWLRQ HPSKDVL]HG E\ WKH SUHFHGLQJ GLVFXVVLRQ WKHUH LV RQH IHDWXUH RI WKH SK\ORJHQHWLF WUHH VKRZQ E\ 1HOLVVHQ HW DO WKDW EHDUV PHQWLRQ 7KHUH LV D KLQW RI WULFKRWRP\ ZLWKLQ WKH JURXSLQJ RIWKH WHQ KHWHURF\VWRXV IRUPV 2QH FOXVWHU LQFOXGHV Nostoc, Anabaena, Cylindrospermum DQG Nodularia VWUDLQV 2UGHU 1RVWRFDOHV sensu .RPiUHN DQG$QDJQRVWLGLV 6HFWLRQ,9 sensu 5LSSNDHW DO $ VHFRQG LQFOXGHV Fischerella DQG Chlorogloeopsis VWUDLQV 2UGHU6WLJRQHPDWDOHV sensu $QDJQRVWLGLV DQG .RPiUHN 6HFWLRQ 9 sensu 5LSSND HW DO DQG D WKLUG FRPSULVHV WKH VHTXHQFHV IURP Scytonema DQG Calothrix VWUDLQV 2UGHU1RVWRFDOHV sensu .RPiUHN DQG $QDJQRVWLGLV 6HFWLRQ,9 sensu 5LSSNDHWDO 'HYHORSLQJ DQ XQGHUVWDQGLQJ RI WKH SK\ORJHQ\ RI KHWHURF\VWRXVF\DQREDFWHULDUHTXLUHVPRUHH[KDXVWLYH PROHFXODU VXUYH\V RI QHZ LVRODWHV IURP ZHOO GRFXPHQWHG ILHOG ORFDWLRQV 5XGL HW DO IRFXVHG RQ WKH FRPSDUDWLYH VHTXHQFH DQDO\VLV RI '1$HQFRGLQJWKHYDULDEOHUHJLRQV99DQG9RI 6 U51$ LQ VWUDLQV RINostoc, Anabaena DQG Aphanizomenon. ,W ZDV FRQFOXGHG WKDW WKH VWUDLQ FDWHJRU\ WKH\ UHSUHVHQWHG ZDV PRQRSK\OHWLF EXW FRQVLVWHGRIWKUHHHYROXWLRQDU\GLIIHUHQWOLQHDJHVWZR Nostoc DQG RQHAnabaena/Aphanizomenon). 'HVSLWH WKH YHU\OLPLWHG GDWDVHWWKH KHWHURJHQHLW\ ZLWKLQ WKH Nostoc FDWHJRU\ SDUWLFXODUO\ WKH GLYHUJHQFH RI Nostoc VS 1,9$&<$ Anabaena VS 3&& IURP RWKHU QDPHG Nostoc VWUDLQV FROOHFWHG IURP ILHOG VLWHV LQ 1RUZD\ DQG $QWDUFWLFD ZDV REYLRXV ,QDVXEVHTXHQWVWXG\WKHDXWKRUVDGGHGWZR VWUDLQV N. commune DQG N. commune YDU flagelliforme ERWKLVRODWHGIURPILHOG VLWHVLQ&KLQD WRWKHGDWDVHW5XGLHWDO &RPSDULVRQRIWKH SK\ORJHQ\ EDVHG RQ 6 U'1$ DQG WKH WUDQVODWHG DPLQR DFLG VHTXHQFH RI rbcX UHYHDOHG VHYHUDO GLVFUHSDQFLHVEXWERWKKDGWKHVDPHFOXVWHULQJRIILYH LQGHSHQGHQW Nostoc LVRODWHV (Nostoc OLQHDJH ,, %RWK SK\ORJHQLHV HPSKDVL]HG WKH FOHDU GLYHUJHQFHV EHWZHHQ OLQHDJH ,, OLQHDJH , (Anabaena/ Aphanizomenon) DQG OLQHDJH,,,(Anabaena VS3&&
7HPSRRI6SHFLDWLRQ :KDW RI WKH WLPH VLQFH Nostoc DFTXLUHG LWV PRUSKRORJ\" 6WXOS GHVFULEHG WKUHH SRVVLEOH VFHQDULRVWRDFFRXQWIRUWKHDQWLTXLW\RIF\DQREDFWHULD DQG WKHLU KLJK GHJUHH RI JHQHWLF UHODWHGQHVV (LWKHU
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³VWDVLV´ VHH *RXOG DQG (OGUHGJH 6FKRSI SURSRVHG WKDW WKH PRUSKRORJ\ DQG SK\VLRORJ\ RI F\DQREDFWHULD HYROYHG OLWWOH RU QRW DW DOO RYHU WKRXVDQGV RI PLOOLRQV RI \HDUV 6FKRSI +\SHUEUDG\WHO\ UHIHUV WR D VORZ RU VXVSHQGHG WHPSR RIHYROXWLRQ 7KH SURSRVDO 6HFWLRQ ,,,$, WKDWWKH ODVW FRPPRQ DQFHVWRU RISURNDU\RWHV DQG HXNDU\RWHV RFFXUUHG QR HDUOLHU WKDQ ELOOLRQ \HDUV DJR FRQWUDGLFWV VXFK K\SHUEUDG\WHO\ :DV WKH JHQHWLF FRPSOH[LW\ DQG SK\VLRORJLFDO FDSDFLW\ RI DQFLHQW Nostoc VLPLODU LI QRW HTXLYDOHQW WR WKDW RI H[WDQW IRUPV VXFK DV N. commune RU KDYH H[WDQW IRUPV UHWDLQHG WKHLU PRUSKRORJLFDO DSSHDUDQFH GHVSLWH D KLJKO\ HYROYHG JHQRPH" 5XGL HW DO LQ GHIHQVH RI K\SHUEUDG\WHO\ VXJJHVWHG WKDW RQO\ WKH KLJKO\ FRQVHUYHG JHQHV XVHG LQ WKH HVWLPDWLRQV RI 'RROLWWOHDQG FROOHDJXHVFRXOG VKDUHDSSUR[LPDWHO\ ELOOLRQ \HDU ROG FRPPRQ DQFHVWRUV 7KH\ SURSRVHG WKDW WKH GLYHUJHQFH EHWZHHQ GLIIHUHQW OLQHDJHV RI KHWHURF\VWRXV F\DQREDFWHULD VXFK DV IRU WKH rbc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
&KDSWHU 1RVWRF $UFKDHD %DFWHULD DQG (XNDU\D 7KH RWKHU WZR SRVVLELOLWLHVZDUUDQWIXUWKHUFRPPHQW 7KH UHOHYDQFH RIUHFRPELQDWLRQ LQ WKH HYROXWLRQ RI EDFWHULDLQJHQHUDO DQG LQ F\DQREDFWHULDLQSDUWLFXODU DQGLWVUROH LIDQ\ LQ SRSXODWLRQV RIWKH ODWWHU ZKHQ WKH\ JURZ LQ VLWX LV H[FHHGLQJO\ KDUG WR HYDOXDWH 5HFRPELQDWLRQGRHV RFFXULQ VWUDLQV RIF\DQREDFWHULD PDQLSXODWHG XQGHU ODERUDWRU\ FRQGLWLRQV F\DQREDFWHULD GR FRQWDLQ rec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³SHULRGLFVHOHFWLRQ´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ostoc IRUPV 7KH ODWWHU SURFHVV FRQWULEXWHG WR WKH KRPRJHQL]DWLRQ LQ QHXWUDOO\HYROYLQJ JHQHV ZKRVH
DQDO\VLV LQ H[WDQW IRUPV PD\ EH SUREOHPDWLF ZKHQ DWWHPSWLQJWRGHWHUPLQHGLYHUJHQFHWLPHV :LWK UHVSHFW WR PRUSKRORJLFDOO\FRPSOH[ F\DQREDFWHULDVXFKDVNostoc, RQHPXVWTXHVWLRQKRZ PRUSKRORJLFDO FKDUDFWHUV ZKLFK PD\ KDYH DULVHQ HDUO\ HJ VKHDWK IRUPDWLRQ DQG FRORQ\ VWUXFWXUH HYROYHG LQ UHVSRQVH WR HQYLURQPHQWDO VHOHFWLRQ SUHVVXUH LI LQGHHGWKH\GLG
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¶VJHQRPHZKLFK PLUURU LWV HQYLURQPHQW ,Q WKH FDVH RI Nostoc VSS DQG RWKHU WHUUHVWULDO RU DHURSK\WLF IRUPV ZKRVH JHQRPHV DUH UHJXODUO\ GHVLFFDWHG WKHUH PD\ EH FKDUDFWHULVWLF JHQRPH ³VLJQDWXUHV´ .DUOLQ HW DO FRPSDUHG FRPSRVLWLRQDO ELDVHV GLVWULEXWLRQV RI VKRUW ROLJRQXFOHRWLGHV DQG GLQXFOHRWLGH UHODWLYH DEXQGDQFH GLIIHUHQFHV LQ SURNDU\RWHV IRU ZKLFK VXEVWDQWLDO LQ PDQ\ FDVHV FRPSOHWH JHQRPH VHTXHQFH GDWD ZHUH DYDLODEOH 7KHVH DQDO\VHV GHPRQVWUDWHG WKDW WKH F\DQREDFWHULD Synechocystis 3&& Synechococcus VS DQGAnabaena VS GLG QRWIRUPDFRKHUHQWJURXS DQG ZHUH DV IDU IURPHDFK RWKHU DV JHQHUDO JUDPQHJDWLYH VHTXHQFHV DUH IURP JHQHUDO JUDPSRVLWLYH VHTXHQFHV .DUOLQ HW DO ,W LV QRW XQUHDVRQDEOH WR VXJJHVW WKDW WKH VHTXHQFLQJ RI WKH FRPSOHWH JHQRPH RI N. commune FRXOGUHYHDOPXFKDERXWWKHFDSDFLW\RIWKLVRUJDQLVP IRUGHVLFFDWLRQWROHUDQFH 7KH DSSHDUDQFH RI WKH FKURPRVRPDO '1$ RI Nostoc commune FROOHFWHG IURP QDWXUDO SRSXODWLRQV LV YDULDEOH DQG GHSHQGV LQ ODUJHSDUWXSRQ WKH GHJUHH RI K\GUDWLRQ RI WKH QXFOHRSODVP )LJ 7KH QXFOHRLG RIAnabaena VS 3&& KDGDVWUXFWXUH WKDW LQFOXGHG SURWUXVLRQV OREHV DQG GLVWLQFW LVRODWHG HOHPHQWV 3LQHYLFK DQG *ULJRU\HYD 7KH ILJXUHV SURYLGHG E\ WKHVH DXWKRUV DUH VRPHZKDW UHPLQLVFHQW RI WKH FKDUDFWHULVWLF ³GRXJKQXW´VKDSHG QXFOHRLGV SUHVHQW LQ Dinococcus radiodurans - %DWWLVWDSHUVFRPP
Fig. 4. $SSHDUDQFH RI WKH QXFOHRSODVP LQ N. commune LV LQIOXHQFHGE\GHJUHHRIK\GUDWLRQ(OHFWURQPLFURJUDSKVRIFHOOVRI ILHOG PDWHULDOV RIN. commune: D 'HVLFFDWHG IRU \HDUV W WK\ODNRLG E 5HK\GUDWHG K SS SRO\SKRVSKDWH Q QXFOHRSODVP DUURZV GHQRWH PHPEUDQHERXQG ULERVRPHV F 5HK\GUDWHG GD\V DUURZV GHQRWH DW\SLFDO GLVFRQWLQXLW\ VSKHULFDO LQ WKH QXFOHRSODVP SRVVLEO\ SRO\SKRVSKDWH" 0DJQLILFDWLRQDSSUR[[ &RXUWHV\RI$ODQ3HDW
0DOFROP3RWWV 7KURXJK DQDORJ\ ZLWK WKH JHQRPH RIA. variabilis LW LV DVVXPHG WKDW WKH JHQRPHV RI Nostoc VSS DUH FLUFXODU +HUGPDQ 7KH JHQRPH VL]HV RI Nostoc VWUDLQV GHSRVLWHG LQ WKH 3DVWHXU &XOWXUH &ROOHFWLRQZHUHPHDVXUHG DV WR [ GDOWRQ RU [ ES +HUGPDQ HW DO 5DSLGO\ UHQDWXULQJ IUDFWLRQV LQ Nostoc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ostoc VSS WKLV UHVSRQVH PD\ EH RI SK\VLRORJLFDO DQG VWUXFWXUDO VLJQLILFDQFH ,Q WKLV UHJDUGLW ZRXOGEHRILQWHUHVWWRGHWHUPLQHWKHHIIHFWV RIFHOO GHK\GUDWLRQ LQ Anabaena VWUDLQV $% DQG &3%WKDWKDYHORVWWKHFDSDFLW\WRV\QWKHVL]HWKH KLVWRQHOLNH SURWHLQ +8 D SURWHLQ WKRXJKW WR SOD\ D UROHLQVSHFLILFUHFRPELQDWLRQHYHQWV.KXG\DNRYDQG :RON ,PSRUWDQWO\ E. coli hupAB PXWDQWV ZKLFK ODFNHG +8 ZHUH VHQVLWLYH WR 89 UDGLDWLRQ EHFDXVH RI WKH UROH RI +8 LQ KRPRORJRXV UHFRPELQDWLRQ/LDQG:DWHUV Nostoc JHQRPHV OLNH WKRVH RIRWKHU F\DQREDFWHULD DUH RIWHQ KHDYLO\ PHWK\ODWHG -DJHU DQG 3RWWV )LJ 6XFKPRGLILFDWLRQ LV NQRZQWRLQIOXHQFH WKH VHQVLWLYLW\ RI FHOOV WR : LUUDGLDWLRQ DQG SKRWRSURGXFWV VXFK DV +2
&KDSWHU 1RVWRF
Fig 5. *HQRPLF'1$RIN. commune FRQVLVWVRIK\SR , DQGK\SHUPHWK\ODWHG,, IUDFWLRQV-DJHUDQG3RWWV 'HVLFFDWHGFRORQLHV ZHUHUHK\GUDWHGIRUHLWKHUPLQK KRU GD\VEHIRUHSXULILFDWLRQRI'1$IUDFWLRQV)UDFWLRQVZHUHWUHDWHGZLWKHLWKHUDpnI ' RU MboI 0 )RUHDFKVHWRIUHK\GUDWLRQVVDPSOHVZHUHORDGHGDVIROORZV )UDFWLRQ , DpnI, MboI; )UDFWLRQ,, DpnI, MboI. /DPEGD+LQG,,, PDUNHUVLQODQHVDQGRIHDFKJHO&RXUWHV\.DULQ-DJHU
0DOFROP3RWWV
Fig. 6. 'LVWLQFW VL]HELDV DQG SHULRGLFLW\ LQ WKH VL]HV RIF\DQREDFWHULDOSRO\SHSWLGHV $OOWKHSRWHQWLDO SRO\SHSWLGHV HQFRGHG E\WKH JHQRPH RISynechocystis 3&& ZHUH LQFOXGHG LQ WKH FDOFXODWLRQ .DQHNR HW DO $UURZV LQGLFDWH PXOWLSOHV Q RIWKH EDVLF UHSHDWLQJXQLWRIN'D
$V WKH JHQRPLF VHTXHQFHV RI Nostoc VSS EHFRPH DYDLODEOH LQ WKH IXWXUH LW ZLOO EH SRVVLEOH WR DVVHVV WKH YDOLGLW\ RIWKH ODWWHUWKHRU\ DV ZHOO DV WKH SRVVLEOH UHOHYDQFH RI ODWHUDO WUDQVIHU WKURXJK FRPSDULVRQV RIWKH PRO *& FRQWHQWV RIVHOHFWHG JHQHV 6RPH SXWDWLYH 25)V ZHUH LGHQWLILHG IROORZLQJ'1$VHTXHQFHDQDO\VLVRIWKHFKURPRVRPH RI Synechocystis VS3&&.DQHNRHWDO 7KH FKURPRVRPHV RINostoc VSS DUHDSSUR[LPDWHO\ WZR WR WKUHH WLPHV ODUJHU WKDQ WKDW RI Synechocystis VS 3&& 7KHVH GDWD PD\ VXJJHVW WKDW WKH Nostoc JHQRPH FRXOG HQFRGH DV PDQ\ DV SRO\SHSWLGHV *HQHV DVVRFLDWHG VSHFLILFDOO\ ZLWK WKH GLIIHUHQWLDWLRQ RI KHWHURF\VWV UHSUHVHQW RQH FODVV RI JHQHV WKDW GR QRW DSSHDU WR EH SUHVHQW LQ QRQ KHWHURF\VWRXVIRUPVVXFKDVSynechocystis DUHWKHUH
RWKHUV" Nostoc VSS DUH FDSDEOH RI D QXPEHU RI GHYHORSPHQWDO RSWLRQV DQG SHUKDSV DOO RI WKHVH UHTXLUH QRYHO JHQHV &RKHQ HW DO &HUWDLQO\ RQHWKDWEHDUVFRQVLGHUDWLRQLVWKHFDSDFLW\IRUNostoc VSS WR LQYDGH ORZHU DQG KLJKHU SODQWV DQG WR HVWDEOLVKVWDEOHUHODWLRQVKLSV ZLWK WKHP&KDSWHU 2WKHUZLVH GR Nostoc JHQRPHV VLPSO\ FDUU\ D FRQVLGHUDEOHIUDFWLRQRIUHGXQGDQW'1$"$WWKLV WLPH RIZULWLQJ KRZHYHU WKH VHTXHQFHV RI RQO\ RU VR Nostoc VSS SURWHLQV ZHUH GHSRVLWHG LQ GDWD EDQNV DQG VRPH RI WKHVH UHIOHFW WKH GXDO QDPLQJ RI Anabaena VS3&&DVNostoc VS 3&& :KLOH WKHUH DUH QR VSHFLILF GDWD DYDLODEOH DW WKLV WLPHWRDVVHVVWKHQXPEHUVRISURWHLQVHQFRGHGE\WKH JHQRPHVRINostoc VSSLWLVXVHIXOWRFRQVLGHUFHUWDLQ GDWDLQ WKH FRQWH[W RIWKH HYROXWLRQ RIF\DQREDFWHULDO JHQRPHV 6RPH RIWKH VDPSOHGSURWHLQV LQ
&KDSWHU 1RVWRF E. coli KDYH PROHFXODU PDVVHV RIQ [ GDOWRQ Q HWF 6DYDJHDX 7KLV ZDV LQWHUSUHWHGDVWKHFRQVHTXHQFHRISRO\SHSWLGHVDULVLQJ IURP D IXQGDPHQWDO VWUXFWXUDO XQLW :LWK WKH DYDLODELOLW\ RI WKH FRPSOHWH '1$ VHTXHQFH RI Synechocystis VS 3&& .DQHNRHWDO LW ZDV SRVVLEOH WR DQDO\]H DOO RI WKH SRWHQWLDO SRO\SHSWLGHVHTXHQFHV RIWKLV F\DQREDFWHULXP 7KH SRO\SHSWLGHSURGXFWVLGHQWLILHGLQSynechocystis VS 3&& DSSHDU WR EH GHVLJQHG RQ D VWDQGDUG XQLW RI N'D HVVHQWLDOO\ DQ LGHQWLFDO VL]H WR WKDW LGHQWLILHGLQE. coli, ZLWK REYLRXVSHDNVRIQ Q )LJ :LOO WKLV SURYH WR EH WKH FDVH IRU RWKHU F\DQREDFWHULD" 'LG LQFUHPHQWDO FKDQJHV WR WKH EDVLF XQLW IDFLOLWDWH PLFURHYROXWLRQ DQG GLG TXDQWXP FKDQJHV WR WKH QH[W VWDEOH VL]H RI SURWHLQ DFFRPPRGDWHPDFURHYROXWLRQ"
1XFOHLF$FLG6WDELOLW\8QGHU6WUHVV &RXOG DORZUDWH RIHYROXWLRQDU\ FKDQJH EH WKH SULFH SDLG RU UDWKHU WKH FRQVHTXHQFH IRU EHLQJ DEOH WR FRORQL]H H[WUHPH HQYLURQPHQWV ZKHUH '1$ GDPDJH PD\ EH DFFHOHUDWHG" 895 IUHH UDGLFDO JHQHUDWLRQ SKRWR[LGDWLRQ DQG GHVLFFDWLRQ DOO OHDG WR '1$ GDPDJHDQGSUHVXPDEO\WKH\ LPSRVHG DFRQVLGHUDEOH UHVWUDLQWXSRQWKHFRORQL]DWLRQRIKDELWDWVE\HDUO\OLIH IRUPV LQFOXGLQJ WHUUHVWULDO Nostoc VSS 3RWWV $FRQVLGHUDWLRQRIWKHFRQVHTXHQFHV RI'1$GDPDJH PD\ VKHG OLJKW XSRQ VHOHFWLRQ SUHVVXUHV LPSRVHG XSRQ SRSXODWLRQV RIF\DQREDFWHULD 2QHSURFHVV WKDW FDXVHV FRQVLGHUDEOH GDPDJH WR FHOOXODU '1$ LV H[SRVXUH WR LRQL]LQJ UDGLDWLRQ &R LUUDGLDWLRQ LQWURGXFHV GDPDJH WR '1$ SULQFLSDOO\ WKURXJK WKH LQWURGXFWLRQ RI GRXEOHVWUDQGHG EUHDNV '6%¶V &\DQREDFWHULD DUH NQRZQ WR EH KLJKO\ UHVLVWDQW WR &R DQG &RLUUDGLDWLRQKDV VRPHWLPHVEHHQXVHGWR UHQGHU FXOWXUHV RI F\DQREDFWHULD D[HQLF LQ KLV ODERUDWRU\ 7KH SUDFWLFH ZDV QRW ZLGHVSUHDG SUHVXPDEO\ GXH WR FRQFHUQV DERXW HQKDQFHPHQW RI PXWDWLRQ IUHTXHQFLHV :HUH WKRVH IHDUV IRXQGHG " 7KH IROORZLQJ VXJJHVWV SHUKDSV WKH\ ZHUH QRW 6SHFLHV RI WKH EDFWHULDO JHQXV Deinococcus, HVSHFLDOO\ D. radioduruns, KDYH D VWDUWOLQJ UHVLVWDQFHWRLRQL]LQJDQGXOWUDYLROHWUDGLDWLRQDVZHOO DV PXWDJHQV WKDW IRUP DGGXFWV DQG FURVV OLQNV LQ '1$ 7KH PHDQV E\ ZKLFK WKH\ GR VR KDV EHHQ VWXGLHG LQ GHWDLO $JRVWLQL HW DO DQG D FRQVLGHUDWLRQ RI WKH GDWD LV LPSRUWDQW WR DOORZ VSHFXODWLRQ DERXW FHUWDLQ SK\VLRORJLFDO UHVSRQVHV RI Nostoc VSS )RU H[DPSOH IROORZLQJ H[SRVXUH WR 0UDG*\ LWZDVGHWHUPLQHGWKDWWKHJHQRPH
RI D. radiodurans VXVWDLQVDERXW '6%VZKLFKDUH UHSDLUHGZLWKLQKZLWKRXWOHWKDOLW\PXWDJHQHVLVRU UHDUUDQJHPHQWV 'DO\ DQG 0LQWRQ - %DWWLVWD SHUV FRPP )ROORZLQJ DQ H[SRVXUH WR 0UDG ZKLFK GHJUDGHV WKH IRXU FKURPRVRPHV RI D FHOO LQWR DERXW IUDJPHQWV DV PDQ\ DV FURVVRYHUV SHU FKURPRVRPH FURVVRYHUV SHU QXFOHRLG XQGHUZHQW UHSDLU 'DO\ DQG 0LQWRQ D E $ PRGHO WR H[SODLQ KRZ D. rudiodurans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radioduruns WR OHYHOV RI LRQL]LQJ UDGLDWLRQ WKDW GR QRW DQG KDYH QHYHU H[LVWHG RQ (DUWK LV VLPSO\ D
Fig. 0RGHO RI KRZ WKH Deinococcus JHQRPH LV UHSDLUHG IROORZLQJ GDPDJH IURP LRQL]LQJ UDGLDWLRQ RU GHVLFFDWLRQ +\SRWKHWLFDO GRXEOHFKURPRVRPHVWUXFWXUHVKRZLQJ GRXEOHVWUDQG '1$ EUHDNV DW ORFL $% DQG &' KHOG LQ DOLJQPHQW WKURXJK SHUVLVWHQW +ROLGD\MXQFWLRQV 'DO\ DQG 0LQWRQ &RXUWHV\ RIWKHDXWKRUVZLWKSHUPLVVLRQRIWKHSXEOLVKHU
UHIOHFWLRQ RIWKHFDSDFLW\ RIWKH VWUDLQIRUGHVLFFDWLRQ WROHUDQFH 0DWWLPRUH DQG %DWWLVWD 7KH FDSDFLW\ RI N. commune IRU GHVLFFDWLRQ WROHUDQFH PXVWUHIOHFW LQ SDUW LWV FDSDFLW\ WR SHUIRUPHIILFLHQW '1$ UHSDLU LQ WKLV UHJDUG PRUH LQIRUPDWLRQ LV QHHGHG RQ WKH UROH RIUHGXQGDQW '1$ VHTXHQFHV LQ F\DQREDFWHULD VXFK DV PXOWLSOH JHQRPHV 6755 V +,3 VHTXHQFHV &KDSWHU DQG WKH UROH RI FKURPRVRPHVWUXFWXUHDQG'1$PHWK\ODWLRQLQWKHVH RUJDQLVPV
*HQHVIRU&DUERK\GUDWH0HWDEROLVP 3DWKZD\V RI FDUERK\GUDWH PHWDEROLVP DQG WKHLU FRQVWLWXHQWHQ]\PHV DUH WKRXJKW WR KDYH DULVHQ HDUO\ GXULQJ HYROXWLRQ 0DUVK DQG /HEKHU] $ FRQVLGHUDWLRQ RI WKH JHQHV HQFRGLQJ FDUERK\GUDWH PRGLI\LQJHQ]\PHVLQNostoc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ostoc muscorum, Anabaena variabilis, Plectonema VS DQGAnabaenopsis VS DQGWKHDFWLYLW\ZDV IRXQG WR EH XVHG VWULFWO\ IRU KH[RVH V\QWKHVLV GXULQJ WKH OLJKWLQGHSHQGHQW UHDFWLRQV RISKRWRV\QWKHVLV 5XWWHU :LOODUG DQG *LEEV 7KH RUJDQL]DWLRQ RI WKHIGDUHJLRQLQN. commune LVDW\SLFDO-RDUGDU HW DO XQSXEOLVKHG GDWD ,Q WKH %DFWHULD JHQHV HQFRGLQJ &DOYLQ &\FOH HQ]\PHV DUH VRPHWLPHV GXSOLFDWHGWKH\DUHRIWHQFOXVWHUHGOLQNHGWLJKWO\DQG RULHQWDWHG LQ WKH VDPH GLUHFWLRQ RQ WKH FKURPRVRPH $OHIRXQGHU HW DO &KHQ HW DO *LEVRQ DQG7DELWD *LEVRQHWDO 6FKDIHUMRKDQQ HWDO 6LQJHUHWDO YDQGHQ%HUJKHWDO 5HFHQWO\LWZDVSURSRVHGWKDWWKHUHPD\EHD
0DOFROP3RWWV GLFKRWRP\ LQ WKH HYROXWLRQ RI WKH EDFWHULDO FODVV ,, DOGRODVHV 6FKlIHUMRKDQQ HW DO 2XU DQDO\VLV DQG FRPSDULVRQV RI)GD VHTXHQFHV LQFOXGLQJ WKDW RI N. commune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nabaena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ucA, HQFRGLQJ VXFURVH V\QWKDVH DQG WKRXJKW WR EH LQYROYHG LQ VXFURVH EUHDNGRZQ OHDG WR D )L[ SKHQRW\SH LQ Anabaena VS 3&& %XLNHPD HW DO ,W VHHPV XQOLNHO\ KRZHYHU WKDW IRUPV VXFK DV Nostoc DFTXLUHGWKHDELOLW\WRPDNHVXFURVHVROHO\WR VXSSRUW QLWURJHQIL[DWLRQ ,QIDFWDJHQHFRUUHVSRQGLQJWRWKDW ZKLFKHQFRGHVWKHVXFURVHSKRVSKDWH V\QWKDVHRIZea mays ZDVLGHQWLILHGLQSynechocystis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
&KDSWHU 1RVWRF GHVLFFDWHG DQG UHK\GUDWHG FHOOV RI N. commune &+(1 (Tmdry & DQG Tmwet &UHVSHFWLYHO\+LOO HWDO 7KHUHLVUHDVRQWRVXSSRVHWKDWVXFURVH V\QWKHVLV LQ F\DQREDFWHULD DURVH WKURXJK WKH QHHG WR SURWHFW FHOOV GXULQJ GHVLFFDWLRQ SUREDEO\ WKH SULQFLSOH VWUHVV LPSRVHG XSRQ DQFLHQW WHUUHVWULDO DQG DHURSK\WLFIRUPV
'1$ 7UDQVIHU .QRZOHGJH RI WKH JHQHWLFV RI KHWHURF\VWRXV F\DQREDFWHULD ZDV PDGH SRVVLEOH LQ ODUJH SDUW WKURXJK WKH GHYHORSPHQW RID FRQMXJDWLRQ V\VWHP IRU ILODPHQWRXV F\DQREDFWHULD DQG LWV DSSOLFDWLRQ WR Anabaena VS 3&& :RON HW DO +DVHONRUQ 7KH GHYHORSPHQWRIPHWKRGV XVHG WR PDQLSXODWHWKLV VWUDLQ ZHUHPDGHSRVVLEOH WKURXJK LVRODWLRQ RI WKH UHSOLFDWLRQ RULJLQ RI D SODVPLG WKDW ZDV HQGLJHQRXV WR Nostoc VS VWUDLQ 3&& 6FKPHWWHUHUDQG:RON $Q DX[RWURSKLF YDOLQHUHTXLULQJ VWUDLQ RI N. muscorum ZDV UHSRUWHGWR EHWUDQVIRUPHGZLWK'1$ LQ DSURFHVV WKDW ZDV LQVHQVLWLYH WR ULERQXFOHDVH DQG SURWHDVH DQG GHSHQGHQW XSRQ '1$ FRQFHQWUDWLRQ 7UHKDQ DQG 6LQKD (QYLURQPHQWDO IDFWRUV VXFK DV 89 LUUDGLDWLRQ DQG GHVLFFDWLRQ PD\ KDYH GLUHFW HIIHFWV XSRQ WKH JHQRPLF LQWHJULW\ RI Nostoc VSSDQGRWKHUF\DQREDFWHULD&KDSWHU JURZLQJLQ VLWX EXW WKH HIIHFWV RI PRUH LQGLUHFW IDFWRUV XQGHU WKHVHFRQGLWLRQV VXFKDVF\DQRSKDJHVDQGH[RJHQRXV '1$ UHPDLQODUJHO\XQNQRZQ&KDSWHU
& $TXLVLWLRQ RIWKH+HWHURF\VW 6HOHFWLYH3UHVVXUH 6WUXFWXUDO SK\VLRORJLFDO DQG PROHFXODU SURSHUWLHV RI KHWHURF\VWV DUH XQGHUVWRRG LQ GHWDLO :RON HW DO EXW ZKDW VHOHFWLYH DGYDQWDJH ZDV LPSRVHG XSRQ Nostoc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ostoc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nabaena variabilis $7&& 7KLHO HW DO 6FKUDXWHPHLHU HW DO 7KH 1LI V\VWHP IXQFWLRQVERWKLQKHWHURF\VWV DQG YHJHWDWLYH FHOOV EXW RQO\ XQGHU DQDHURELF FRQGLWLRQV 7KH 1LI V\VWHP ZDV DEVHQW IURP D QXPEHU RI VWUDLQV FODVVLILHG DV Anabaena VSS LQFOXGLQJ Anabaena VS 3&& DQG DOVR DEVHQW IURP D QXPEHU RI Nostoc VWUDLQV LQFOXGLQJ IUHHOLYLQJ LVRODWHV IURP Azolla DQG Mucrozamia. 7KH V\VWHP ZDV GHWHFWHG LQ ILYH LQGHSHQGHQW LVRODWHV FODVVLILHG DV Anabaena IURP Azolla. 7KHVHGDWDLPSO\WKDWWKHSUHVHQFHRIWKH1LI V\VWHP UHIOHFWV D UHVWULFWHG SK\VLRORJLFDO FDSDFLW\ D PRUH WKRURXJK DQDO\VLV RI LWV GLVWULEXWLRQ LV QHHGHG EHIRUHLWFDQEHGHWHUPLQHGZKHWKHUSRVVHVVLRQRIWKLV V\VWHP LV D XVHIXO WD[RQRPLF PDUNHU ZLWKLQ WKH Anabaena/Nostoc JURXS 7KLVILQGLQJRIDVHFRQG 1LI V\VWHP LQ YHJHWDWLYH FHOOV RI VRPH KHWHURF\VWRXV IRUPLQJ VWUDLQV UDLVHG WKH TXHVWLRQ RI ZKHWKHU 1LIO DQG1LIDURVHLQGHSHQGHQWO\DTXHVWLRQZKLFKLVQRW \HWUHVROYHGZLWKFHUWDLQW\ 7KHUHVWULFWHGGLVWULEXWLRQ RI WKH 1LI V\VWHP PD\ VXJJHVW D ODWHUDO WUDQVIHU HYHQW
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ostoc VS 3&& DQG A. cylindrica ZHUH IRXQG WR EH PRUH UHVLVWDQW WR GU\LQJ DQG IUHH]LQJ WKDQYHJHWDWLYHFHOOV VHH+HUGPDQQ ,Q IDFW HOHFWURQ PLFURVFRSH DXWRUDGLRJUDSK\ XVLQJ +WK\PLGLQH LQGLFDWHG WKDW DNLQHWHV ZHUH QRW TXLHVFHQW DQG KDG D PRGH RI '1$ V\QWKHVLV WKDW PDWFKHG WKDW LQ JHUPLQDWLQJ DNLQHWHV DQG YHJHWDWLYH FHOOV )DYDOL DQG &DLROD *HLWOHU FRPPHQWHGWKDWKH³VSRWWHG´DNLQHWHVLQN. commune RQ RQO\ RQH RFFDVLRQ DQG TXHVWLRQHG ZKHWKHU RWKHU UHSRUWV RIDNLQHWHV LQ WKLV F\DQREDFWHULXP FRQFHUQHG RWKHU IRUPV ,Q IDFW KH DWWULEXWHG WKH UDUH GHYHORSPHQW RI DNLQHWHV LQ N. commune WR WKH UHVLVWDQFH RIWKH FRORQ\ LH WKH YHJHWDWLYH FHOOV WR GHVLFFDWLRQ DQG IUHH]LQJ %\ GRLQJ VR KH LPSOLHG D IXQFWLRQIRU DNLQHWHV +RZHYHU LWGRHVQRWVHHPWKDW WKH IXQFWLRQRIDNLQHWHVLVVSHFLILFDOO\IRUGHVLFFDWLRQ WROHUDQFH RU IUHH]LQJ WROHUDQFH 3RWWV $NLQHWHVZHUHDEVHQWIURPDOORIWKHILHOGPDWHULDOVRI N. commune ,FROOHFWHGGXULQJWKHSDVW \HDUV DQG WKH\ GR QRW RFFXU LQ ODERUDWRU\ FXOWXUHV RI N. commune '5+O RU 87(; XQSXEOLVKHG REVHUYDWLRQV 7KH YHJHWDWLYH FHOOV RI N. commune '5+O GR VKRZ HOHFWURQ WUDQVOXFHQW ZDOO OD\HUV WKDW UHVHPEOH WKRVH RI DNLQHWHV 3RWWV DQG RWKHU Nostoc VWUDLQV GR SURGXFH DNLQHWHV 0ROOHQKDXHU E $NLQHWHV DUH UHVWLQJ VWDJHV WKDW PD\ FRQIHU VRPH VHOHFWLYH DGYDQWDJH %XW QRQKHWHURF\VWRXV F\DQREDFWHULD DUH DV HTXDOO\ GLVWULEXWHG DQG VHHPLQJO\ DV VXFFHVVIXO DV KHWHURF\VWRXV F\DQREDFWHULD \HW WKH\ KDG QR UHFRXUVH WR GHYHORS DNLQHWHV:K\"
0DOFROP3RWWV *HQH ([SUHVVLRQ LQ WKH 'LIIHUHQWLDWHG +HWHURF\VW 1HDUO\ RI WKH Anabaena variabilis $7&& '1$ VHQVH VWUDQG GLIIHUHQW P51$ VSHFLHV LV WUDQVFULEHG E\ WHUPLQDOO\GLIIHUHQWLDWHG KHWHURF\VWV DQG PRUH WKDQ KDOI RI WKH WUDQVFULEHG P51$V DSSUR[LPDWHO\ JHQHV DUH KHWHURF\VW VSHFLILF/\QQHW DO %XLNHPDDQG +DVHONRUQ 7KH DELOLW\ WR GLIIHUHQWLDWH D KHWHURF\VW WKXV UHTXLUHV DQ DPRXQW RI '1$ WKDW LV JUHDWHU WKDQ WKH WRWDO JHQRPH VL]H RI D VPDOO SURNDU\RWH HJ Mycoplasma genitalium NES ,Q YLHZ RI WKH SURSRVHG LQRUGLQDWHO\ VORZ UDWH RI HYROXWLRQ RI F\DQREDFWHULD &KDSWHU WKH FRPSDUDWLYHO\ UHFHQW DSSHDUDQFH RIIRUPV FDSDEOH RIGLYHUWLQJ VRPH RI WKH FRGLQJ FDSDFLW\ RI WKHLU JHQRPH WR D VLQJOHSK\VLRORJLFDOIXQFWLRQLV LQFRQJUXRXV 'LGWKH FDSDFLW\ WR GLIIHUHQWLDWH D KHWHURF\VW HYROYH VORZO\ WKURXJK JUDGXDO UHDUUDQJHPHQWV DQG PRGLILFDWLRQV RI H[LVWLQJ JHQHV DQG SDWKZD\V RU GLG WKH HYHQW RFFXU UDSLGO\ IROORZLQJ DFTXLVLWLRQ RI '1$ E\ ODWHUDO WUDQVIHU" 7KH JHQRPH RI Anabaena VS 3&& LV DSSUR[LPDWHO\ 0E DQG ZDV SDUWLDOO\ PDSSHG XVLQJ KRPRORJRXV DQG KHWHURORJRXV JHQH SUREHV :RON 2QO\ D UHODWLYHO\ VPDOO QXPEHU RI JHQHV ZHUH PDSSHG EXW WKH SRVLWLRQV RI D VXIILFLHQW QXPEHU RI WKRVH UHTXLUHG IRU KHWHURF\VW GLIIHUHQWLDWLRQ ZHUH ORFDWHG )URP WKHVH DQDO\VHV LW DSSHDUV WKDW JHQHV HQFRGLQJ FRPSRQHQWV RI WKH KHWHURF\VW GLIIHUHQWLDWLRQ PDFKLQHU\ DUH VFDWWHUHG WKURXJKRXWWKHJHQRPH )RUH[DPSOHZLWKUHVSHFWWR SRVLWLRQ 2 RQ D FLUFXODU PDS hetR %XLNHPD DQG +DVHONRUQ LV ORFDWHG DW SRVLWLRQ 0E JHQHV RI WKH nifB DQG nifH RSHURQV 0XOOLJDQ DQG +DVHONRUQ DUHFOXVWHUHG DW0E DQGhepA LV ORFDWHG DW SRVLWLRQ 0E +ROODQG DQG :RON &DQZHH[SHFWDVLPLODUSDWWHUQRIGLVWULEXWLRQ RIKHWHURF\VWVSHFLILF JHQHV LQNostoc VSS DQG RWKHU KHWHURF\VWRXVIRUPV" 8QOLNHQRQKHWHURF\VWRXVIRUPVNostoc, Anabaena DQG RWKHU KHWHURF\VWRXV F\DQREDFWHULD FRQWDLQ PXOWLSOH FRSLHV RI JHQHV ZKLFK HQFRGH SURGXFWV VLPLODUWRHXNDU\RWLF51$ELQGLQJSURWHLQV0XOOLJDQ HWDO 5DWKHUWKDQ WKHSUHVHQFH RIKHWHURF\VWV WKLV PD\ UHIOHFW WKH ODUJHU JHQRPHV RIWKH ODWWHU DQG PRUHFRPSOH[SK\VLRORJ\VXFKDVWKHSUHVHQFHRI OLIH F\FOHV
&KDSWHU 1RVWRF 6LJQLILFDQFHLQ3ODQW6\PELRVHV :K\GR V\PELRVHV EHWZHHQSODQWV DQG F\DQREDFWHULD LQYROYH KHWHURF\VWRXV DQG QRW QRQKHWHURF\VWRXV IRUPV DQG ZK\ LV LW WKDW Nostoc VSS DUH WKH SUHGRPLQDQW KHWHURF\VWRXV IRUPV LQ VXFK DVVRFLDWLRQV" 6\PELRWLF F\DQREDFWHULD GR KDYH WKH FDSDFLW\WRIRUPPRWLOHKRUPRJRQLDDQGWKLVSURSHUW\ LVWKRXJKWWREHLPSRUWDQWIRUWKHHVWDEOLVKPHQWRIDQ DVVRFLDWLRQ ZLWK SODQWV 7DQGHDX GH 0DUVDF &KDSWHU ,Q IDFW VSHFLILF JHQHV LQ Nostoc punctiforme $7&& hrmUA, PD\ UHJXODWH KRUPRJRQLXP IRUPDWLRQ WKURXJK PHWDEROLVP RI D PHWDEROLWH IURP WKH V\PELRQW Anthoceros punctatus &RKHQ DQG 0HHNV +RZHYHU DOWKRXJK QXPHURXV F\DQREDFWHULD SURGXFH PRWLOH KRUPRJRQLD WKH DELOLW\ WR HQWHU LQWR V\PELRVHV ZLWK SODQWV LV UHVWULFWHG WR D IHZ JHQHUD ,QWHUHVWLQJ GDWD ZHUH SURYLGHG UHFHQWO\ E\ 0HHNV DQG FROOHDJXHV Nostoc VS VWUDLQ8&' LVDWUDQVSRVRQLQGXFHGPXWDQWRI Nostoc VS VWUDLQ $7&& WKDW LV XQDEOH WR IL[ QLWURJHQLQDLUEXWLWGRHVVRXQGHUDQR[LFFRQGLWLRQV 7KHPXWDQWSKHQRW\SHUHVXOWVIURPGLVUXSWLRQRIdevR WKDW LV HVVHQWLDO WR WKH GHYHORSPHQW RI PDWXUH KHWHURF\VWV EXW QRW DNLQHWHV &DPSEHOO HW DO 7KHUHIRUH WKH DELOLW\ RI WKH PXWDQW WR LQYDGH DQG HVWDEOLVK DQ DVVRFLDWLRQ ZLWK Anthoceros punctatus VXJJHVWVWKDWSURWHFWLRQDJDLQVWR[\JHQLQDFWLYDWLRQLV SURYLGHG ZLWKLQ WKH WLVVXHV RI WKH KRVW $ UDQJH RI RWKHU DQFLOODU\ IDFWRUV PXVW EH WDNHQ LQWR DFFRXQW ZLWK UHVSHFW WR R[\JHQ VHQVLWLYLW\ )RU H[DPSOH Nostoc cordubensis 3URVSHUL FDQ IRUP FXOWXUHVZLWK RU ZLWKRXWPXFLODJH PXFK RIZKLFK LV SURGXFHG E\ KHWHURF\VWV 3URVSHUL 0XFLODJLQRXV KHWHUR F\VWV UHWDLQHG QLWURJHQIL[LQJ FDSDELOLW\ DW KLJK R[\JHQ FRQFHQWUDWLRQV ZKLOH KHWHURF\VWV ODFNLQJ PXFLODJH ZHUH XQDEOH WR IL[ QLWURJHQ DW R[\JHQ FRQFHQWUDWLRQV KLJKHU WKDQ $W WKLV WLPH WKHUH DSSHDUV WR EH QR DGHTXDWH H[SODQDWLRQ IRU WKH SUHYDOHQFHRINostoc VSSLQSODQWDVVRFLDWLRQV
,9 3K\VLRORJ\DQG*URZWK $ /LIH&\FOHV 7KHOLIHF\FOHVRINostoc VSS UHSUHVHQWSK\VLRORJLFDO DQGSUHVXPDEO\JHQHWLF FDSDFLWLHVWKDWDUHXQNQRZQ LQ RWKHU JHQHUD .RPDUHN DQG $QDJQRVWLGLV 7KHVH FKDUDFWHULVWLFV LQFOXGH LQ VLPSOH WHUPV WKH IRUPDWLRQ RIPRWLOH KRUPRJRQLD ZLWK RU ZLWKRXW JDV YHVLFOHV WKH IRUPDWLRQ RI VHULDWH ILODPHQWRXV DQG DVHULDWH EDOOOLNH SKDVHV IRUPDWLRQ RI PLFUR DQG
PDFUR FRORQLHV UHSURGXFWLRQ WKURXJK EXGGLQJ DQG IUDJPHQWDWLRQ RI FRORQLHV RU GHYHORSPHQW RI KRUPRJRQLD 0ROOHQKDXHUE 0ROOHQKDXHU 'RGGV HW DO 7DQGHDX GH 0DUVDF /D]DURII 3RWWV DQG %RZPDQ 6R GLVWLQFW DUH WKHVH OLIH F\FOHV WKDW LW LV QRW VXUSULVLQJ WKH\ ILJXUHG SURPLQHQWO\ LQ FODVVLFDO GHVFULSWLRQV RI Nostoc DQG ZHUH WKXVXVHG H[WHQVLYHO\ LQWD[RQRPLF VFKHPHV *HLWOHU :KLOH WKH YDOLGLW\ RI WKH PRUH HODERUDWH RIWKH ODWWHU DUH TXHVWLRQDEOH WKHUH LV QR GRXEW WKDW WKH JHQHWLF GHWHUPLQDQWV RI WKHVH OLIH F\FOHV ZLOO EH IRXQG WR KDYH VROLG WD[RQRPLF XWLOLW\ IRUWKHGLVFULPLQDWLRQRIFHUWDLQNostoc VSS 6WXGLHV ZLWK Nostoc commune VWUDLQ 87(; Nostoc muscorum $ DQG Nostoc commune VXJJHVWHG WKDW WKH VSHFWUDO TXDOLW\ RI LQFLGHQW UDGLDWLRQ DV ZHOO DV WKH SUHVHQFH RI H[FUHWHG PHWDEROLWHV RI XQNQRZQ LGHQWLW\ LQIOXHQFHG PRUSKRORJLFDO FKDQJHV LQFOXGLQJ WKH SURGXFWLRQ RI KRUPRJRQLD,VRQRDQG)XMLWD /D]DURII 5RELQVRQ DQG 0LOOHU 0DUWLQH] HW DO 7DQGHDX GH 0DUVDF +RZHYHU PDQ\ GHWDLOV DUH ODFNLQJ DQG LQ VRPH FDVHV WKH GDWD RI GLIIHUHQW DXWKRUV DUH FRQWUDGLFWRU\ )XUWKHU DGYDQFHV LQ WKLV DUHD PD\ EHQHILW IURP WKH LQFUHDVLQJ DZDUHQHVV RI DQG LQWHUHVW LQ WKH FRPSRQHQWV RI SURWHLQ SKRVSKRU\ODWLRQ DQG GHSKRVSKRU\ODWLRQ QHWZRUNV LQ EDFWHULDO SK\VLRORJ\ DQG WKHLU UROHV LQ VHQVLQJ .HQQHOO\ DQG 3RWWV 0F&DUWQH\ HW DO )RU H[DPSOH F$03 VWLPXODWHG VLJQLILFDQWO\ WKH DJJUHJDWLRQ DQG JOLGLQJ PRYHPHQW RI ILODPHQWV RI Spirulina platensis, DQG UDSLG FKDQJHV LQ OHYHOV RI F$03 LQ Anabaena cylindrica DFFRPSDQLHG UHVSRQVHV WR HQYLURQPHQWDO IDFWRUV LQFOXGLQJ FKDQJHV LQS+ 2KPRUL 2KPRUL HWDO WKH JHQH HQFRGLQJ DGHQ\ODWH F\FODVH LQ F\D LQ Anabaena cylindrica LVQRZDYDLODEOH.DWD\DPDHWDO 7KH PRUSKRORJLFDO IHDWXUHV RI WKH OLIH F\FOHV RI VHYHUDO Nostoc VSS DUH GRFXPHQWHG LQ GHWDLO 0ROOHQKDXHU /D]DURII 0DUWLQH] HWDO DQGSKRWRJUDSKV RIVRPHUHSUHVHQWDWLYH VWDJHV ZHUHSURYLGHGE\5LSSNDHWDO 7KHIROORZLQJ GHVFULEHV RQO\WKHHVVHQWLDO IHDWXUHV RIWKHOLIH F\FOHV RI Nostoc commune VWUDLQV 87(; DQG '5+ 7KHVH VWUDLQV GLIIHU LQ DVSHFWV RIWKHLU OLIH F\FOH LQ PRUSKRORJ\ DQG LQ D QXPEHU RI GLIIHUHQW JHQHWLF PDUNHUV ,W LV XQFOHDU ZKHWKHU WKH RULJLQDO LVRODWH RI N. commune 87(; GHULYHG IURP FODVVLF ILHOG PDWHULDORIN. commune. Nostoc VS *LEVRQ 7 *LEVRQ 6FRWODQG" 0 '\DU 8 :DVKLQJWRQ 0% $OOHQ 8 &DOLIRUQLD%HUNHOH\ GHSRVLWHG LQ 87(; &XOWXUH
&ROOHFWLRQ DV N. commune 87(; 0 3RWWV FRQWDPLQDWHG FXOWXUH UHQGHUHG D[HQLF E\ 1DQF\ 6 0RUULVRQ)ORULGD6WDWH8QLYHUVLW\ Nostoc commune FROOHFWHG LQ +XQDQ 3URYLQFH +HQ\RQJ'LVWULFW&KLQD 6 6FKHUHU8 .RQVWDQ] *HUPDQ\ 0 3RWWV N. commune VWUDLQ '5+ LVRODWHGE\'RQQD5+LOO9LUJLQLD7HFK 7KHIROORZLQJIHDWXUHVDSSO\WRN. commune 87(; DQG DUHGHVFULEHG LQ SDUW LQ3RWWV DQG %RZPDQ 6WDWLRQDU\ SKDVH OLTXLG FXOWXUHV RI N. commune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ostoc VS LVRODWHG IURP WKH HQGRF\WRELRVLV ZLWK Geosiphon pyriforme %LOJHU HW DO +RUPRJRQLD GLVSOD\HG DQ HQKDQFHG FDSDFLW\IRUQRQUDGLDWLYHGLVVLSDWLRQRIDEVRUEHGOLJKW HQHUJ\ 7KH PDWXUH WKDOOXV RIN. commune LQ QDWXUH KDV D FKDUDFWHULVWLF VWUXFWXUH )LJE 3URORQJHG LQFXEDWLRQ RIFRORQLHV RQ VHPLVROLG PHGLD JHQHUDWHV VPDOO VSKHULFDO FRORQLHV ³SHDUOV´ ³3HUOHQ GHV 5R]HQNUDQ]HV´ 0ROOHQKDXHU D IURP WR PP LQ GLDPHWHU ZKLFK EXG IURP WKH PDWXUH WKDOOXV 7KH PDVV RI VPDOO FRORQLHV KDV WKH DSSHDUDQFH RI FDYLDU )LJ G LQ 3RWWV 3ODWH G $ VWUDLQ ZDV LVRODWHG IURP WKH LQQHU SRUWLRQ RI RQH RI WKHVH FRORQLHV DQG GHVLJQDWHG N. commune VWUDLQ '5+O 6RPH IHDWXUHV RI WKH OLIH F\FOH RI Nostoc commune DQG N. commune VWUDLQ'5+O ZHUHSUHVHQWHGLQ3RWWV )LJ DQG +LOO HWDO E N. commune VWUDLQ '5+O JURZV LQ OLTXLG FXOWXUH DV FRQWRUWHG ILODPHQWV LQ ORRVH DJJUHJDWHV WKDW IRUP VSKHULFDO PDVVHV ,Q WKH LQLWLDO VWDJHV RI WKHLU IRUPDWLRQ LQ OLTXLGFXOWXUHWKHVHFRORQLHVDSSHDUDVVPDOOEDOOVDQG FDQ EH UHVROYHG ZLWK D GLVVHFWLQJ PLFURVFRSH +RPRJHQL]DWLRQ RI OLTXLG FXOWXUHV DQG WUDQVIHU RI DOLTXRWV RI WKH FHOO VXVSHQVLRQ WR VHPLVROLG PHGLD OHDGV WR WKH GHYHORSPHQW RI PDFURVFRSLF VSKHULFDO FRORQLHV³3HDUOV´SODFHGRQVWHULOHDJDUDWRQHVLGHRI D 3HWUL SODWH XOWLPDWHO\ JHQHUDWH SHDUOV DW VRPH GLVWDQFH IURPWKH LQRFXOXPSUHVXPDEO\ DV DUHVXOWRI WKHIRUPDWLRQRIPRWLOHKRUPRJRQLD $ GLVWLQJXLVKLQJ FKDUDFWHULVWLF RI VWUDLQV RI N. commune LV WKH PDVVLYH DPRXQW RI(36 SURGXFHG LQ OLTXLG DQG LGHQWLILFDWLRQ DV D FDUERK\GUDWHPRGLI\LQJ HQ]\PHV +LOO HW DO D E LW VHHPV OLNHO\ WKDW WKH UROH IRU :VS LV WKH PRGLILFDWLRQPRGXODWLRQ RI WKH JO\FDQ VWUXFWXUH SHUKDSV WKURXJK OLPLWHG K\GURO\VLVRIFURVVOLQNV 6WDLQLQJ RI ³SHDUOV´ ZLWK FDUERK\GUDWHVSHFLILF G\HVDWGLIIHUHQWS+VGHPRQVWUDWHGKRZWKHOD\HUVRI
&KDSWHU 1RVWRF
Fig. 8. /LIHF\FOHRINostoc commune 87(; D /DWH DVHULDWHVWDJH 1RWH YHU\ SURPLQHQW VKHDWK RIWKH FHOOSDFNHWV DQG ODFN RI VKHDWKDURXQGWKHKHWHURF\VWDUURZ WKDWMRLQVWKHSDFNHWVEDU DSSUR[ PP E 6WULQJRI PDWXUHDVHULDWHSDFNHWV WKDWGHULYHGIURPD VLQJOHILODPHQWZLWK WZRKHWHURF\VWV F DVHULDWHSDFNHWVVKRZLQJILQHVWUXFWXUHRIWKHVXUIDFHVKHDWK6(0 G +RUPRJRQLDDIWHUWUDQVIHURI DVHULDWHVWDJHVWRIUHVKPHGLD6(0FRPSDUHZLWK)LJ
0DOFROP 3RWWV
Fig. 9. 7KHOLIHF\FOHRI1RVWRF commune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
FDUERK\GUDWH GLIIHU PDUNHGO\ LQ GLIIHUHQW VHFWLRQV RI WKH WKDOOXV +LOO HW DO E ,Q PDWHULDO RI WKDOOL FROOHFWHG IURP$OGDEUD$WROO LW LV SRVVLEOH WRUHVROYH VLQJOHILODPHQWVWRWDOO\FDOFLILHG 7KHVH DUH UHOHDVHG IURPIUDJPHQWHGFRORQLHVDQGPD\LIYLDEOH VHUYHDV D VRXUFH RI LQRFXOD IRU GLVSHUVDO +LOO DQG 3RWWV XQSXEOLVKHGGDWD 7KH FKDUDFWHULVWLF PRUSKRORJLHV RI Nostoc commune, N. commune YDU flagelliforme DQG N.
parmelioides DUH PDLQWDLQHG LQ DOO WKRVH KDELWDWV ZLWKLQ ZKLFK WKHVH IRUPV DFFXPXODWH 7KHVH FKDUDFWHULVWLF PRUSKRORJLHV IRXQG LQ FRORQLHV IURP WKH 7URSLFV WRSRODUUHJLRQVDUH VWULOXQJ DQG PXVW EH GHWHUPLQHG RQO\ LQ SDUW E\ HQYLURQPHQWDO YDULDEOHV
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ostoc commune 87(; UDLVHG LPSRUWDQW TXHVWLRQV DV WR WKH RULJLQ RI W\URVLQH SKRVSKRU\ODWLRQ DQG WKH HYROXWLRQ RI LWV IXQFWLRQ 3RWWV HW DO 0F&DUWQH\ HW DO $ JHQH OLEUDU\RIN. commune 87(;ZDVFRQVWUXFWHGLQ KJW DQG ZDVSODWHG LQWKHSUHVHQFH RIEURPR FKORURLQGRO\O SKRVSKDWH %&,3 D QRQVSHFLILF VXEVWUDWH IRU SKRVSKRPRQRHVWHUDVHV ZKLFK JHQHUDWHV LQGLJRFDUPLQH D FRORUHG SURGXFW XSRQ K\GURO\VLV RIWKH SKRVSKRHVWHU ERQG LQ WKH SUHVHQFH RIR[\JHQ 2QH EOXH SODTXH ZDV UHFRYHUHG IURP D SODWLQJ RI DURXQG SODTXHV $Q (FR5, IUDJPHQW ZDV VXEVHTXHQWO\ UHFRYHUHG VXEFORQHG LQ D SODVPLG YHFWRU DQG LW ZDV IRXQG WKDWWKLV IUDJPHQW GLUHFWHG WKHV\QWKHVLVRIDVHFUHWHGLQGROHSKRVSKDWHK\GURODVH
,SK3 ;LH HW DO ,Q UHWURVSHFW WKLV ILQGLQJ KDV DOO WKH HOHPHQWV RIVHUHQGLSLW\ JW LV QRW DQ H[SUHVVLRQYHFWRU DQG WKH IDFWWKDWWKH JHQH SURGXFW ZDV FRUUHFWO\ SURFHVVHG VHFUHWHG EH\RQG WKH RXWHU PHPEUDQHRIE. coli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almonella zyphimurium. .DQLJD HW DO ,W VHHPVQRWXQUHDVRQDEOHWKDWin vivo WKHWDUJHWRI,SK3 LV SKRVSKRSURWHLQ HLWKHU RQ N. commune 87(; LWVHOIRURQRWKHURUJDQLVPV ZLWKZKLFK LWFRPHV LQWR FRQWDFW $V LV HYLGHQW IURP WKH SUHFHGLQJ GLVFXVVLRQ WKH FRPSRQHQWV RI VLJQDO WUDQVGXFWLRQ QHWZRUNV PRGXODWHG WKURXJK SURWHLQ SKRVSKRU\ODWLRQ DUH EHLQJ XQFRYHUHG LQ F\DQREDFWHULD 7KH ILQGLQJ RI D FDOPRGXOLQOLNH SRO\SHSWLGH LQ Anabaena VWUDLQV 3HWWHUVVRQ DQG%HUJPDQ LV WKHUHIRUH RIVRPH LQWHUHVW EHFDXVH LQ HXNDU\RWLF FHOOV SURWHLQ SKRVSKRU\ODWLRQGHSKRVSKRU\ODWLRQ QHWZRUNV DUH DWWHQXDWHG FRQVLGHUDEO\ E\ FDOPRGXOLQ $GGHG WR WKLV DUH WKH UHSRUWV RI DEVFLVLF DFLG $%$ D SRWHQWLDO FDOFLXP DJRQLVW LQ F\DQREDFWHULD +LUVFK HW DO 0DUViOHNHW DO ,Q N. muscorum LPSRVLWLRQ RI VDOW VWUHVV OHDG WR WKH DSSHDUDQFH RI $%$ LQ WKH H[WUDFHOOXODU PHGLXP DQG LW ZDV FRQVLGHUHG WKDW LW PD\ EHFRPH LPPRELOL]HG LQ WKH (360DUViOHNHWDO 2QHRIWKHPRVWREYLRXV
0DOFROP3RWWV
HIIHFWVRIDGGLQJH[RJHQRXV$%$WRNostoc ZDV D LQFUHDVH LQ LQWUDFHOOXODU FDOFLXP DQG D GRXEOLQJ RI WKH KHWHURF\VW IUHTXHQF\ 6PLWK HW DO 0DUViOHN HW DO +XGGDUW HW DO 7KHVH GDWDUDLVH LPSRUWDQW TXHVWLRQV WR GR ZLWK &D IOX[HV WKDW PD\ KDYH WKH SRWHQWLDO WR LQGXFH SK\VLRORJLFDO FKDQJHV LQ ILHOG SRSXODWLRQV DQG WKH UROHV LIDQ\ RI$%$ 2Q $OGDEUD$WROO DW\SLFDO YDOXH IRU &D LQ IUHVKZDWHU SRROV ZKLFK VXSSRUW Nostoc JURZWKVLVDURXQGPJ/ P03RWWV DYDOXH VOLJKWO\ KLJKHUWKDQ WKDW DW ZKLFK WKH ELSKDVLF XSWDNH RI &D ZDV VDWXUDWHG LQ Nostoc 0$& 3DQGH\ HW DO 7KH PHDQ FRQWHQW RI &D LQWULFKRPHVRINostoc calcicola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ostoc IRUPV RIWHQ JURZ LQ FORVH SUR[LPLW\ WR RWKHURUJDQLVPV LQFOXGLQJKLJKHUDQG ORZHUSODQWV LW QHFHVVDU\ WR HYDOXDWH DQ\ UHSRUWV RI DEVFLVLF DFLG HLWKHU LQ ILHOG FRORQLHV RU ³D[HQLF´ FXOWXUHV PRVW FDXWLRXVO\
6HFRQGDU\3URGXFWV $ZLGHUDQJH RIVHFRQGDU\SURGXFWV DUHSURGXFHG E\ Nostoc VSS *RODNRWLHW DO .D\DHWDO 6HH &KDSWHU %LRDFWLYH FRPSRXQGV SUHVHQW LQ PHWKDQROLF H[WUDFWV DQG H[WUDFHOOXODU SURGXFWV IURP Nostoc muscorum HYRNHG D VLJQLILFDQW LQKLELWLRQ RI JURZWK RIWKHSK\WRSDWKRJHQSclerotinia sclerotiorum 0XOH HW DO 7KH F\FOLF GHSVLSHSWLGHV QRVSHSWLQV $ DQG % ZHUH LVRODWHG IURPN. minutum 1,(6 DQG ZHUH IRXQG WR EH SRWHQW LQKLELWRUV RI HODVWDVHDQGFK\PRWU\SVLQ2NLQRHWDO 7ZR EDFWHULRKRSDQRLGV ZLWK DF\FOLF SHQWRO VLGHFKDLQV ZHUHSXULILHGIURPNostoc VSVWUDLQ3&&=KDR HWDO $%RHVHNHQFRPSOH[RI ERULFDFLGZDV UHSRUWHG IURP WKH PDULQH IRUP Nostoc linkia +HPVFKHLGWHWDO ,QWKLVVWUDLQWKHVWUXFWXUDO VNHOHWRQV RI SDUDF\FORSKDQHV ZHUH SRO\NHWLGHV WKDW IRUPHG IURP WKH GLPHUL]DWLRQ RIDFHWDWHGHULYHG
QRQDNHWLGHV %RE]LQ DQG 0RRUH 7KH V\QWKHVLV RI KHWHURF\VW JO\FROLSLGV LV WKRXJKW WR RULJLQDWH ZLWK SRO\NHWLGHV DQG D JHQH HQFRGLQJ D SRO\NHWLGHV\QWKDVHOLNHHQ]\PHZDV FKDUDFWHUL]HG LQ N. punctiforme VWUDLQ $7&& &DPSEHOO HWDO
C.
Cyanoglobins
7KHJHQHHQFRGLQJF\DQRJORELQ glbN, ZDV LGHQWLILHG WKURXJK VHTXHQFH DQDO\VLV RIWKH nifU-nifH UHJLRQ RI Nostoc commune 87(; 3RWWV HW DO *OE1 LV D PRQRPHULF KHPRJORELQ WKDW ELQGV R[\JHQ UHYHUVLEO\ ZLWK KLJK DIILQLW\ DQG ZLWK QRQ FRRSHUDWLYLW\7KRUVWHLQVVRQHWDO &\DQRJORELQLV V\QWKHVL]HG RQO\ LQ FHOOV VWDUYHG RI FRPELQHG QLWURJHQ ZKHQ WKH\ DUH VXEMHFWHG WR PLFURR[LF RU DQR[LF FRQGLWLRQV +LOO HW DO 7KHDYDLODEOHGDWDVXJJHVWWKDW*OE1PD\IXQFWLRQDV SDUWRIDWHUPLQDOF\WRFKURPHR[LGDVHFRPSOH[WKDWLV QHHGHG XQGHU PLFUR[LF FRQGLWLRQV ZKHQ $73 VXSSO\ IRU QLWURJHQ IL[DWLRQ PD\ EH OLPLWHG +LOO HW DO ¶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aramecium DQG Tetrahymena) DQG WKH JUHHQ DOJD Chlamydomonas, WR D F\DQREDFWHULDO DQFHVWRU 7KH DOLJQPHQW RI D UHVWULFWHG QXPEHU RI VHTXHQFHV LV SURYLGHG KHUH DQG LQFOXGHV LQ DGGLWLRQ WKRVH RI D SXWDWLYH JORELQ RI Synechocystis 3&& D UHSUHVHQWDWLYH RI D QHZ FODVV RI QRQOHJKHPRJORELQ VHTXHQFHV IURP SODQWV $QGHUVVRQ HW DO DQG WKH VHTXHQFH RI WKH F\WRFKURPH R[LGDVH RI Pseudomonas perfectomarina (stutzeri) -LLQJVW HW DO ZKRVH 1WHUPLQDO UHJLRQ VKDUHV VHTXHQFH VLPLODULW\ ZLWK Nostoc *OE1 +LOO HW DO (YLGHQFHWRVXSSRUWDKRUL]RQWDOWUDQVIHUIURP DQ
&KDSWHU 1RVWRF
Fig. 10. $OLJQPHQW RI VHOHFWHG KHPRSURWHLQV DQG F\DQRJORELQV 6HTXHQFHVZHUHDFTXLUHG IURPWKH *HQEDQN3,5DQG&\DQREDVH GDWDEDVHV $EEUHYLDWLRQV7S Tetrahymena pyriformis; 6V Synechocystis 3&& 6803; &U Chlamydomonas reinhardtii; 3F Paramecium caudatum; 1F Nostoc commune 87(; 584; 1V 1RVWRF 081 6F Saccharomyces cerevisiae; 9V Vitreoscilla VS (F Escherichia coli; $H Alcaligenes eutrophus; &Q Candida norvegensis; 3D Parasponia andersonii; *P Glycine PD[ *P Glycine max QRQOHJKHPRJORELQ 3S Pseudomonas perfectomarina; Vp, Vibrio parahaemolyticus; %M Bradyrhizobium japonicum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
DQFHVWUDOFLOLDWHWRDQDQFHVWRURINostoc LVQRWUHDGLO\ DSSDUHQW)LJ DOWKRXJKWKHUHVWULFWHGGLVWULEXWLRQ RI glbN PD\ LQLWVHOIVXJJHVW LW ZDV DFTXLUHG WKURXJK ODWHUDOWUDQVIHU 7KH VHTXHQFH VLPLODULW\EHWZHHQ WKH SURWR]RDQ VHTXHQFHV DQG F\DQREDFWHULDO JORELQV LV DSSDUHQW,QYLHZRIWKHIDFWWKDWSURWR]RDRIWHQRFFXU LQDVVRFLDWLRQZLWKNostoc FRORQLHVDQGWKHUDQJHRI V\PELRWLF DVVRFLDWLRQV EHWZHHQ SURWR]RD DQG F\DQREDFWHULD &DUU DQG :KLWWRQ WKH VXJJHVWLRQ RI WKH WUDQVIHU RI DQ DQFHVWUDO JHQH EHWZHHQ PHPEHUV RIWKHVH WZR JURXSV DSSHDUV WR EH QRWXQUHDVRQDEOH *LYHQWKHLGHQWLILFDWLRQRISURWHLQV RI N'D LQ D UDQJH RI F\DQREDFWHULD WKDW DUH LPPXQRORJLFDOO\UHODWHG WR *OE1 +LOO HW DO DQG WKH LGHQWLILFDWLRQ RI D SXWDWLYH KHPRSURWHLQ LQ Synechocystis VS VWUDLQ 3&& ZKLFK VKRZV
REYLRXVVHTXHQFHVLPLODULW\WRSURWR]RDQJORELQVLWLV SRVVLEOH WKDW ³F\DQRJORELQV´ PD\ EH ZLGHVSUHDG LQ F\DQREDFWHULD WKH\ PD\ KDYH TXLWH GLIIHUHQWUROHV WR ILW GLIIHUHQW HFRORJLFDO FRQWH[WV 7KH JHQH WKDW PD\ HQFRGH WKH JORELQ LQ Synechocystis 3&& LV SRVLWLRQHG ZLWKLQ D FOXVWHU RIJHQHV WKDW PD\ HQFRGH VHQVRU\ WUDQVGXFWLRQ KLVWLGLQH NLQDVHV D VLWXDWLRQ ZKLFKKDVREYLRXVSDUDOOHOVLQRWKHUEDFWHULDOV\VWHPV *LOOHV*RQ]DOH]HWDO 7KH UHVWULFWHG GLVWULEXWLRQ RI glbN DQG *OE1 WR D VPDOO VXEVHW RI Nostoc VWUDLQV VXJJHVWHG WKDW WKH SUHVHQFH RI glbN PD\ SURYLGH D XVHIXO JHQRW\SLF PDUNHU +LOO HW DO 7KH PDUNHU LV SUREDEO\ LQGLFDWLYHRID VKDUHGSK\VLRORJLFDO FDSDFLW\E\WKHVH VWUDLQV DQG RQH WKDWPD\ EH RIHFRORJLFDO UHOHYDQFH ,QWKLVFRQWH[WLWLVRILQWHUHVWWKDWN. commune VWUDLQ '5+ WKH GHULYDWLYH RI ILHOG PDWHULDO RI N. commune VHHPVWRODFN*OE1+LOOHWDO 7KH VHTXHQFH VLPLODULW\ EHWZHHQ WKH UHJLRQV ERWK XSVWUHDP DQG GRZQVWUHDP RI glbN LQ N. commune DQG Nostoc VS LV KLJK ZLWK WKH H[FHSWLRQ WKDW Nostoc VS ODFNV WKH 1WF$ELQGLQJ VLWH LGHQWLILHG LQ WKH nifU-glbN LQWHUJHQLF UHJLRQ RI N. commune +LOO HW DO -DFNPDQQ DQG 0XOOLJDQ &XULRXVO\ WKH 1WF$ ELQGLQJ UHJLRQ LQ N. commune LV IODQNHG E\ VKRUW WDQGHP UHSHDWV$$77$&* DQGLQYHUWHGUHSHDWVSURYLGHWKH SRWHQWLDO IRU WKLV UHJLRQ WR IROG LQWR D VWDEOH VHFRQGDU\VWUXFWXUH+LOOHWDO 7KHIXQFWLRQV RI VXFK HOHPHQWV LV XQUHVROYHG 0DVHSRKO HW DO 3HUKDSVWKH1WF$ELQGLQJVLWHRIN. commune glbN ZDV DGGHG WR WKH SURPRWHU WKURXJK D VHSDUDWH UHFRPELQDWLRQ HYHQW DIWHU DFTXLVLWLRQ RIWKH DQFHVWUDOJHQH ,I DVVHHPV WREHWKHFDVHglbN KDV LWV RZQ SURPRWHU RQH PXVW VWLOO TXHVWLRQ ZK\ WKH JHQH LV SRVLWLRQHG EHWZHHQ WKH nif JHQHV RI WZR FRQWLJXRXV nif RSHURQV Nostoc 081 8820 FRQWDLQV WKH glbN JHQH SRVLWLRQHG EHWZHHQ nifU DQG nifH EXW ODFNV WKH $$77$&* 6755 VHTXHQFHV SUHVHQW LQ WKH nifUglbN UHJLRQ RIN. commune 87(; +LOO HW DO
' 'HVLFFDWLRQ 7ROHUDQFH 6LJQLILFDQFH 7KHSLYRWDOUROHRIZDWHULQWKHHPHUJHQFHRIOLIHLV UHDGLO\DFNQRZOHGJHG ,QFRQWUDVW WKH LPSRUWDQFH RI
0DOFROP3RWWV
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seudoanabaena galeata *HLWOHU /\RSKLOL]DWLRQRIFXOWXUHVRIF\DQREDFWHULD EHORQJLQJWR VHYHQJHQHUDZDVPHDVXUHGLQDYDULHW\ RIVXVSHQGLQJVXEVWDQFHV 7KHUHZDVYDULDWLRQLQWKH VXUYLYDO GHSHQGLQJ RQ WKH PHGLXP :LWK WKH H[FHSWLRQ RI Nostoc muscorum, QRQH RI WKH VWUDLQV VXUYLYHG O\RSKLOL]DWLRQ LQ VNLP PLON HJJ DOEXPLQ PHGLXPRU-DQVHQ VDOWVPHGLXP&RUEHWWDQG3DUNHU
7DEOH SURYLGHVGDWDWKDWHPSKDVL]HWKHORZOHYHO RI K\GUDWLRQ RI WKH F\WRSODVP LQ N. commune DQG RWKHUDQK\GURELRWHV
1DWXUDO&RPPXQLWLHV 7KH FDSDFLW\ RI ILHOG SRSXODWLRQV RI Nostoc WR ZLWKVWDQG F\FOHV RI GHVLFFDWLRQ DQG UHK\GUDWLRQ ZDV NQRZQIRUDFRQVLGHUDEOHDPRXQWRIWLPH6HFWLRQ9 ,W WRRN KRXUV IRU WKH WKDOOL RI GHVLFFDWHG N. commune WR EHFRPH IXOO\ K\GUDWHG LQ ILHOG WHVWV LQ 6FKLUPDFKHU 2DVLV $QWDUFWLFD &KDSWHU 6LPLODUO\ XSRQ GHK\GUDWLRQ ZDWHUORVV ZDV FRPSOHWH ZLWKLQ KRXUV 8QGHU ODERUDWRU\ FRQGLWLRQV ZDWHU JDLQSURFHHGHGDWDUDWHIROG IDVWHUWKDQLQWKHILHOG *XSWDDQG .DVK\DS Nostoc commune PDWV LQ 0F0XUGR 6RXQG $QWDUFWLFD UHFRYHUHG WR SUH GHVLFFDWLRQ UDWHV RI SKRWRV\QWKHVLV DQG UHVSLUDWLRQ ZLWKLQDVOLWWOHDV PLQRIUHZHWWLQJ ZKLOHUHFRYHU\ RI DFHW\OHQH UHGXFWLRQ ZDV VORZHU ! K Phormidium PDWV ZHUH OHVV WROHUDQW RI GHVLFFDWLRQ DQG UHTXLUHG GD\V IRU UHFRYHU\ 7KH SDUWLDO K\GUDWLRQ RI N. commune GXULQJ DHULDO H[SRVXUH WHQGHGWRVORZUHFRYHU\XSRQIXOOUHK\GUDWLRQ+DZHV HWDO 7KH ILUVW GHVFULSWLRQ RI WKH VWUXFWXUDO SURSHUWLHV RI GHVLFFDWHG F\DQREDFWHULD ZDV IRU WKRVH LVRODWHG IURP KRWGHVHUW URFNV 3RWWV HW DO 6XEVHTXHQW VWXGLHV ZLWK Nostoc commune GHVFULEHG WHFKQLTXHV WR SHUPLW WKH IL[DWLRQ DQG SUHSDUDWLRQ RI GHVLFFDWHG FHOOV WKDW DYRLGHG DUWLIDFWV DQG SHUPLWWHG LPPXQRF\WRFKHPLVWU\ 3HDW DQG 3RWWV 3HDW HW DO $SSOLFDWLRQ RI WKH ODWWHU WHFKQLTXH FRQILUPHG WKDW WKH ZDWHU VWUHVV SURWHLQV :VS RIN. commune ZHUH VHFUHWHG SURWHLQV LQWLPDWHO\ DVVRFLDWHG ZLWK WKH H[WUDFHOOXODU JO\FDQ 7KH ILQGLQJ WKDW )H SURWHLQ RI QLWURJHQDVH ZDV VWDEOH LQ FHOOV GHVLFFDWHG IRU PRUH WKDQ D GHFDGH FRPSOHPHQWHG VWXGLHV GLUHFWHG DW WKH DQDO\VLV RI SURWHLQ VWDELOLW\ 3HDWHWD
0HFKDQLVPV )ROORZLQJ WKH WUDQVIHU RI FHOOV RI N. commune 87(; IURP OLTXLG FXOWXUH WR WKH DLU WKH FHOOV ORRVH WKH FDSDFLW\ WR IL[ QLWURJHQ ZLWKLQ DSSUR[LPDWHO\ PLQXWHV EXW PDLQWDLQ D SRRO RI LQWUDFHOOXODU $73 HTXLYDOHQW WR WKDW LQ OLTXLG JURZLQJ FHOOV IRU VHYHUDO KRXUV 8SRQ UHK\GUDWLRQ GHVLFFDWHG FHOOV RI N. commune UHFRYHU WKH FDSDFLWLHVIRUUHVSLUDWLRQSKRWRV\QWKHVLVDQG QLWURJHQ
&KDSWHU 1RVWRF
Table 1. 3URWHLQVLQGHVLFFDWHG N. commune ODFNDPRQROD\HUFRYHUDJHRIZDWHU
3URWHLQ K\GUDWLRQD J + J SURWHLQ
(YHQW
&RQVHTXHQFH IRU SURWHLQ
&HOOK\GUDWLRQ J +JFHOO VROLG
WR
FKDUJHGJURXSK\GUDWLRQ SRODU JURXS K\GUDWLRQ GLVRUGHURUGHU WUDQVLWLRQ
SURWRQUHGLVWULEXWLRQ GLVXOILGHVUHRUGHU SURWHLQ ORRVHQVXS
DQK\GURELRWLFFHOO HJN. commune
WR
DFLGVVDWXUDWH FOXVWHUV PRELOL]H VLGHFKDLQSRODU VDWXUDWLRQ
VLGHFKDLQDQG EDFNERQH FRQIRUPDWLRQDOVKLIWV
WR
SHSWLGH 1+ VDWXUDWHV ZDWHU VWUXFWXUH FRQGHQVDWLRQ DQG UHDUUDQJPHQW
onset of enzyme activity
WR
SRODU JURXS PRQROD\HU FRYHUDJH ILQDOO\ FRPSOHWH PRQROD\HUFRYHUDJH
G\QDPLFLFUHDVH WR VROXWLRQ YDOXH ZLWK LQFUHDVH LQ DFWLYLW\
WR
water content of protein crystals
0.02 to 0.05
decreasing
H2O
Bacillus VSRUH 0.21 to 0.58 DW 0.3 VHQVLWLYH EDFWHULD GLH
D6RPH GDWD DGDSWHG IURP UHIV LQ 3RWWV
IL[DWLRQ LQ WKDW RUGHU IRU UHIV VHH 3RWWV 0HO QLNRYD QRWHG WKDW RQH RIWKH HDUO\ HYHQWV WR RFFXUXSRQGHK\GUDWLRQ RINostoc ZDV DQ LQFUHDVH LQ DQ DELOLW\ RIWKH QXFOHDU PDWHULDO WR ELQG DFULGLQH RUDQJH \RXQJ JURZLQJ FHOOV DSSHDUHG PRUH UHVLVWDQW WKDQPDWXUHFHOOVDERXWWRGLYLGH &KDUDFWHUL]DWLRQ RI GHVLFFDWHG FRORQLHV RI Nostoc commune LGHQWLILHG D QRYHO FODVV RI DFLGLF SRO\SHSWLGHV ± WKHVH ZHUH WHUPHG ZDWHU VWUHVV SURWHLQV:VS DQGWKH\FRQVWLWXWHG VRPHRIWKH WRWDO VROXEOH SURWHLQ RI GHVLFFDWHG FHOOV :VS SRO\SHSWLGHVDUHKLJKO\VWDEOHLQGHVLFFDWHGFHOOV DQG WKH\DUHLQGXFHGLQOLTXLGJURZQFXOWXUHVLIWKRVHFHOOV DUH VXEMHFWHG WR PXOWLSOH F\FOHV RI GU\LQJ 6FKHUHU DQG3RWWV ,QLWLDOO\LWZDVQRWDSSUHFLDWHGWKDW :VSSURWHLQV DUHVHFUHWHGE\N. commune, EXWLWZDV QRWHG WKDW WKH DFFXPXODWLRQV RI :VS LQ ODERUDWRU\ JURZQ FXOWXUHV QHYHU DFKLHYHG WKRVH SUHVHQW LQ ILHOG PDWHULDOV 6FKHUHU DQG 3RWWV SURYLGH D SUHOLPLQDU\FKDUDFWHUL]DWLRQRI:VS 5HFHQWO\LWZDV VKRZQ WKDW :VS SRO\SHSWLGHV DUH VHFUHWHG DQ XQH[SHFWHG ILQGLQJ JLYHQ WKH FRQFHQWUDWLRQV RIWKHVH SURWHLQV :VS DVVRFLDWH ZLWK D ' [\ODQ[\ODQRK\GURODVH DFWLYLW\ WKDW LV DOVR VHFUHWHG IURP WKH FHOOV WKH\ IRUP FRPSOH[HV ZLWK VHFUHWHG
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co 5, IUDJPHQW ZDV LVRODWHG UHFHQWO\ IURP D OLEUDU\ RI 1 commune 87(; '1$ LQS7UF$ 7KH IUDJPHQW ZDV FRQWDLQHG LQ D FORQH WKDW VKRZHG LQWHQVH FURVVUHDFWLRQ ZLWK :VS DQWLERGLHV 2IWKH 25)V WKDWKDYH EHHQ LGHQWLILHG RQH VKRZV FRQVSLFXRXV GHULYHG DPLQR DFLG VHTXHQFH VLPLODULW\ WR SURWHLQV LQYROYHG LQ WKH VHFUHWLRQ RI F\FOLF JOXFDQV E\ Agrobacterium tumefaciens DV ZHOO DV +HS$ IURPAnabaena 3&& D SURWHLQ
0DOFROP3RWWV
LQYROYHG LQ WKH GHYHORSPHQW RI WKH KHWHURF\VW FDUERK\GUDWH ZDOO OD\HU+ROODQG DQG :RON 7KH RWKHU 25)V DOO VKRZ FRUUHVSRQGHQFH ZLWK FDUERK\GUDWHPRGLI\LQJHQ]\PHVXQSXEOLVKHGGDWD ,W DSSHDUV WKDWWKH FDSDFLW\ WR PDLQWDLQ SURWHLQV LQ D IXQFWLRQDOO\VWDEOH VWDWH LV D FHQWUDO IHDWXUH RI WKH GHVLFFDWLRQ WROHUDQFH RIN. commune 3RWWV )RUH[DPSOH WKHRQVHW RIWKHV\QWKHVLV RIDOOFODVVHV RIOLSLG LQGHVLFFDWHGFHOOVUHVXPHVXSRQUHK\GUDWLRQ LQVWDQWDQHRXVO\ 7DUDQWR HW DO 6RPH RIWKH SRWHQWLDO UROHV RI SURWHLQV LQ UHVSRQVHV RI FHOOV WR ZDWHUGHILFLWDUHVXPPDUL]HGLQ7DEOH 3URGXFWLRQ RI (36 DV GHVFULEHG LQ 3RWWV PD\ DOVR UHSUHVHQW D PHFKDQLVP IRU GHVLFFDWLRQ WROHUDQFH7KH(36 RINostoc commune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
(36 WR SUHYHQW PHPEUDQH OHDNDJH WKH DW\SLFDO VWUXFWXUH RI WKH RXWHU PHPEUDQH DQG WKH PDUNHG GHSUHVVLRQ LQ WKH PHPEUDQH SKDVH WUDQVLWLRQ WHPSHUDWXUH RIGHVLFFDWHG FHOOV ZKLFK ZDV PHDVXUHG in vitro WKURXJK )RXULHU WUDQVIRUP LQIUDUHG VSHFWURVFRS\FRQVWLWXWHSURWHFWLYHPHFKDQLVPVZKLFK SUHYHQW PHPEUDQH GDPDJH GXULQJ GHVLFFDWLRQ RU IUHH]HGU\LQJ DQGUHK\GUDWLRQ+LOOHWDO
9 3DWHQWVDQG$SSOLFDWLRQV $ VPDOO QXPEHU RI SDWHQWV GHVFULEHG PHWKRGV WKDW HPSOR\ ZKROH FHOOV RU FRPSRQHQWV RI Nostoc VSS 7KH OLIH F\FOH RI Nostoc muscorum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ostoc DQG Anabaena DV KRVWFHOOV ZLWKLQ ZKLFK WR V\QWKHVL]H WKH Bacillus thuringiensis VVS israeliensis cryIVD SURGXFW 7KH ODWWHU LV DFWLYH DV D SHVWLFLGH DJDLQVW'LSWHUD *HQHV HQFRGLQJ WKH 1VS9 UHVWULFWLRQ PRGLILFDWLRQ HQ]\PHV IURP Nostoc 3&& ZHUH LVRODWHG DQG D PHWKRG ZDV GHYLVHG WR H[SUHVV WKH DFWLYHHQ]\PHVLQE. coli 8HQRHWDO Nostoc DQG RWKHUF\DQREDFWHULD IRUP WKH EDVLV RID VXEVWUDWH GHVLJQHG WR SURGXFH YHJHWDWLRQ RQ EDUH WHUUDLQ
Table2. 3RWHQWLDOUROHVIRUSURWHLQVLQWKHGHVLFFDWLRQUHVSRQVHRIN. commune
VSHFLILF SHSWLGDVH VSHFLDO VLJPD IDFWRUV FRQWURO GHUHSUHVVLRQ RI FDWDEROLF HQ]\PHV FDWDEROLF HQ]\PHV SURWHLQV IRU VXUYLYLQJ VWUHVV
VHQVRU\ SURWHLQV VWUXFWXUDO SURWHFWLYH SURWHLQV
SURWHLQ WDUJHWLQJ HJ XELTXLWLQ WR LQGXFH H[SUHVVLRQ RI VWUHVV JHQHV WR HVFDSH VWUHVV WR SURYLGH HQHUJ\ WR PDLQWDLQ KRPRHRVWDVLV WR PDLQWDLQ S+ HWF WR VWDELOL]H '1$ DQG 51$ WR XQFRXSOH YLDELOLW\ IURP FHOO GLYLVLRQ WR UHJXODWH UHFUXLWPHQW RI P51$ WR UHJXODWH FDUERK\GUDWH PHWDEROLVP WR WUDQVGXFH WKH VWUHVV WR DFKLHYH WROHUDQFH SURWHFWLRQ "
&KDSWHU 1RVWRF &KLDIIUHGR DQG )LJXUHDX &HOO H[WUDFWV RI Nostoc DQG RWKHU F\DQREDFWHULD ZHUH XVHG LQ D IORFFXODWLRQ SURFHVV WR WUDS PLQHUDOV /HVOLH HW DO DQG DOVR WR SURGXFH D GULHG IORZDEOH ZDWHU VXVSHQGLEOH FRPSRVLWLRQ IRU WKH WUHDWPHQW RI VRLOV 6FKDHIHU DQG %R\XP 0HUFN DQG &R ,QF GHYHORSHG VHYHUDO SDWHQWV IRU QDWXUDO SURGXFWV RI Nostoc VSS WKDW DUH HIIHFWLYH DJDLQVW Cryptococcus VS DFDXVDODJHQWRIVHFRQGDU\ IXQJDO LQIHFWLRQV LQ SDWLHQWVZLWK$,'6+LUVFK HWDO Nostoc punctiforme ZDV IRXQG WR EH HIIHFWLYH LQ WUHDWLQJ HIIOXHQW IURP WKH SRO\ILEUH LQGXVWU\ WKDW FRQWDLQHG GRPHVWLF VHZDJH +RVHWWL DQG 3DWLO ,Q YLHZRILWV JURZWK FKDUDFWHULVWLFV LQ DQHZJURZWK PHGLXP %: D Nostoc VS ZDV FRQVLGHUHG WR EH SURPLVLQJ DV D VRXUFH RI SK\FRELOLSURWHLQV IRU WKH SURGXFWLRQRIQDWXUDOG\HV6LOYDHWDO 7KH ZLGH VFRSH RI WKHVH WHFKQLTXHV VXJJHVW FRQVLGHUDEOH DSSOLFDWLRQV IRU ZKROH FHOOV RI Nostoc VSSDVZHOODVWKHLUQDWXUDOSURGXFWV
9, $&KURQRORJ\RI Nostoc 7KH GLVFRYHU\ DQG VXEVHTXHQW VWXG\ RI Nostoc FRQVWLWXWHVDIDVFLQDWLQJDQGDKXPRURXVVWRU\$GGHG WRWKLVDUHDQXPEHURISHFXOLDULWLHVRILWV LQWHUDFWLRQV ZLWKKXPDQVWKDWZDUUDQWGLVFXVVLRQ
7& +XDQJ SHUV FRPP <7 :RQJ SHUV FRPP &77HQJSHUVFRPP 7KH %HQ&DR*DQJ0X RU 3HQ7VDR.DQJ0X *HQHUDO 2XWOLQHV DQG 'LYLVLRQV RI+HUEDO 0HGLFLQH E\ 6KL=KHQ /L " RI WKH 0LQJ '\QDVW\ /L KDV YROXPHV DQG OLVWV DW OHDVW PHGLFLQHVDPRQJZKLFKDUHNostoc commune DQG N. commune YDU flagelliforme. 7KH ODWWHU LV UHIHUUHG WR DV ³)D&DL´ LQ PDQGDULQ &KLQHVH )D PHDQV KDLU RI KXPDQKHDG &DLPHDQVYHJHWDEOH 7KHGHULYDWLRQ RI WKH ZRUG LV UHDGLO\ H[SODLQHG IURP WKH KDLUOLNH PRUSKRORJ\ RI GHVLFFDWHG FRORQLHV $UXJD (DUOLHVWGHVFULSWLRQV RI)D&DL GDWH EDFN WR WKH 6XQJ '\QDVW\$' 7KHSURQXQFLDWLRQRI)D&DL RU )D7VDL LV VLPLODU WR WKH SURQXQFLDWLRQ RI D SKUDVH WKDW PHDQV ³DFTXLULQJ ZHDOWK´ RU ³PDNH PRQH\´ $ GLIIHUHQW ZRUG EXW ZLWK WKH VDPH SURQXQFLDWLRQ LV DOVR SDUW RI WKH JUHHWLQJ ³JRQJ[L IDFDL´ ZKLFK PHDQV ³ZLVK \RX SURVSHULW\´ 7KHVH GLIIHUHQW SOD\V RQ ZRUGV DFFRXQW IRU WKH WUDGLWLRQDO HDWLQJRIN. commune YDU flagelliforme DVDGHOLFDF\ DWWKHWLPHRIWKH&KLQHVH1HZ
$ 7KH+DLU9HJHWDEOH %HFDXVH Nostoc commune JURZVLQWKHZLOGZLWKRXW URRWV VWHPV RU OHDYHV DQG IORDWV RQ ZDWHU WKH &KLQHVHFRPSDUHGLWWRWKHLPPRUWDOV ZKRDUHVDLG WR WUDYHO WKH ZRUOG DW ZLOO Nostoc commune EHFDPH NQRZQ DV 7LDQ;LDQ0L 5LFH RI +HDYHQO\ ,PPRUWDOV&KLQHVHQDPHVDQGSODFHVZHUHWUDQVODWHG DFFRUGLQJ WR WKH 3LQ\LQ V\VWHP SURSRVHG E\ WKH 3HRSOH¶V 5HSXEOLF RI &KLQD RU 7LDQ;LDQ&DL 9HJHWDEOH RI +HDYHQO\ ,PPRUWDOV $FFRUGLQJ WR OHJHQG +XQJ*H DQ DOFKHPLVW SK\VLFLDQDQG7DRLVW WKHRUHWLFLDQRIWKH(DVWHUQ-LQ'\QDVW\ $' XVHG N. commune DV D VWDSOH IRRG GXULQJ SHULRGV RI IDPLQH ZKHQ KH ZDV D KHUPLW LQ 6RXWKHUQ &KLQD +XQJ *H DWWULEXWHG KLV KHDOWK WR WKH FRQVXPSWLRQ RI N. commune DQGKH ODWHULQWURGXFHGN. commune WR WKH (PSHURU IROORZLQJ DQ LQYLWDWLRQ WR &RXUW 7KH DLOLQJ (PSHURU LV VDLG WR KDYH UHJDLQHG KLV KHDOWK DIWHU HDWLQJ Nostoc DQG ZDV so JUDWHIXO WKDW KH JUDQWHGWKHQDPH*H;LDQ0L5LFHRI,PPRUWDO *H IRU WKH F\DQREDFWHULXP LQ KRQRU RI +XQJ *H WKH QDPH LVUHWDLQHG WR WKLV GD\ -6 &KHQ SHUV FRPP
Fig. I1. *RRG KHDOWK DQG ZHDOWK 7KH FRYHU RI D ER[ RI N. commune YDU flagelliforme VROG DW WKH WLPH RI WKH &KLQHVH 1HZ
7KH IROORZLQJ ZDV WUDQVODWHG IURP D SRSXODU &KLQHVH QHZVSDSHU “In Ninxia, Northwest China, he pointed to a barren hill beyond the ditch and indicated to us that this is where Fa-Cui is grown. The place shown is surrounded by barbwire and guarded by a vicious dog and a guard hut. I walked up the hill and saw for the first time the cultivated Fa-Cui which is also called black gold” (for its price; Suen, 1994). Its price is based on gram
quantities. With the price skyrocketing, it has been collected aggressively to near extinction. A one-year medium-scale test showed that from one Mu (a Chinese unit of land measurement equivalent to about 14.8 acres), the equivalent to about US$140 (in 1994) worth of Fa-Cai could be produced.” 7KHUH VHHPV WR EH TXLWH D EXVLQHVV LQ Nostoc. ,Q WKHPDUNHWRI,ORFRV1RUWH/DRDJ&LW\3KLOLSSLQHVN. commune LVVROGLQGULHGIRUP FXULRXVO\LQWKHILVK VHFWLRQUDWKHUWKDQWKHYHJHWDEOHVHFWLRQRIWKHPDUNHW DQGVWRUHGGU\LQNJMXWHVDFNV0DUWLQH] 7KH ILQGLQJ RI Nostoc JURZWKV LV WDNHQ ZLWK WKH XWPRVWVHULRXVQHVVLQ&KLQD $FFRUGLQJWRDUHSRUWLQ WKH RYHUVHDV HGLWLRQ RI The People’s Daily: “good news came recently from the Shi-Wu-Tou Village, Xin-Hua County, Shen-Nueng-Jia Forest District, Hubei Province (in Southeast China). The rare, precious Ge-Xian-Mi has been found to grow in about ten Mu (about 148 acres) of water-filled paddy in that village. The annual yield reached five kilograms. According to the literature, the only other places that also produce Ge-Xian-Mi are: six Mu (about 90 acres) in Africa and 0.7 Mu (about 10 acres) of water-filled paddy or swamp in Xiang-Fan of Hubei Province. At present, there is no whole-sale quantity of Ge-Xian-Mi on the domestic or overseas market, so the price for it is very high. ¶¶ /LNH DOO WKLQJV LQ &KLQD WKH HDWLQJ RI Nostoc LV OLNHO\ WR KDYH JRQH RQ IRU D FRQVLGHUDEOH SHULRG RI WLPH VRPHWKLQJ WKDW *HLWOHU DSSHDUHG WR EH XQDZDUH RI ZKHQ KH LPSOLHG WKDW QDWLYHV VKRXOG SXW N. commune WRFXOLQDU\XVH*HLWOHU Nostoc LV D FRPPRQ GLHWDU\ VXSSOHPHQW IRU WKH HQGLJHQRXV SRSXODWLRQV RI 7KDLODQG 3HUX &KLQD (FXDGRU )LML -DYD -DSDQ 0H[LFR 0RQJROLD DQG 6LEHULD ZKR KDYH ORQJ DSSUHFLDWHG WKH SODQWOLNH QXWULWLRQDO SURSHUWLHV RID QXPEHU RI GLIIHUHQW IRUPV LQFOXGLQJ N. commune YDU flagelliforme, N. parmelioides (edule), N. ellipsosporum, N. verrucosum DQG N. pruniforme -DVVE\ 3ODWH OL 'RHVLWQRWVHHPUHDVRQDEOHWR TXHVWLRQZKHWKHU SULPLWLYHPDQDOVRVDPSOHGVXFKGHOLFDFLHV"
7KH3KLORVRSKHU¶V6WRQH 'HVFULSWLRQV RINostoc IURP (XURSH DSSHDUHG VRPH \HDUVDJR ZHOO EHIRUH WKH VFRXUJH RIWKH %ODFN 'HDWK LQ WKH V DQG PRUH WKDQ \HDUV EHIRUH $QWRQLH YDQ /HHXZHQKRHN GHVFULEHG DQLPDOFXOHV XQGHUKLVSURWRW\SHPLFURVFRSHLQ3RWWV ,W LV FOHDU ZK\ Nostoc FDPH WR SURPLQHQFH DW WKLV WLPH FRORQLHV RI N. commune LQ VRLOV DSSHDU DV
0DOFROP3RWWV EODFNHQHG EULWWOH QRQGHVFULSW FUXVWV ZKHQ WKH\ DUH GULHG EXW EHFRPH FRQVSLFXRXV DQG VZROOHQ DV GDUN EOXHJUHHQ RU ROLYHJUHHQ PDVVHV ZLWK WKH FRQVLVWHQF\ RI ILUP JHODWLQ ZKHQ WKH\ DUH ZHWWHG )LJ OE 7KH VZHOOLQJ FRORQLHV LV UDSLG WKHUH LV D FRQVLGHUDEOH LQFUHDVH LQ WKHLU EXON DQG RIWHQ UHK\GUDWLRQ LQGXFHV WKH UHOHDVH RI WKH FKDUDFWHULVWLF ³HDUWK\´ VPHOO RI trans- GLPHWK\OWUDQV GHFDORO RU JHRVPLQ 1DHV $Q\WKLQJ WKDW LV FRQVSLFXRXV DQG WKDW KDV DQ XQXVXDO VPHOO WHQGV WR FDWFK SHRSOH¶V DWWHQWLRQ 7KH UDSLG DSSHDUDQFH RI JURZWKVRIN. commune DIWHUWKXQGHUVKRZHUVOHDGWR WKHFRPPRQ EHOLHIGXULQJ WKH 0LGGOH $JHV WKDW VXFK FRORQLHVIHOOIURPWKHVN\DQGWKH\ZHUHUHIHUUHGWRDV ³6WHUQVFKQXSSHQ´ VKRRWLQJ VWDUV 0ROOHQKDXHU D “Nostoch is that which we call a falling star, a kind of gelly or slime found oftentimes in the Summer in fields, and meadowes” )UHQFK 7KH GLIIHUHQW QDPHV XVHG WR GHVFULEH WKH FRORQLHV RI WKLV F\DQREDFWHULXP 7DEOH DWWHVW QRW RQO\ WR WKH OLYHO\ QDWXUH RIKXPDQ VXSHUVWLWLRQ DQG LPDJLQDWLRQ WKH\ HPSKDVL]H WKDW Nostoc VSS PD\ KDYH KDG D ZLGHVSUHDG RFFXUUHQFH LQ VRLOV SULRU WR WKH LQWURGXFWLRQDQGH[WHQVLYHXVHRIFKHPLFDOIHUWLOL]HUV 7KH WK FHQWXU\ VFLHQWLVW SKLORVRSKHU DQG DOFKHPLVW$XUHROLV3KLOOLSXV7KHRSKUDVWXV %RPEDVWXV YRQ +RKHQKHLP 3DUDFHOVXV LQYHQWHG WKH QDPH ³1RVWRFK´3RWWV “Nostoch Pollution of some plethoricall and wanton Star, or rather excrement blown from the nostrills of some rheumatick planet” 3DUDFHOVXVLQ &KDUOWRQ 7KLV REVHUYDWLRQ E\ 3DUDFHOVXV LV QRW WKH RXWUDJHRXV VWDWHPHQWLWPD\ VHHP UDWKHULW KLG XQWLO UHFHQWO\ WKH FOXH WR WKH VRXUFH RI D ZRUG ZKLFK GHILHGHW\PRORJLFDODQDO\VLVIRU\HDUV1RVWRFKLV DSOD\RQWZRZRUGVE\DYHU\FOHYHUPDQDQ(QJOLVK ZRUG DQGD*HUPDQZRUG ZKLFKERWKGHVFULEHDSDUW RI WKH KXPDQ DQDWRP\ LQWLPDWHO\ DVVRFLDWHG ZLWK H[WUDFHOOXODUSRO\VDFFKDULGH 1RVWUK\O DQG1DVHQORFK QRVWULO 3RWWV ,Q IDFW RQH FDQ UHDGLO\ DSSUHFLDWHWKHSUREDEOH RULJLQRIWKH2OG'XWFKZRUG ³VQRW´ DYXOJDUHXSKHPLVPIRU QDVDO VHFUHWLRQ $V TXLWH FRUUHFWO\ SRLQWHG RXW E\ :ROIJDQJ .UXPEHLQ 3DUDFHOVXV ZDV WKH ILUVW WR UHFRJQL]H DQG VWXG\ ELRILOPV.UXPEHLQ ,WLVSHUKDSV LURQLF WKDWVRPH \HDUVDIWHUEHLQJ FODVVLILHG DV D SODQW LQ DQFLHQW &KLQD Nostoc, LQ ³HQOLJKWHQHG´ (XURSH ZDV UHOHJDWHG WR WKH UHDOP RI WKH VXSHUQDWXUDO DQGSHRSOHZHUH OHDG WR EHOLHYH WKDW “Nostoch understandeth the nocturnall” 3DUDFHOVXV &KDUOWRQ WUDQVODWLRQ (XURSHDQV ZHUH PRVW
&KDSWHU 1RVWRF
Table 1DWLRQDORULJLQVRIZRUGVXVHGWR GHVFULEH Nostoc commune *(50$1<
)5$1&(
81,7(' .,1*'20
6&$1',1$9,$
3+,/,33,1(6
6 WHPVFKQXSSHQ 6WLHPVFKRWW (UGEOXPH (UGJDOOHUW *ODVJDOOHUWH .XFNXFNVVSHLFKHO +LPPHOVEODWW +LPPHOVEOXPH 6FKOHLPOLQJ =LWWHUDOJH =LWWHUWDQJ
DUFKHH FHOHVW EHXUUH PDJLTXH FUDFKDW G¶XQ GLDEOH FUDFKDW GH OD OXQH FUDFKDW GH 0DL HFXPH SULQWLDUH IHXLOOH GX FLHO IOHXU GX VROHLO PHUGH GX FRXFRX VDOLYH GH FRXFRX
IDLULHV¶ EXWWHU IDOOHQ VWDUV ORFNOXEEHUW \ VFRRP VKRW VWDU VSLWWOH RIWKH VWDUV VWDUMHOO\ VWDU VKRW VWDUVOLPH VWDU VORXJK VWDUVOXEEHU ZLOOR¶WKHZLVS ZLWFKHV EXWWHU
WUROGVP¡U VN\IDOOVDOJHQ
EDWNLO EDNDWH EDOEDODG\RN NXOLQWDED WDEDWDED
FHUWDLQO\ FROOHFWLQJ Nostoc IRU VWXG\ DW WKLV WLPH DOWKRXJK ZH KDYH WR ZRQGHU ZKDW WKH\ ZHUH GRLQJ ZLWKLW DV DUHIHUHQFHLQDGLDU\FRQILUPV “a bottle of Nostock” 'H /D 3U\PH ,W WRRN D SURWUDFWHG DPRXQW RI WLPH IRU Nostoc WR DJDLQ EH DFFHSWHGDVDPHPEHURIWKHSODQWZRUOG
)URP%OXH*UHHQ$OJDWR &\DQREDFWHULXP 7KH KXPEOLQJ UHDOL]DWLRQ WKDW UDLQZDWHU OHDG WR WKH P\VWHULRXV DSSHDUDQFH RIN. commune LV DWWULEXWHG WR 5HQH $QWRLQH )HUFKDXOW GH 5HDPXU LQ DW D WLPH ZKHQ FODVVLFDO ERWDQ\ ZDV HQMR\LQJ LWV YRJXH 0ROOHQKDXHU D ,W LV KDUG WR EHOLHYH QHYHUWKHOHVV WKDW WKRVH SHRSOH ZKR ZHUH FROOHFWLQJ N. commune PXFKHDUOLHUWKDQ WKLV GDWH GLGQ¶WFDWFK RQ WR WKH UHK\GUDWLRQ SKHQRPHQRQ 7KH IRUP RIWKH N. commune PDFURVFRSLF FRORQ\ VWLOO KRZHYHU SRVHGSUREOHPVIRUWD[RQRPLVWVLQWKHVLWVWLOOLV WRGD\ ,Q Nostoc ZDVFRQVLGHUHGWREHDIRUPRI “Tremella,” DIXQJDOJHQXVZKRVHQDPHGHULYHVIURP WKHODWLQ tremere, PHDQLQJWRWUHPEOHRUTXLYHULQWKH VDPH IDVKLRQ WKDW WKH WXUJLG Nostoc FRORQ\ GRHV LI PRYHG 0ROOHQKDXHU D 6XFK VXSHUILFLDO VLPLODULWLHVWRIXQJLZHUHVXEVHTXHQWO\GLVFRXQWHGDQG Nostoc’s ZHOIDUH ZDV WDNHQ RYHU ILUVW E\ $OEUHFKW :LOKHOP 5RWK ZKR ILUVW SURYLGHG D GHILQLWLRQ RIWKH WHUP ³DOJDH´ DQG WKHQ PRUH DYLGO\ E\ D FOHUJ\PDQ DQG ERWDQLVW -HDQ 3LHUUH (WLHQQH 9DXFKHU ZKR LV FUHGLWHG ZLWK WKH ILUVW FOHDUFXWGHILQLWLRQRIWKHJHQXVNostoc 0ROOHQKDXHU DQG .RYiFLN
*XVWDYH$GROSKH 7KXUHW DQG -HDQ %DSWLVWH (GRXDUG %RUQHW SURYLGHG WKH VWDUWLQJSRLQWIRU D QHZ SKDVH LQ SK\FRORJ\ ZLWK WKH SXEOLFDWLRQ IURP WKURXJK RI WKHLU ³5HYLVLRQ GHV 1RVWRFDFpHV KpWpURF\VWpHV 0ROOHQKDXHU DQG .RYiFLN %RUQHW DQG )ODKDXOW 7KH 5HYLVLRQ IRUPV WKH ODWHU VWDUWLQJ SRLQW IRU WKH WD[RQRP\ RIWKH JURXS DV ZDV UDWLILHG E\ WKH ,QWHUQDWLRQDO %RWDQLFDO &RQJUHVV LQ 7KXUHWDQG%RUQHWFOHDUO\VKDUHG DSDVVLRQ IRU WKHLU ZRUN DQG WKLV SDLU RI SK\FRORJLVWV IRU PDQ\ \HDUV OLYHG DQG ZRUNHG WRJHWKHU DW 7KXUHW¶V 0HGLWHUUDQHDQ YLOOD DW $QWLEHV 7KH ILJXUHV DQG GUDZLQJV RINostoc VSS DQG RWKHU F\DQREDFWHULDWKDW DSSHDULQWKHLUSXEOLFDWLRQVDUHRXWVWDQGLQJDQG DUHRI WKHKLJKHVWTXDOLW\LPDJLQDEOH 'URXHW FODLPHG EHFDXVH WKH 5HYLVLRQ ZDV VR WRWDOO\ DFFHSWHG E\ ERWDQLVWV RI WKH WLPH WKDW SK\FRORJ\ DV D SURILWDEOH FDUHHU DWWUDFWHG QR GHGLFDWHG UHVHDUFKHUV GXULQJ WKH IROORZLQJ \HDUV Nostoc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
9,, (SLORJXH ,WLVILWWLQJE\ZD\RIDVXPPDU\ WRFRQILUPWKDWDQ DSSUHFLDWLRQ RI WKH HFRORJLFDO VLJQLILFDQFH DQG VXFFHVV RI Nostoc GLG QRW HVFDSH WKRVH RXWVLGH WKH VFLHQWLILF FRPPXQLW\ 7KH DYDQWJDUGH PXVLFDO JURXS =RYLHW)UDQFH XVHG WKH WLWOH ³*DWKHULQJ QRVWRF´IRUDSLHFHRIPXVLFRQWKHLUDOEXP/RK/DQG UHFRUGHGLQ$SULO LQ$PVWHUGDP=RYLHW)UDQFH %HQ 3RQWRQ WKH FRIRXQGHU RI WKH JURXS H[SODLQHG WKDW WKH XVH RI WKH ZRUG QRVWRF ZDV GHOLEHUDWH LW “included a sense of closeness to the earth, of belonging to and living within the biosphere, a sense that has in contemporary times become atrophied and displaced. Nostoc is a fine example of (an organism) living literally hand to mouth” %HQ3RQWRQSHUVFRPP
$FNQRZOHGJPHQWV 7KLVZRUNZDVPDGHSRVVLEOHE\WKH1DWLRQDO6FLHQFH )RXQGDWLRQ ,WKDQNDOOFROODERUDWRUVZKRFRQWULEXWHG WR VWXGLHV RQ Nostoc. 'LHWHU 0ROOHQKDXHU %HQ 3RQWRQ -LDQQ6KLQ &KHQ 7DQ&KL +XDQJ
5HIHUHQFHV $EGHODKDG 1 %D]]LFKHOOL * DQG 9HQWROD ) $Q HOHFWURQ PLFURVFRSLFLQYHVWLJDWLRQRQJUDQXOHVREVHUYHGLQWKHVKHDWKRI Nostoc commune 9$8&+ $UFK +\GURELRO6XSSO $JRVWLQL+-&DUUROO-'DQG0LQWRQ.: ,GHQWLILFDWLRQDQG FKDUDFWHUL]DWLRQ RI uvrA, D '1$ UHSDLU JHQH RI Deinococcus radiodurans. -%DFWHULRO $OHIRXQGHU 35 6$ %DOGZLQ 51 3HUKDP DQG 1- 6KRUW &ORQLQJ VHTXHQFHDQDO\VLV DQGRYHUH[SUHVVLRQRIWKHJHQHIRU WKH FODVV ,, IUXFWRVH ELVSKRVSKDWHDOGRODVH RI Escherichia coli. %LRFKHP- $OOHQ 00 &\DQREDFWHULDO FHOO LQFOXVLRQV $QQ 5HY 0LFURELRO $QDJQRVWLGLV DQG .RPiUHN 0RGHUQ DSSURDFK WR WKH FODVVLILFDWLRQ V\VWHP RI F\DQRSK\WHV 6WLJRQHPDWDOHV $UFK +\GURELRO6XSSO$OJRORJLFDO6WXGLHV $QGHUVRQ&5 -HQVHQ(2 /OHZHOO\Q '- 'HQQLV (6 DQG 3HDFRFN :- $ QHZ KHPRJORELQ JHQH IURP VR\EHDQ D UROH IRU KHPRJORELQ LQ DOO SODQWV 3URF 1DWO $FDG 6FL 86$ $UXJD < +DELWDW DQG GLVWULEXWLRQ RI ³IDFDL´ Nostoc flagelliforme &\DQRSK\WD -DS-3K\FRO6RUXL %D]]LFKHOOL * $EGHODKDG 1 DQG 9HQWROD ) 6WUXFWXUDO PRGLILFDWLRQV LQ WKH H[WUDFHOOXODU LQYHVWPHQW RI Nostoc
0DOFROP3RWWV commune 9DXFK GXULQJWKH OLIH F\FOH ,0RWLOHDQGQRQPRWLOH KRUPRJRQLXPELVHULDWHVWDJH-8OWUDVWUXF5HV %D]]LFKHOOL * $EGHODKDG 1 DQG 9HQWROD ) 6WUXFWXUDO PRGLILFDWLRQV LQ WKH H[WUDFHOOXODU LQYHVWPHQW RI Nostoc commune 9DXFK GXULQJ WKH OLIH F\FOH OO 0LFURFRORQLHV DQG ³EDOOV´RIILODPHQWV-8OWUDVWUXF0RO6WUXF5HV %HHVOH\&( 6PLWK5-7HPSOH6-DQG/HD 3- &ORQLQJDQG QXFOHRWLGH VHTXHQFH RI WKH JHQH HQFRGLQJ GLQLWURJHQDVH UHGXFWDVH nifH IURP WKH F\DQREDFWHULXP Nostoc %LRFKHP%LRSK\V$FWD %HUQDUG ) 'HQVLW\ RIIODJHOODWHV DQG 0\[RSK\FHDH LQ WKH KHWHURWURSKLF OD\HUV UHODWHG WR HQYLURQPHQW ,Q 2SHQKHLPHU & HG 6\PSRVLXPRQ0DULQH0LFURELRORJ\SS &KDUOHV &7KRPDV6SULQJILHOG,OOLQRLV %HUQDUG ) DQG /HFDO - 3ODQFWRQ XQLFHOOXODLUH UHFROWp GDQV ¶2FHDQ ,QGLHQ SDU OH &KDUFRW HW OH 1RUVHO %XOO,QVW2FHDQRJU0RQDFR %LOJHU : %LLGHO % 0ROOHQKDXHU 5 DQG 0ROOHQKDXHU ' 3KRWRV\QWKHWLF DFWLYLW\RIWZRGHYHORSPHQWDO VWDJHVRIDNostoc VWUDLQ &\DQREDFWHULD LVRODWHG IURP Geosiphon pyriforme 0\FRWD -3K\FRO %RE]LQ 6& DQG 0RRUH 5( %LRV\QWKHWLF RULJLQ RI F\FORSKDQHV IURP F\DQREDFWHULD 7HWUDKHGURQ /HWW %RKP*$3IOHLGHUHU: %RJHU3DQG6FKHUHU6 6WUXFWXUH RI D QRYHO ROLJRVDFFKDULGHP\FRVSRULQHDPLQR DFLG XOWUDYLROHW $% VXQVFUHHQ SLJPHQW IURP WKH WHUUHVWULDO F\DQREDFWHULXP Nostoc commune. -%LRO&KHP %RUQHW e DQG )ODKDXOW & 5HYLVLRQ GHV 1RVWRFDFpHV KpWpURF\VWpHVFRQWHQXHVGDQVOHVSULQFLSDX[KHUELHUVGH)UDQFH $QQ 6FL 1DW%RW %URDG\3$ (FRORJ\DQGWD[RQRP\RIWKHWHUUHVWULDODOJDHRI WKH 9HVWIROG +LOOV ,Q 3LFNDUG - HG $QWDUFWLF 2DVLV SS $FDGHPLF3UHVV %LLGHO % :HVVHOV '&- DQG 0ROOHQKDXHU ' 0DVVHQHQWZLFNOXQJ YRQ Nostoc FI microscopicum &DUPLFKDHO +DUYH\ H[ %RPHW )ODKDXOW LQ DONDOLVFKHP VDO]KDOWLJHP 0LOLHX HLQHU +RKOH LQ GHQ 'UDNHQVEHUJHQ 6XGDIULNDV *ROGHQ *DWH +LJKODQGV 1DWLRQDO 3DUN $UFK 3URWLVWHQNG %XLNHPD :- *XR < =XWKHU ( +DJHPDQQ 0 DQG +DVHONRP 5 6XFURVH V\QWKDVH LQ Anabaena LV UHTXLUHG IRU KHWHURF\VW IXQFWLRQ $EV 9,WK &\DQREDFWHULDO :RUNVKRS $VLORPDU % %XLNHPD :- DQG +DVHONRUQ 5 &KDUDFWHUL]DWLRQ RID JHQH FRQWUROOLQJ KHWHURF\VW GLIIHUHQWLDWLRQ LQ WKH F\DQREDFWHULXP Anabaena *HQHV'HYHO %XLNHPD :- DQG +DVHONRUQ 5 0ROHFXODU JHQHWLFV RI F\DQREDFWHULDOGHYHORSPHQW $QQX5HY3ODQW3K\VLRO3ODQW0RO %LRO %XUNH '+ +HDUVW -( DQG 6LGRZ $ (DUO\ HYROXWLRQ RI SKRWRV\QWKHVLV FOXHV IURP QLWURJHQDVH DQG FKORURSK\OO LURQ SURWHLQV 3URF1DWO$FDG6FL86$ &DPSEHOO (/ &RKHQ 0) DQG 0HHNV -& $ SRO\NHWLGH V\QWKDVH OLNH JHQH LV LQYROYHG LQ WKH V\QWKHVLV RI KHWHURF\VW JO\FROLSLGV LQ Nostoc punctiforme VWUDLQ $7&& $UFK 0LFURELRO &DPSEHOO(/+DJHQ.' &RKHQ0)6XPPHUV0/DQG0HHNV-& 7KH GHY5 JHQH SURGXFW LV FKDUDFWHULVWLF RIUHFHLYHUV RI WZR FRPSRQHQW UHJXODWRU\ V\VWHPV DQG LV HVVHQWLDO IRU KHWHURF\VW GHYHORSPHQW LQ WKH ILODPHQWRXV F\DQREDFWHULXP Nostoc VSVWUDLQ$7&&-%DFWHULRO
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scillatoria rubescens '& $UFK +\GURELRO &KDUOWRQ 9DQ+HOPRQW¶VSDUDGR[HV 7UDQVO6XSSO &KD]DO '( DQG 6PLWK *' &KDUDFWHUL]DWLRQ RI D EURZQ Nostoc VSHFLHV IURP-DYDWKDWLVUHVLVWDQWWRKLJKOLJKWLQWHQVLW\ DQG890LFURELRORJ\ &KHQ -+ *LEVRQ -/ 0F&XH /$ DQG 7DELWD )5 ,GHQWLILFDWLRQ H[SUHVVLRQ DQG GHGXFHG SULPDU\ VWUXFWXUH RI WUDQVNHWRODVH DQG RWKHU HQ]\PHV HQFRGHG ZLWKLQ WKH IRUP ,, &2 IL[DWLRQ RSHURQ RIRhodobacter sphaeroides. - %LRO &KHP &KLDIIUHGR 0 DQG )LJXUHDX & 6XEVWUDWH FRQWDLQLQJ &\DQRSK\FHDH DQG %U\RSK\WH SURWRQHPDV IRU SURGXFLQJ YHJHWDWLRQRQEDUHWHUUDLQ8QLWHG6WDWHV3DWHQW &OHJJ -6 7KH SK\VLFDO SURSHUWLHV DQG PHWDEROLF VWDWXV RI Artemia F\VWV DW ORZ ZDWHU FRQWHQWV 7KH ³ZDWHU UHSODFHPHQW K\SRWKHVLV´,Q/HRSROG$&HG 0HPEUDQHVPHWDEROLVPDQG GU\ RUJDQLVPV SS &RPVWRFN 3XEOLVKLQJ $VVRFLDWHV ,WKDFDDQG/RQGRQ &RKDQ )0 7KH UROH RI JHQHWLF H[FKDQJH LQ EDFWHULDO HYROXWLRQ$601HZV &RKHQ 0) DQG 0HHNV -& $ KRUPRJRQLXP UHJXODWLQJ ORFXV hrmUA, RIWKHF\DQREDFWHULXPNostoc punctiforme VWUDLQ $7&&DQGLWVUHVSRQVHWRDQH[WUDFWRIDV\PELRWLFSODQW SDUWQHU$QWKRFHURV SXQFWDWXV 0RO 3ODQW0LFUREH ,QW &RKHQ 0) :DOOLV -* &DPSEHOO (/ DQG 0HHNV -& 7UDQVSRVRQ PXWDJHQHVLV RINostoc VS VWUDLQ $7&& D ILODPHQWRXV F\DQREDFWHULXP ZLWK PXOWLSOH FHOOXODU GLIIHUHQWLDWLRQDOWHUQDWLYHV0LFURELRORJ\ &RUEHWW // DQG 3DUNHU '/ 9LDELOLW\ RI O\RSKLOL]HG F\DQREDFWHULD EOXHJUHHQ DOJDH $SSO (QYLURQ 0LFURELRO &XSDF 6 DQG *DQWDU 0 &KHPLFDO FRPSRVLWLRQ DQG PRUSKRORJLFDO VWUXFWXUH RI WKH PXFLODJLQRXV VKHDWK RI WKH F\DQREDFWHULXP Nostoc '%LRPHG/HWW &XUWLV 6( DQG 0DUWLQ -$ 7KH WUDQVFULSWLRQ DSSDUDWXV DQG WKH UHJXODWLRQ RIWUDQVFULSWLRQDO LQLWLDWLRQ ,Q %U\DQW '$ HG 7KH 0ROHFXODU%LRORJ\ RI&\DQREDFWHULD SS .OXZHU $FDGHPLF3XEOLVKHUV7KH1HWKHUODQGV 'DO\ 0- DQG 0LQWRQ .: D ,QWHUFKURPRVRPDO UHFRPELQDWLRQ LQ WKH H[WUHPHO\ UDGLRUHVLVWDQW EDFWHULXP Deinococcus radiodurans. -%DFWHULRO 'DO\0-DQG0LQWRQ.:E 5HVLVWDQFHWRUDGLDWLRQ 6FLHQFH 'DO\ 0- DQG 0LQWRQ .: $Q DOWHUQDWLYH SDWKZD\ RI UHFRPELQDWLRQ RI FKURPRVRPDO IUDJPHQWV SUHFHGHV recAGHSHQGHQW UHFRPELQDWLRQ LQ WKH UDGLRUHVLVWDQW EDFWHULXP Deinococcus radiodurans. -%DFWHULRO 'H/D3U\PH /HW,Q'LDU\6XUWHHV S 'HVLNDFKDU\ 79 &\DQRSK\WD ,QGLDQ &RXQFLO RI $JULFXOWXUDO5HVHDUFK1HZ'HOKL
'RGGV:.DQG&DVWHQKRO]5: (IIHFWVRIJUD]LQJDQGOLJKW RQWKHJURZWK RINostoc pruniforme &\DQREDFWHULD %U3K\FRO - 'RGGV:.*XGGHU'$DQG0ROOHQKDXHU' 7KHHFRORJ\RI Nostoc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¶$QWRQLR 7KH HIIHFWV RIVXEVWUDWH WH[WXUH JUD]LQJ DQG GLVWXUEDQFH RQ PDFURDOJDO HVWDEOLVKPHQW LQ VWUHDPV(FRORJ\ (KOLQJ6KXO] 0 %LOJHU : DQG 6FKHUHU 6 89%LQGXFHG V\QWKHVLV RI SKRWRSURWHFWLYH SLJPHQWV DQG H[WUDFHOOXODU SRO\VDFFKDULGHV LQ WKH WHUUHVWLDO F\DQREDFWHULXP Nostoc commune. -%DFWHULRO (PUDOLQR *$ DQG 5RGXOIR %5 &XOWXUDO VWXGLHV RQ EOXH JUHHQ DOJDH ,,, Nostoc microscopicum &DUPLFKDHO 3KLOOLS - 6FL (UQVW$&KHQ7:DQG%RJHU3 &DUERK\GUDWHIRUPDWLRQLQ UHZHWWHGWHUUHVWULDOF\DQREDFWHULD2HFRORJLD )DJHUEDNNH .0 +HOGDO 0 DQG 1RUODQG 6 9DULDWLRQ LQ HOHPHQWDO FRQWHQW DPRQJ DQG ZLWKLQ WULFKRPHV LQ Nostoc calcicola :$ PHDVXUHG E\ [UD\ PLFURDQDO\VLV )(06 0LFURELRO/HWW )DYDOL 0$ DQG &DLROD 0* $XWRUDGLRJUDSK\ DQG XOWUDVWUXFWXUDO VWXG\ RQ Nostoc VSHFLHV ODEHOHG ZLWK ¶+ WK\PLGLQH0LFURELRV )HQJ ') &KRS * DQG 'RROLWWOH 5) 'HWHUPLQLQJ GLYHUJHQFH WLPHV ZLWK D SURWHLQ FORFNXSGDWH DQG UHHDYOXDWLRQ 3URF1DWO$FDG6FL86$ )HUQDQGH]SLQDV ) 0DWHR 3 DQG %RQLOOD , 8OWUDVWUXFWXUDO FKDQJHV LQGXFHG E\ VHOHFWHG FDGPLXP FRQFHQWUDWLRQV LQ WKH F\DQREDFWHULXP Nostoc 8$0-3ODQW3K\VLRO )MHUGLQJVWDG ( )MHUGLQJVWDG ( DQG )MHUGLQJVWDG (- (YLGHQFH IRUWKH SUHVHQFH RIFROODJHQ LQ WKH VKHDWKV RIDEOXH JUHHQ DOJD DQ HOHFWURQ PLFURVFRSLFDO DQG KLVWRFKHPLFDO VWXG\ RI Scytonema myochrous VWDWXV SHWDORQHPDWRLGHV $UFK +\GURELRO )R[*(:LVRW]NH\-'DQG-XUWVKXN3 +RZFORVHLV FORVH 6 U51$VHTXHQFHLGHQWLW\PD\QRWEHVXIILFLHQWWRJXDUDQWHH VSHFLHVLGHQWLW\,QW-6\VW%DFWHULRO )UDOHLJK 3& DQG %XUQKDP -& 0\[RFRFFDO SUHGDWLRQ RQ F\DQREDFWHULDO SRSXODWLRQV QXWULHQW HIIHFWV /LPQRO 2FHDQRJU )UHQFK- &K\PQ'LFW *DQWDU0 .HUE\1:DQG5RZHOO3 &RORQL]DWLRQ RIZKHDW (Triticum vulgare / E\ 1IL[LQJ F\DQREDFWHULD $Q XOWUDVWUXFWXUDOVWXG\1HZ3K\WRO *DR . &KLQHVH VWXGLHV RQ WKH HGLEOH EOXHJUHHQ DOJD Nostocflagelliforme: DUHYLHZ-$SSO3K\FRO *DR . 4LX% ;LD -<X $DQG/L< (IIHFWRIZLQGVSHHG RQ ORVV RI ZDWHU IURP Nostoc flagelliforme FRORQLHV - $SSO 3K\FRO *HLWOHU / &\DQRSK\FHDH ,Q .RONZLW] HG 'U / 5DEHQKRUVW¶V .U\SWRJDPHQIORUD YRQ 'HXWVFKODQG gVWHUUHLFK XQGGHU6FKZHL]$NDGHPLVFKH9HUODJVJHVHOOVFKDIW/HLS]LJ
*HLWOHU/ =XU .HQQWQLV GHU2VFLOODWRULDFHH Pseudanabaena galeata &\DQRSK\WD 3ODQW6\VWDQG(YRO *LEVRQ -/ )DOFRQH '/ DQG 7DELWD )5 1XFOHRWLGH VHTXHQFH WUDQVFULSWLRQDO DQDO\VLV DQG H[SUHVVLRQ RI JHQHV HQFRGHG ZLWKLQ WKH IRUP ,&2 IL[DWLRQ RSHURQ RIRhodobacter sphaeroides. -%LRO&KHP *LEVRQ-/DQG7DELWD)5 /RFDOL]DWLRQDQGPDSSLQJRI&2 IL[DWLRQ JHQHV ZLWKLQ WZR JHQH FOXVWHUV LQ Rhodobacter sphaeroides. -%DFWHULRO *LOOHV*RQ]DOH] 0$ 'LWWD *6 DQG +HOLQVNL '5 $ KDHPRSURWHLQ ZLWK NLQDVH DFWLYLW\ HQFRGHG E\ WKH R[\JHQ VHQVRURIRhizobium meliloti. 1DWXUH *LRYDQQRQL 6- 7XUQHU 6 2OVHQ *- %DUQV 6 /DQH '- DQG 3DFH 15 (YROXWLRQDU\UHODWLRQVKLSV DPRQJF\DQREDFWHULDDQG JUHHQFKORURSODVWV-%DFWHULRO *RODNRWL72JLQR- +HOW]HO&( +XVHER7/-HQVHQ&0 /DUVHQ /.3DWWHUVRQ*0/0RRUH5(0RREHUU\6/ &RUEHWW7+DQG 9DOHULRWH )$ 6WUXFWXUH GHWHUPLQDWLRQ FRQIRUPDWLRQDO DQDO\VLV FKHPLFDOVWDELOLW\VWXGLHV DQGDQWLWXPRUHYDOXDWLRQ RI WKHFU\SWRSK\FLQV ,VRODWLRQRI QHZDQDORJVIURPNostoc VS VWUDLQ*69-$P&KHP6RF *RXOG6-DQG(OGUHGJH1 3XQFWXDWHGHTXLOLEULXPFRPHVRI DJH1DWXUH *URLVPDQ ($ 6DLHU -U 0+ DQG 2FKPDQ + +RUL]RQWDO WUDQVIHURIDSKRVSKDWDVHJHQHDVHYLGHQFH IRUPRVDLF VWUXFWXUH RIWKHSalmonella JHQRPH(0%2- *URVVPDQ $5 6FKDHIHU 05 &KLDQJ ** DQG &ROOLHU -/ 7KH UHVSRQVHV RI F\DQREDFWHULD WR HQYLURQPHQWDO FRQGLWLRQV OLJKWDQGQXWULHQWV ,Q %U\DQW'$HG 7KH0ROHFXODU%LRORJ\ RI &\DQREDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 1HWKHUODQGV *XSWD $ DQG 6LQJKDO *6 &KDQJHV LQ XOWUDVWUXFWXUH DQG LQKLELWLRQ RIHOHFWURQWUDQVSRUW DFWLYLW\ LQGXFHG E\ 7KDOOLXP LQ Nostoc muscorum. 3KRWRV\QWKHWLFD *XSWD 5. DQG .DVK\DS $. 8SWDNH DQG ORVV RIZDWHU E\ $QWDUFWLF F\DQREDFWHULXP Nostoc commune XQGHU ILHOG DQG ODERUDWRU\FRQGLWLRQV3K\NRV *XSWD56%XVWDUG.)DODK0DQG6LQJKW' 6HTXHQFLQJRI KHDW VKRFN SURWHLQ 'QD. KRPRORJV IURP Deinococcus proteolyticus DQG Thermomicrobium roseum DQG WKHLU LQWHJUDWLRQ LQ D SURWHLQEDVHG SK\ORJHQ\ RI SURNDU\RWHV - %DFWHULRO *XUHYLW] 0 2VLHZDF] +' DQG .HUHQ < 0ROHFXODU HYLGHQFH IRU LQWHUFKURPRVRPDO UHFRPELQDWLRQ LQ WKH F\DQREDFWHULXP 6\QHFKRF\VWLV VS 3&& 3ODQW 6FL +DELFKW.6 DQG&DQILHOG'( 6XOSKXULVRWRSHIUDFWLRQDWLRQ LQ PRGHP PLFURELDO PDWV DQG WKH HYROXWLRQ RI WKH VXOSKXU F\FOH1DWXUH +DJHPDQQ0*ROOGDFN'%LJJLQV-DQG(UGPDQQ1 6DOW GHSHQGHQW SURWHLQ SKRVSKRU\ODWLRQ LQ WKH F\DQREDFWHULXP Synechocystis 3&&)(060LFURELRO/HWW +DJHQ .' DQG 0HHNV -& *HQHWLF HYLGHQFH IRU WKH LQYROYHPHQWRIDSKRVSKRUHOD\V\VWHP LQKHWHURF\VWPDWXUDWLRQ LQ Nostoc punctiforme. $EV 9,WK &\DQREDFWHULDO :RUNVKRS $VLORPDU% +DUGLVRQ5& $EULHIKLVWRU\RIKHPRJORELQV3ODQWDQLPDO SURWLVWDQGEDFWHULD3URF1DWO$FDG6FL86$ +DVHJDZD 0 DQG )LWFK :0 'DWLQJ WKH FHQDQFHVWHU RI RUJDQLVPV6FLHQFH +DVHONRP 5 'HYHORSPHQWDOO\ UHJXODWHG JHQH UHDUUDQJHPHQWVLQSURNDU\RWHV$QQX5HY*HQHW
0DOFROP3RWWV +DZHV ,+RZDUG:LOOLDPV&DQG9LQFHQW:) 'HVLFFDWLRQ DQG UHFRYHU\ RI $QWDUFWLF F\DQREDFWHULDO PDWV 3RODU %LRO +HPVFKHLGW 7 3XJOLVL 03 /DUVHQ /. 3DWWHUVRQ *0/ 0RRUH 5( 5LRV -/DQG &ODUG\- 6WUXFWXUHDQG ELRV\QWKHVLV RI ERURSK\FLQ D QHZ %RHVHNHQ FRPSOH[ RI ERULF DFLG IURP D PDULQHVWUDLQRIWKHEOXHJUHHQDOJDNostoc linckia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ostoc VSS-%DFWHULRO +LOO '5 +ODGXQ + 6FKHUHU 6 DQG 3RWWV 0 D :DWHU VWUHVV SURWHLQV RINostoc commune &\DQREDFWHULD DUH VHFUHWHG ZLWK 89$%DEVRUELQJ SLJPHQWV DQG DVVRFLDWH ZLWK E' [\ODQ[\ODQRK\GURODVHDFWLYLW\-%LRO&KHP +LOO '5 .HHQDQ 7: +HOP 5) 3RWWV 0 / 0 &URZH DQG -+ &URZH([WUDFHOOXODUSRO\VDFFKDULGHRINostoc FRPPXQH &\DQREDFWHULD LQKLELWV IXVLRQ RI PHPEUDQH YHVLFOHV GXULQJ GHVLFFDWLRQ -$SSO3K\FRO +LOO '5 3HDW $ DQG 3RWWV 0 E %LRFKHPLVWU\ DQG VWUXFWXUH RIWKHJO\FDQVHFUHWHGE\GHVLFFDWLRQWROHUDQW Nostoc FRPPXQH &\DQREDFWHULD 3URWRSODVPD +LUDWD.1DNDJDPL+7DNDVKLQD-0DKPXG7.RED\DVEL 0 ,Q < ,VKLGD 7 DQG 0L\DPRWR . $ QRYHO YLROHW SLJPHQW QRVWRFLQH $ DQ H[WUDFHOOXODU PHWDEROLWH IURP F\DQREDFWHULXP Nostoc spongiaeforme. +HWHURF\FOHV +LUVFK &) /LHVFK-0 6DOYDWRUH 0- 6FKZDUW] 5(DQG 6HVLQ ') $QWLIXQJDO IHUPHQWDWLRQ SURGXFW DQG PHWKRG 8QLWHG 6WDWHV3DWHQW +LUVFK5+DUWXQJ:DQG*LPPOHU+ $EVFLVLFDFLGFRQWHQW RIDOJDHXQGHUVWUHVV %RW$FWD +RGJHV &) D ,QWHUDFWLRQ RI F\DQREDFWHULD DQG VXOIDWH UHGXFLQJ EDFWHULD LQ VXEVXUIDFH EODFNOD\HU IRUPDWLRQ LQ KLJK VDQGFRQWHQWJROIJUHHQV6RLO%LRO%LRFKHP +RGJHV &) E 3DWKRJHQLFLW\ RI Pythium torulosum WR URRWV RI Agrostis palustris LQ EODFNOD\HUHG VDQG SURGXFHG E\ WKH LQWHUDFWLRQ RIWKH F\DQREDFWHULD VSHFLHV Lyngbya, Phormidium, DQG Nostoc ZLWK Desulfovibrio desulfuricans. &DQ - %RW +ROODQG'DQG:RON&3 ,GHQWLILFDWLRQDQGFKDUDFWHUL]DWLRQ RI hetA, D JHQH WKDW DFWV HDUO\ LQ WKH SURFHVV RIPRUSKRORJLFDO GLIIHUHQWLDWLRQRIKHWHURF\VWV-%DFWHULRO +RVHWWL %% DQG 3DWLO +6 ,PSDFW RI JURZWK RI Nostoc punctiforme DQG Escherichia coli RQ SRO\ILEUH ZDVWHZDWHU - (QYLURQ%LRO +RXJK / -RQHV /.1 DQG :DGPDQ :+ $Q LQYHVWLJDWLRQ RI WKH SRO\VDFFKDULGH RI FRPSRQHQWV RI FHUWDLQ IUHVKZDWHU DOJDH-&KHP6RF +RZHOO /' *ULIILWKV & 6ODGH /: 3RWWV 0 DQG .HQQHOO\ 3- 6XEVWUDWH VSHFLILFLW\ RI ,SK3 D F\DQREDFWHULDO GXDO VSHFLILFLW\ SURWHLQ SKRVSKDWDVH ZLWK 0$3 NLQDVH SKRVSKDWDVH DFWLYLW\%LRFKHPLVWU\ +X 7/ .HDQ &. DQG /HRQJ 26 2GRURXV SUREOHP DVVRFLDWHG ZLWK Nosroc VS LQ ZDWHU VXSSO\ V\VWHP RI FHQWUDO 7DLZDQ ,QW&RQI$VLDQ:DWHU7HFK
&KDSWHU 1RVWRF +XDQJ= /LX < 3DXOVHQ 63DQG .ODYHQHVV 'VXEPLWWHG 6WXGLHV RQ SRO\VDFFKDULGHV IURP WKUHH VSHFLHV RI HGLEOH Nostoc &\DQRSK\WD &\DQREDFWHULD ZLWK GLIIHUHQW FRORQ\ PRUSKRORJLHV &RPSDULVRQ RI PRQRVDFFKDULGH FRPSRVLWLRQ DQGYLVFRVLW\EHWZHHQILHOGFRORQLHVDQGVXVSHQVLRQFXOWXUHV +XGGDUW + 6PLWK 5- /DQJWRQ 3' +HWKHULQJWRQ $0 DQG 0DQVILHOG 7$ ,V DEVFLVLF DFLG D XQLYHUVDOO\ DFWLYH FDOFLXPDJRQLVW"1HZ3K\WRO ,NHKDUD.$PDGD)
.HQQHOO\ 3- DQG 3RWWV 0 )DQF\PHHWLQJ\RXKHUH DIUHVK ORRN DW ³SURNDU\RWLF´ SURWHLQ SKRVSKRU\ODWLRQ - %DFWHULRO .KXG\DNRY , DQG :RON &3 (YLGHQFH WKDW WKH hanA JHQH FRGLQJIRU+8 SURWHLQLV HVVHQWLDO IRUKHWHURF\VWGLIIHUHQWLDWLRQ LQ DQG F\DQRSKDJH $/ VHQVLWLYLW\ RI Anabaena VS VWUDLQ 3&&-%DFWHULRO .OHLQKDXV 6 DQG .DLVHU $' (FRORJ\ DQG ELRPHFKDQLFDO FRQVHTXHQFHVRIOLYLQJWRJHWKHU ,QGXFHG PRUSKRORJLFDOFKDQJH LQDNostoc DQG Cricotopus V\PELRVLV $P=RRO$ .RFK$/ :KDWVL]HVKRXOGDEDFWHULXPEH" $TXHVWLRQ RI VFDOH$QQX5HY0LFURELRO .RPi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pV ) *HQHWLF HYLGHQFHWKDW KHS$JHQH LV LQYROYHG LQ WKH QRUPDO GHSRVLWLRQ RIWKH HQYHORSH RI ERWK KHWHURF\VWV DQG DNLQHWHV LQ Anabaena variubilis $7&& )(06 0LFURELRO/HWW /HVOLH -) *UHDVKDP 5/ DQG +XOEHUW 0+ %LRORJLFDO IORFFXODWLRQRIPLQHUDOVOLPHV 8QLWHG6WDWH3DWHQW /L 6 DQG:DWHUV 5 Escherichia coli VWUDLQV ODFNLQJSURWHLQ +8 DUH 89 VHQVLWLYH GXH WR D UROH IRU +8 LQ KRPRORJRXV UHFRPELQDWLRQ-%DFWHULRO /L 6& 3HQ 7V¶DR .DQJ 0X &KLQHVH 3KDUPDFHXWLFDO &RPSHQGLXP UHSULQWHG E\ 3HRSOH¶V 0HGLFDO 3XEOLVKLQJ+RXVH %HLMLQJ /WWJH 8 &\DQREDFWHULDO Tintenstrich FRPPXQLWLHV DQG WKHLUHFRORJ\1DWXQYLVVHQVFKDIWHQ /\QQ 0( %DQWOH -$ DQG 2ZQE\ -' (VWLPDWLRQ RIJHQH H[SUHVVLRQ LQ KHWHURF\VWV RI Anabaena variabilis E\ XVLQJ '1$51$K\EULGL]DWLRQ-%DFWHULRO 0DQQ 1+ 3URWHLQ SKRVSKRU\ODWLRQ LQ F\DQREDFWHULD 0LFURELRORJ\ 0DQQ 1+ 5LSSND 5 DQG +HUGPDQQ 0 5HJXODWLRQ RI SURWHLQSKRVSKRU\ODWLRQ LQ WKHF\DQREDFWHULXP Anabaena VWUDLQ 3&&-*HQ0LFURELRO 0DUVDOHN% =DKUDGQtFNRYiDQG +URQNRYi0 ([WUDFHOOXODU DEVFLVLF DFLG SURGXFHG E\ F\DQREDFWHULD XQGHU VDOW VWUHVV - 3ODQW3K\VLRO 0DUVK--DQG/HEKHU]+* )UXFWRVHELVSKRVSKDWHDOGRODVHV DQHYROXWLRQDU\KLVWRU\7,%6 0DUWLQH] 0 Nostoc commune 9DXFK D QLWURJHQIL[LQJ EOXHJUHHQ DOJD DV VRXUFH RI IRRG LQ WKH 3KLOLSSLQHV 3KLO $JULFXO 0DUWLQH]05 DQG 4XHULMHUR 10% 0DFURFRORQ\ IRUPDWLRQ LQ WKH QLWURJHQIL[LQJ EOXHJUHHQ DOJD Nostoc FRPPXQH 9$8&+3KLO$JUL 0DUWLQH] 05 4XHULMHUR 10% DQG 9LOODUHDO&DVWLOOR / 3KRWRFRQWUROOHG JURZWK QLWURJHQIL[DWLRQ DQG PRUSKRORJLFDO
GHYHORSPHQWDOOLIH1RVWRFFRPPXQH 3KLO- 6FL 0RQRJUDSK 0DVHSRKO % *RUOLW] . DQG %RKP + /RQJ WDQGHPO\ UHSHDWHG UHSHWLWLYH /755 VHTXHQFHV LQ WKH ILODPHQWRXV F\DQREDFWHULXP $QDEDHQD VS 3&& %LRFKLP %LRSK\V $FWD*HQH6WUXFWXUHDQG([SUHVVLRQ 0DWWLPRUH 9 DQG %DWWLVWD -5 5DGLRUHVLVWDQFH RI 'HLQRFRFFXV UDGLRGXUDQV IXQFWLRQV QHFHVVDU\ WR VXUYLYH LRQL]LQJ UDGLDWLRQ DUH DOVR QHFHVVDU\ WR VXUYLYH SURORQJHG GHVLFFDWLRQ-%DFWHULRO 0F$UWQH\%+RZHOO' .HQQHOO\3-DQG3RWWV 0 3URWHLQ W\URVLQH SKRVSKRU\ODWLRQ LQ $QDEDHQD VS 3&& &\DQREDFWHULD -%DFWHULRO 0HO QLNRYD 99 5HVSRQVH RI SRLNLOR[HURSKLOLF EOXHJUHHQ ZDWHU SODQWV 1RVWRF FRPPXQH 9DXFK WR GHK\GUDWLRQ 7VWLWRORJL\D 0LQWRQ .: 5HSDLU RI LRQL]LQJUDGLDWLRQ GDPDJH LQ WKH UDGLDWLRQUHVLVWDQW EDFWHULXP 'HLQRFRFFXV UDGLRGXUDQV 0XW 5HV 0LQWRQ.:DQG'DO\ 0- $PRGHO IRUUHSDLURIUDGLDWLRQ LQGXFHG '1$ GRXEOHVWUDQG EUHDNV LQ WKH H[WUHPH UDGLRSKLOH 'HLQRFRFFXVUDGLRGXUDQV %LR(VVD\V 0RHQV / 9DQIOHWHUHQ - 9DQ GH 3HHU < 3HHWHUV . .DSS &]HOX]QLDN-*RRGPDQ0%OD[WHU0DQG9LQRJUDGRY6 *ORELQV LQ QRQYHUWHEUDWH VSHFLHV 'LVSHUVDO E\ KRUL]RQDO JHQH WUDQVIHU DQG HYROXWLRQ RI WKH VWUXFWXUHIXQFWLRQ UHODWLRQVKLSV 0RO%LRO(YRO 0RM]VLV 6- $UUKHQLXV * 0F.HHJDQ .' +DUULVRQ 70 1XWPDQ $3 DQG )ULHQG &5/ (YLGHQFH IRU OLIH RQ (DUWK EHIRUH PLOOLRQ\HDUVDJR1DWXUH 0ROOHQKDXHU' %HLWUDJH ]XU.HQQWQLV GHU*DWWXQJ1RVWRF $EKVHQFNHQEQDWXUIRUVFK*H 0ROOHQKDXHU'D %ODXDOJHQGHU*DWWXQJ1RVWRF LKUH5ROOH LQ )RUVFKXQJ XQG :LVVHQVFKDIWVJHVFKLFKWH , 1DWXU XQG 0XVHXP 0ROOHQKDXHU'E %ODXDOJHQGHU*DWWXQJ1RVWRF LKUH5ROOH LQ )RUVFKXQJ XQG :LVVHQVFKDIWVJHVFKLFKWH ,, 1DWXU XQG 0XVHXP 0ROOHQKDXHU'D %ODXDOJHQGHU*DWWXQJ1RVWRF LKUH5ROOH LQ )RUVFKXQJ XQG :LVVHQVFKDIWVJHVFKLFKWH ,,, 1DWXU XQG 0XVHXP 0ROOHQKDXHU'E %ODXDOJHQGHU*DWWXQJ1RVWRF LKUH5ROOH LQ )RUVFKXQJ XQG :LVVHQVFKDIWVJHVFKLFKWH ,9 1DWXU XQG 0XVHXP 0ROOHQKDXHU ' DQG .RYi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
0DOFROP3RWWV 1HOLVVHQ % 'H %DHUH 5 :LOPRWWH $ DQG 'H :DFKWHU 5 3K\ORJHQHWLF UHODWLRQVKLSV RI QRQD[HQLF ILODPHQWRXV F\DQREDFWHULDOVWUDLQVEDVHGRQ 6U51$ VHTXHQFHDQDO\VLV - 0RO(YRO 2KPRUL. +LURVH0DQG2KPRUL 0 )XQFWLRQRIF$03DV DPDWIRUPLQJIDFWRULQ WKH F\DQREDFWHULXP 6SLUXOLQDSODWHQVLV 3ODQW&HOO3K\VLRO 2KPRUL 0 F$03 LQ $QDEDHQD F\OLQGULFD UDSLG FKDQJHV LQ FHOOXODU OHYHOV LQ UHVSRQVH WR FKDQJHV LQ H[WUDFHOOXODU HQYLURQPHQWV3ODQW&HOO3K\VLRO 2NLQR 7 4L 6 0DWVXGD + 0XUDNDPL 0 DQG
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ostoc commune 3K\VLRO 3ODQW 5RGULJXH] 5 *DUFLD*RQ]iOH] 0 *XHUUHUR 0* DQG /DUD & $PPRQLXPVHQVLWLYH SURWHLQ NLQDVH DFWLYLW\ LQ SODVPD PHPEUDQHV RI WKH F\DQREDFWHULXP Anacystis nidulans. )(%6 /HWW 5XGL . 6NXOEHUJ 20 DQG -DNREVHQ .6 (YROXWLRQ RI F\DQREDFWHULD E\ H[FKDQJH RI JHQHWLF PDWHULDO DPRQJ SK\OHWLFDOO\UHODWHGVWUDLQV-%DFWHULRO 5XGL . 6NXOEHUJ 20 /DUVHQ )DQG -DNREVHQ .6 6WUDLQ FKDUDFWHUL]DWLRQ DQG FODVVLILFDWLRQ RIR[\SKRWREDFWHULD LQ FORQH FXOWXUHVRQ WKHEDVLVRI6U51$VHTXHQFHV IURPWKHYDULDEOH UHJLRQV 9 9 DQG 9 $SSO (QYLURQ 0LFURELRO 5XWWHU:- (YROXWLRQ RIDOGRODVHV )HG 3URF )HG $P 62& ([S%LRO 6DYDJHDX 0$ 3URWHLQV RI Escherichia coli FRPH LQ VL]HV WKDWDUHPXOWLSOHV RI N'D GRPDLQ FRQFHSWV DQGHYROXWLRQDU\ LPSOLFDWLRQV3URF1DWO$FDG6FL86$ 6FKDHIHU -U -: DQG %R\XP .: 0LFURELDO FRPSRVLWLRQV DQGPHWKRGVIRUWUHDWLQJVRLO8QLWHG6WDWHV3DWHQW 6FKDIHUMRKDQQ -
PLFURVFRS\ &/60 DQG HOHFWURQ PLFURVFRS\ 3URWRSODVPD 6FKXEOHU$ 6FKQHSI( 0ROOHQKDXHU' DQG .OXJH 0 7KH IXQJDO EODGGHUV RI WKH HQGRF\DQRVLV Geosiphon pyriforme, D *ORPXVUHODWHG IXQJXV FHOO ZDOO SHUPHDELOLW\ LQGLFDWHV D OLPLWLQJSRUHUDGLXVRIRQO\QP3URWRSODVPD 6HDEXUJ .* 3DUNHU %& 3UHVFRWW *: DQG :KLWIRUG /$ 7KH DOJDH RI 6RWKHUQ 9LFWRULDODQG $QWDUFWLFD %LEOLRWKHFLD 3K\FRORJLFD %DQG 7UDXVV DQG &UDPHU *PE+ +LUVFKEHUJ *HUPDQ\ 6LOYD +- &RUWLQDV 7, DQG (UWROD 5- (IIHFW RI QXWULWLRQDO IDFWRUV RQ WKH FXOWXUH RI Nostoc VS DV D VRXUFH RI SK\FRELOLSURWHLQV $SSO0LFURELDO%LRWHFKQRO 6LPRQ 5' ,QFOXVLRQ ERGLHV LQ WKH F\DQREDFWHULD F\DQRSK\FLQ SRO\SKRVSKDWH SRO\KHGUDO ERGLHV ,Q )D\ 3 DQG 9DQ %DDOHQ & HGV 7KH &\DQREDFWHULD SS (OVHYLHU 6FLHQFH3XEOLVKHUV%9%LRPHGLFDO'LYLVLRQ 6LQJHU035RVVPLHVVO%0/LHENH&+DQG*URVV&$ 7KH Escherichia coli ts8 PXWDWLRQ LV DQ DOOHOH RI fda, WKH JHQH HQFRGLQJ IUXFWRVHGLSKRVSKDWH DOGRODVH - %DFWHULRO 6PLWK5-+REVRQ6DQG(OOLVIR 7KHHIIHFWRIDEVFLVLFDFLG RQ FDOFLXPPHGLDWHG UHJXODWLRQ RI KHWHURF\VW IUHTXHQF\ DQG QLWURJHQDVHDFWLYLW\LQNostoc 1HZ3K\WRO 6WDQLHU5< 7KHMRXUQH\ QRWWKH DUULYDO PDWWHUV $QQ 5HY 0LFURELRO 6WDQQDFK . &\DQRSK\WD 6LQLFH *ODXFRSK\WD*ODXNRILW\ ,Q 6WDUPDFK . +UVJ )ORUD VORGNRZRGQD 3ROVNL 3DQVWZRZ :\GDZQLFWZR1DXNRZH :DUV]DZD,6 6WHYHQV -U 6( DQG 0XUSK\ 5& 5HFRPELQDQW ELRSHVWLFLGH DQGPHWKRGRIXVHWKHUHRI 8QLWHG6WDWH3DWHQW 6WXOS %. 0RUSKRORJLFDO DQG PROHFXODU DSSURDFKHV WR WKH WD[RQRP\ RI WKH JHQXV Anabaena &\DQRSK\FHDH &\DQREDFWHULD 3K'7KHVLV8QLYHUVLW\RI*URQLQJHQ 6XHQ 0+ (QJOLVK WUDQVODWLRQ 2QVLWH QHZV 6FL 7HFK 'DLO\&KLQD 7DQGHDX GH 0DUVDF 1 'LIIHUHQWLDWLRQ RI KRUPRJRQLD DQG UHODWLRQVKLSV ZLWK RWKHU ELRORJLFDO SURFHVVHV ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU$FDGHPLF3XEOLVKHUV7KH1HWKHUODQGV 7DUDQWR3.HHQDQ7:DQG3RWWV0 5HK\GUDWLRQLQGXFHVD UDSLG RQVHWRIOLSLG ELRV\QWKHVLV LQ GHVLFFDWHG Nostoc commune &\DQREDFWHULD %LRFKLP%LRSK\V$FWD 7HDVH%(DQG:DONHU5: &RPSDUDWLYHFRPSRVLWLRQ RIWKH VKHDWK RI WKH F\DQREDFWHULXP Gloeothece $7&& FXOWXUHG ZLWK DQG ZLWKRXW FRPELQHG QLWURJHQ - *HQ 0LFURELDO 7KLHO 7 /\RQV (0 (UNQFU -& DQG (UQVW $ $ VHFRQG QLWURJHQDVH LQ YHJHWDWLYH FHOOV RI D KHWHURF\VWIRUPLQJ F\DQREDFWHULXP3URF1DWO$FDG6FL86$ 7KRUVWHLQVVRQ 09 %HYDQ '5 (EHO 5( :HEHU5( DQG 3RWWV 0 6SHFWURVFRSLFDO DQG IXQFWLRQDO FKDUDFWHUL]DWLRQ RI WKH KHPRJORELQ RI Nostoc commune 87(; &\DQREDFWHULD %LRFKLP%LRSK\V$FWD 7UHKDQ . DQG 6LQKD 8 '1$PHGLDWHG WUDQVIRUPDWLRQ LQ Nostoc muscorum, D QLWURJHQIL[LQJ F\DQREDFWHULXP $XVW - %LRO6FL 8HQR 7 ,WR + .RWDQL + DQG 1DNDMLPD . 1639 UHVWULFWLRQPRGLILFDWLRQJHQHV 8QLWHG6WDWHV3DWHQW YDQGHQ%HUJK(5(%DNHU6&5DJJHUV 5- 7HUSVWUD3:RXGVWUD (& 'LMNKXL]HQ/DQG 0HLMHU:* 3ULPDU\ VWUXFWXUH DQG SK\ORJHQ\ RI WKH FDOYLQ F\FOH HQ]\PHV WUDQVNHWRODVH DQG IUXFWRVH ELVSKRVSKDWH DOGRODVH RI Xanthobacter flavus. %DFWHULRO
:DOVE\$( *DVYHVLFOHV0LFURELRO5HY :DUG '0 %DWHVRQ 00 :HOOHU 5 DQG 5XII5REHUWV $/ 5LERVRPDO 51$ DQDO\VLV RI PLFURRUJDQLVPV DV WKH\ RFFXU LQ QDWXUH$GY0LFURE(FRO :DUQHU .0 DQG *6 %XOOHMDKQ /LJKWGHSHQGHQW W\URVLQH SKRVSKRU\ODWLRQ LQ WKH F\DQREDFWHULXP Prochlorothrix hollandica. 3ODQW3K\VLRO :KLWWRQ %$ 'RQDOGVRQ $ DQG3RWWV 0 1LWURJHQ IL[DWLRQ E\ Nostoc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nabaena VSVWUDLQ3&& ,Q GH%UXLMQ)- /DSVOHL-5DQG:HLQVWRFN*HGV %DFWHULDO*HQRPHV6WUXFWXUH DQG$QDO\VLV&KDSPDQDQG+DOO :RON &3 (UQVW$DQG(OKDL- +HWHURF\VWPHWDEROLVP DQG GHYHORSPHQW ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU$FDGHPLF 3XEOLVKHUV 7KH 1HWKHUODQGV ;LH:4:KLWWRQ%$ 6LPRQ-:-DJHU.5HHG'DQG 3RWWV0 Nostoc commune 87(; JHQH H[SUHVVLQJ LQGROH SKRVSKDWH K\GURODVH DFWLYLW\ LQ Escherichia coli. - %DFWHULRO
0DOFROP3RWWV
&KDSWHU A rthrospira (Spirulina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
6XPPDU\ 1RWZLWKVWDQGLQJ WKH RIILFLDO UHFRJQLWLRQ LQ RI WKH F\DQREDFWHULD Arthrospiru DQG Spirulina DV GLVWLQFW JHQHUD WKH WHUP "Spirulina" KDV EHHQ XVHG WR LQGLFDWH WKH VSHFLHV RIERWK JHQHUD LQGLIIHUHQWO\ SDUWLFXODUO\ WKH VSHFLHV S platensis" DQG S. maxima" FXOWLYDWHG DQG VROG DV IRRG IHHG DQG D VSHFLDOW\ SURGXFW VRXUFH 7KH VXFFHVVIXO FRPPHUFLDO H[SORLWDWLRQ RIArthrospira, EHFDXVH RILWV KLJK QXWULWLRQDO YDOXH FKHPLFDO FRPSRVLWLRQ B A Whitton and M Potts (eds). The Ecology of cyanobacteria, © Kluwer Academic Publishers Printed in the Netherlands
$9RQVKDNDQG/7RPDVHOOL
DQG WKH VDIHW\ RILWV ELRPDVV KDV PDGH LW RQH RIWKH PRVW LPSRUWDQW LQGXVWULDOO\ FXOWLYDWHG PLFURDOJDH 7KLV FKDSWHU GHVFULEHV WKH HFRORJ\ RIArthrospira, WRJHWKHU ZLWK PRUSKRORJLFDO DQG XOWUDVWUXFWXUDO IHDWXUHV UHOHYDQW IRU VXSSRUWLQJ WKH V\VWHPDWLF SRVLWLRQ RIWKLV RUJDQLVP :KLOH WKH FRQIXVHG WD[RQRP\ RIArthrospira DQG LWV UHODWLRQVKLS ZLWK Spirulina KDYH EHHQ UHVROYHG E\ 6 U51$ VHTXHQFH DQDO\VLV WKH ORQJ GHEDWHG SUREOHP RI VSHFLHV GHILQLWLRQ LV VWLOO RQJRLQJ 2QH VWXG\ RIWHQ VWUDLQV VXJJHVWHG WKDW WKH PRUSKRORJLFDO FULWHULD XVHG WR LGHQWLI\ WKH VSHFLHV $ maxima, A. platensis FRUUHVSRQGHG WR WKH PROHFXODU GDWD REWDLQHG E\ WRWDO '1$ UHVWULFWLRQ SURILOH DQDO\VHV +RZHYHU DQRWKHU VWXG\ EDVHG RQ D ZLGHU UDQJH RI FODVVLFDO VSHFLHV DQG IRUW\ GLIIHUHQWVWUDLQVIDLOHGWRVKRZ DFOHDUFRUUHVSRQGDQFHZLWKPROHFXODUGDWDREWDLQHGIRURQH SDUWRIWKHJHQRPH 7KLV HPSKDVLVHV WKH QHHG WR DGGUHVV WKHSUREOHPRIWKHGHILQLWLRQRIDVSHFLHV E\ XVLQJ DSRO\SKDVLF DSSURDFK .QRZOHGJH RI WKH HFRSK\VLRORJ\ RI Arthrospira, HVVHQWLDO IRU XQGHUVWDQGLQJ WKH JURZWK UHTXLUHPHQWV RI WKLV DONDOLSKLOLF RUJDQLVP LQ WKH QDWXUDO HQYLURQPHQW KDV EHHQ XVHG LQ GHYHORSLQJ VXLWDEOH WHFKQRORJLHV IRU PDVV FXOWLYDWLRQ 7KHUHODWLRQVKLSVEHWZHHQHQYLURQPHQWDO DQGFXOWXUDOIDFWRUV ZKLFKJRYHUQSURGXFWLYLW\ LQ RXWGRRU FXOWXUHV DUH GLVFXVVHG LQ FRQQHFWLRQ ZLWK JURZWK \LHOG DQG HIILFLHQF\ 7KH UHVSRQVH RI Arthrospira DQG LWV PRGLILFDWLRQXQGHUVWUHVV DUHGHVFULEHG WRJHWKHU ZLWKWKH VWUDWHJ\ RIRVPRWLF DGMXVWPHQWDQG WKHPHFKDQLVP RI LQWHUQDOS+UHJXODWLRQWRDONDOLQLW\ 7KHPHWDEROLFSODVWLFLW\ RIWKHUHVSRQVH RIWKLV F\DQREDFWHULXPWR GLVSDUDWH HQYLURQPHQWDO VWLPXOLLVGHPRQVWUDWHGLQWKHQDWXUDOHQYLURQPHQW EXWLV DOVR ZHOOH[SUHVVHGLQWKHPDLQWHQDQFH RIKLJKO\SURGXFWLYHPRQRFXOWXUHLQLQWHQVLYHRXWGRRUFXOWLYDWLRQV\VWHPV
, ,QWURGXFWLRQ (YHUVLQFHArthrospira ZDV ILUVWUHSRUWHG LQ E\ 6WL]HQEHUJHU PDQ\ VSHFLHV RIWKLV JHQXV RIKHOLFDOO\ FRLOHG F\DQREDFWHULD KDYH EHHQ GHVFULEHG DQG LVRODWHG +RZHYHU WKH FODVVLILFDWLRQ RI WKLV JHQXV KDV ORQJ EHHQ D VRXUFH RIFRQIXVLRQ *HLWOHU LQYDOLGDWHG WKH JHQXV Arthrospira LQ KLV UHYLVLRQ RI WKH&\DQRSK\FHDHDQGLQFOXGHGDOOUHJXODUO\KHOLFDOO\ FRLOHG RVFLOODWRULDQ RUJDQLVPV LQ WKH SUHYLRXVO\ GHVFULEHG JHQXV Spirulina 7XUSLQ 6LQFHWKHQ WKH WHUP Spirulina KDV PRVWO\ EHHQ XVHG WR LQGLFDWH VSHFLHV RI HLWKHU JHQXV LQGLIIHUHQWO\ $V GHVFULEHG ODWHU LW LV QRZ UHFRJQL]HG WKDW WKHUH DUH WZR YHU\ GLVWLQFWJHQHUD&DVWHQKRO] 7KHUHIRUHPDQ\ VSHFLHV WKDW FXUUHQWO\ EHDU WKH JHQHULF HSLWKHW Spirulina, VKRXOGEHUHLQFOXGHGLQWKHRULJLQDO JHQXV Arthrospira. +RZHYHU WKH WHUP 6SLUXOLQD LV ZLGHO\ XVHG WR LQGLFDWH WKH HFRQRPLFDOO\ LPSRUWDQW VSHFLHV "S. platensis" DQG "S. maxima", VRZHXVHLWLQSDUWV RI WKLV FKDSWHU WR DYRLG FRQIXVLRQ ZLWK SUHYLRXV VWXGLHV Arthrospira KDV EHHQ UHSRUWHG WR H[LVW LQ HQYLURQPHQWV YDU\LQJ LQ WKHLU RVPRWLFXP WHPSHUDWXUH DQG VDOW FRQFHQWUDWLRQV PRVW RI WKHP EHLQJ RI KLJK DONDOLQLW\ ,OWLV D E %XVVRQ )LODPHQWV RIWKH JHQXV Spirulina LQ WKH VWULFW VHQVHDOVRRFFXULQPDQ\RIWKHVHHQYLURQPHQWV 7KHUH LV JUHDW LQWHUHVW LQ SDVW XVH RI WKH XVH RI Arthrospira DV D WUDGLWLRQDO IRRG 'DQJpDUG %UDQGLO\ /pRQDUG DQG /pRQDUG DQG &RPSpUH DOO GHVFULEHG KRZ $IULFDQ WULEHV OLYLQJ DORQJ /DNH &KDG FROOHFW WKLV PLFURVFRSLF DOJD $ platensis). ,W LV FROOHFWHG IURP ZDWHU
ERGLHV QHDU WKH ODNH DQG VXQGULHG RQ WKH VKRUHV WR SURGXFHDKDUGHQHGGDUNFDNHFDOOHG GLKp ZKLFKLV EURNHQ LQWR VPDOO SLHFHV DQG XVHG LQ GLIIHUHQW IRUPV E\WKHORFDOSRSXODWLRQVDVSDUWRIWKHLUGDLO\GLHW $W DERXW WKH VDPH WLPH DQG RQ WKH RWKHU VLGH RI WKH JOREH Arthrospira ZDV DOVR UHFRUGHG LQ WKH ZDWHU RI /DNH 7H[FRFR 0H[LFR +HUH LW KDG EHHQ XVHG DV IRRG E\ WKH QDWLYHV OLYLQJ LQ WKH DUHD &OpPHQW 7UDYHOHUV WR 0H[LFR GXULQJ WKH WK FHQWXU\ GHVFULEHG KRZ WKH $]WHFV XVHG D VRIW D EOXHJUHHQ PDWHULDO KDUYHVWHG ZLWK ILQH QHWV IURP WKH ODNH IRU PDNLQJ D NLQG RI EUHDG FDOOHG WHFXLWODWO &LIHUUL ,W LV FOHDU WKDW WKHVH WZR GLIIHUHQW FXOWXUHV OLYLQJ IDU DSDUW GLVFRYHUHG WKH QXWULWLRQDO YDOXH RI Arthrospira LQGHSHQGHQWO\ /DWHU DWWHQWLRQ ZDV UHIRFXVHG RQ "Spirulina" E\ WKHSLRQHHULQJZRUNGRQHDWWKH,QVWLWXWH)UDQoDLVHGX 3pWUROH ZLWK VWXGLHV RQ WKH F\DQREDFWHULDO EORRPV RFFXUULQJ LQ WKH HYDSRUDWLRQ SRQGV RI WKH LQGXVWULDO VRGD SURGXFWLRQ IDFLOLW\ DW /DNH 7H[FRFR QHDU 0H[LFR &LW\ 7KLV OHG WR WKH ILUVW GHWDLOHG VWXG\ RI WKH JURZWK UHTXLUHPHQWV DQG SK\VLRORJ\ RI Arthrospiru. 7KH3K' VWXG\E\=DUURXN ZDV WKH EDVLV IRU HVWDEOLVKLQJ WKH ILUVW ODUJHVFDOH SURGXFWLRQ SODQW RI Arthrospira. 7KH ZRUN ZDV IROORZHG XS E\ VHYHUDO JURXSV LQ ,WDO\ )UDQFH DQG ,VUDHO DQGZDVVXPPDUL]HGLQDGHWDLOHGUHYLHZSDSHU E\ &LIHUUL 7KH H[WHQVLYH UHVHDUFK RQ FHOO ELRORJ\ ELRFKHPLVWU\ DQG ELRWHFKQRORJ\ FDUULHG RXW VLQFHWKHQLVUHYLHZHGLQWKHERRNHGLWHGE\9RQVKDN D 7KLV FKDSWHU SURYLGHV D SHUVSHFWLYH RQ WKH V\VWHPDWLFV DQG WKH HFRSK\VLRORJ\ RI "Spirulina",
&KDSWHU $UWKURVSLUD 6SLUXOLQD QRZ FRUUHFWO\ UHDVVLJQHG 6WL]HQEHUJHU
WR
Arthruspira
,, 0RUSKRORJ\ $ 7KH+HOLFDO6KDSHRIWKH)LODPHQW 7KHPDLQ PRUSKRORJLFDO IHDWXUHRIArthruspira LV WKH SDWWHUQHG DUUDQJHPHQW RILWV PXOWLFHOOXODU F\OLQGULFDO WULFKRPHV LQ DQ RSHQ KHOL[ 7KH WULFKRPHV DUH FRPSRVHG RI F\OLQGULFDO FHOOV WKDW XQGHUJR ELQDU\ ILVVLRQ LQ D VLQJOH SODQH SHUSHQGLFXODU WR WKH PDLQ D[LV 7ULFKRPH HORQJDWLRQ RFFXUV WKURXJK PXOWLSOH LQWHUFDODU\ FHOO GLYLVLRQ DORQJ WKH HQWLUH ILODPHQW 0XOWLSOLFDWLRQ RFFXUV RQO\ E\ IUDJPHQWDWLRQ 7ULFKRPH EUHDNDJH LV WUDQVFHOOXODU DQG LQYROYHV GHVWUXFWLRQ RIDQ LQWHUFDODU\FHOO 7KH PHFKDQLVPDV LW RFFXUV LQ "S. maxima" DQG LQ "S. platensis" ZDV GHVFULEHG LQ GHWDLO E\ 7RPDVHOOL HW DO ,Q Spirulina, WULFKRPH EUHDNDJH LV LQWHUFHOOXODU 5LSSND HWDO $QDJQRVWLGLVDQG.RPi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rthruspira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rthruspira DOVR VXJJHVWV WKDWWKH KHOLFDO FKDUDFWHU PD\ EH FDUULHG RQ SODVPLGV EXW QR RQH KDV \HW GHPRQVWUDWHG WKH H[LVWHQFH RI SODVPLGV LQ Arthrospira RU Spirulina
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pirulina VSHFLHV DOVRVWUDLJKWHQ DIWHU WKH\ KDYH EHHQ VWUHDNHG RQ SODWHV GXULQJ LVRODWLRQ SURFHGXUHV RU DIWHU UHSHDWHG WUDQVIHUV RQWR VROLGPHGLD/7RPDVHOOLXQSXEOLVKHG
% 8OWUDVWUXFWXUH 7KH FHOO RUJDQL]DWLRQ RI Arthruspira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
$9RQVKDNDQG/7RPDVHOOL
Fig. 1. 7KLQVHFWLRQVKRZLQJWKHWLQHVWUXFWXUHRIArthrospira maxima. 1RWH FHOOZDOOFZ VHSWXPV DQGWK\ODNRLGVW %DUPDUNHU PSKRWRFRXUWHV\RI053DODQGUL
F\DQRSK\FLQJUDQXOHV 6WUXFWXUHV OLNHWKHF\OLQGULFDO ERGLHV UHSRUWHG IRU VRPH RWKHU PHPEHUV RI WKH 2VFLOODWRULDFHDH DUH RIWHQ SUHVHQW 7KHVH LQFOXVLRQVZKRVH H[SODQDWLRQ LV VWLOO XQNQRZQ DUH ODFNLQJLQSpirulina 7RPDVHOOLHWDO
,,, 6\VWHPDWLFV 7KH ILUVW UHSRUW RID WD[RQ ZKLFK FDQ EH DVVLJQHG WR Arthrospira 6WL]HQEHUJHU GDWHV EDFN WR PRUH WKDQ \HDUV DJR ZKHQ :LWWURFN DQG 1RUGVWHGW UHSRUWHG WKH RFFXUUHQFHQHDU0RQWHYLGHR RID EOXHJUHHQ DOJD ZLWKKHOLFDO VKDSHG ILODPHQWV ZKLFK WKH\ GHVFULEHG DV "Spirulina jenneri I platensis", HYHQ WKRXJK LW SRVVHVVHG VHSWD $W WKDW WLPH WKH JHQXV Spirulina ZDV FRQVLGHUHG WR EH XQLFHOOXODU 6RPH \HDUV ODWHU 6WL]HQEHUJHU ZKR KDG REVHUYHG VHSWD LQ VRPH KHOLFDOO\ FRLOHG RVFLOODWRULDFHDQ IRUPV SURSRVHGWKDW WKH IRUPV ZLWK DPXOWLFHOOXODUVWUXFWXUHEHLQFOXGHGLQDQHZ JHQXV Arthrospira. 7KLV GLVWLQFWLRQ ZDV FRQILUPHG E\ *RPRQW LQ KLV WD[RQRPLF VWXG\ RQ WKH 2VFLOODWRULDFHDH +H OHIW WKH DSSDUHQWO\ DVHSWDWH IRUPV LQ Spirulina, DQG SODFHG WKH VHSWDWH IRUPV LQ Arthrospiru 6WL]HQEHUJHU +RZHYHU *HLWOHU V UHYLVLRQ RI WKH &\DQRSK\FHDH UHXQLILHG WKH PHPEHUV RI WKHVH WZR JHQHUD LQ Spirulina 7XUSLQ +H EDVHG FODVVLILFDWLRQ RQWKHFORVH VLPLODULW\ LQ PRUSKRORJ\ DQG LJQRUHG WKH SUHVHQFH RI VHSWD
YLVLEOH RU RWKHUZLVH DQG WKXV PDGH QR GLVWLQFWLRQ EHWZHHQ WKH IRUPV SUHYLRXVO\ UHFRJQL]HG DV VHSDUDWH JHQHUD *HLWOHU DFWXDOO\ VSOLW WKH JHQXV LQWR WZR VXEJHQHULF WD[D 6HFWLRQ , $UWKURVSLUD DQG 6HFWLRQ (XVSLUXOLQD RQ WKH EDVLV RI WKH FULWHULRQ RULJLQDOO\XVHGE\6WL]HQEHUJHU WRVHSDUDWHWKH JHQHUD Spirulina DQG Arthrospira, LH YLVLEOH RU QRQ YLVLEOH FURVVZDOOV 7KH IRUPV ZLWK VHSWD WKDW ZHUH HDVLO\ REVHUYDEOH XQGHU D OLJKW PLFURVFRSH ZHUH FODVVLILHG LQ WKH 6HFWLRQ $UWKURVSLUD DQG WKRVH ZLWK XQOLNHO\ RU RQO\ DUWLILFLDOO\ REVHUYDEOH VHSWD DIWHU WU\SVLQ WUHDWPHQWRU VWDLQLQJ ZLWK QHXWUDO UHG LQ WKH 6HFWLRQ(XVSLUXOLQD ,Q %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ &DVWHQKRO] DWWHPSWHG WRUHVROYHWKH PDWWHUE\ UHVWRULQJ Arthrospira WR LWV RULJLQDO UDQN RI JHQXV +H VXJJHVWHG XVLQJ WKUHH IHDWXUHV WR VHSDUDWH Spirulina DQG Arthrospira LQ 6XEVHFWLRQ ,,, RI WKH 2UGHU 2VFLOODWRULDOHV ZKLFK VKDUHV WKH SURSHUW\ RI KHOLFDOO\FRLOHGWULFKRPHV 7KHVHDUH GHJUHH RI LQFOLQDWLRQ RI WKH SLWFK RI WULFKRPH KHOL[IURPWUDQVYHUVHD[LV DVSHFW DQG YLVLELOLW\ OLJKW PLFURVFRSH RIFURVV ZDOOVEHWZHHQWKHFHOOVLQWKHILODPHQW GLVWULEXWLRQ HOHFWURQ PLFURVFRS\ RIMXQFWLRQDO SRUHVLQWKHFHOOZDOO 7KXV Arthrospira FDQEHGLIIHUHQWLDWHGIURPSpirulina RQ WKH EDVLV RIWKH GHJUHH RILQFOLQDWLRQ RIWKH SLWFK RI WKH WULFKRPH KHOL[ ZKLFK IRUPV DQ DQJOH !
&KDSWHU $UWKURVSLUD 6SLUXOLQD
Table 1. 0DLQIHDWXUHVVHSDUDWLQJWKHJHQHUDArthrospira 6WL]HQEHUJHUDQGSpirulina 7XUSLQ &ULWHULD
$UWKURVSLUD
6SLUXOLQD
7ULFKRPH GLDPHWHU
P
P
7\SH RI KHOLFLW\
ORRVHO\ FRLOHG
WLJKWO\ FRLOHG
&URVVZDOOV
YLVLEOH XQGHU OLJKW PLFURVFRSH
LQYLVLEOH XQGHU OLJKW PLFURVFRSH
&HOO ZDOO SRUH SDWWHUQ
VLQJOH URZ DURXQG WKH WULFKRPH
VHYHUDO URZV RQ FRQFDYH VLGH RI WKH FRLO
0RGH RI WULFKRPH IUDJPHQWDWLRQ
LQWUDFHOOXODU QHFULGLXP
LQWHUFHOOXODU
&\OLQGULFDO ERGLHV
SUHVHQW
DEVHQW
$QR[\JHQLF SKRWRV\QWKHVLV
DEVHQW
SUHVHQW LQ VRPH VWUDLQV
&SK\FRHU\WKULQ
QRW IRXQG
SUHVHQW LQ VRPH VWUDLQV
\OLQROHQLF DFLG
SUHVHQW
DEVHQW
IURPWKHWUDQVYHUVH D[LV WKH SUHVHQFH RIHDV\ YLVLEOH VHSWD DQG WKH GLVWULEXWLRQ RIMXQFWLRQDO SRUHV LQ RQH FLUFXODUURZDURXQGWKHFURVVZDOOV 7RPDVHOOL HWDO VXJJHVWWKDWWKHUHDUH RWKHU PHDQLQJIXO FULWHULD WR GLVWLQJXLVK EHWZHHQ WKH WZR LQFOXGLQJ 6 U51$ VHTXHQFH GDWD ZKLFK FOHDUO\ VKRZ WKDW Spirulina DQG Arthrospira DUH SK\ORJHQHWLFDOO\ GLVWDQW *LRYDQQRQL HW DO 1HOLVVHQ HW DO $QRWKHU XVHIXO FULWHULRQ LV WKDW XQOLNH Spirulina, Arthrospiru FRQWDLQV WKH XQVDWXUDWHG IDWW\ DFLG OLQROHQLF DFLG &RKHQ DQG 9RQVKDN Spirulina DQG Arthrospiru DUH ERWK DTXDWLF EXW XQOLNH Spirulina, Arthrospiru KDV JDV YDFXRODWHGFHOOV 7KHPDLQFULWHULDIRUGLVWLQJXLVKLQJ WKHWZRJHQHUDDUHOLVWHGLQ7DEOH $OWKRXJK Arthrospira LV KHOLFDOO\ FRLOHG LQ QDWXUDO SRSXODWLRQV WKH RFFXUUHQFH RI VWUDLJKW RU VOLJKWO\ VWUDLJKW ILODPHQWRXV IRUPV VLPLODU WR Oscillatoriu, KDYH EHHQ UHSRUWHG UHSHDWHGO\ LQ FXOWXUH PDNLQJ VLPSOH PRUSKRORJLFDO REVHUYDWLRQ LQYDOLG LQ WKH LGHQWLILFDWLRQ RI Arthrospiru /HZLQ 7KH SUHVHQFH RI VWUDLJKW IRUPV RI Arthrospiru DQG WKHLU VLPLODULW\ WR VRPH PHPEHUV RI WKH WULEH 2VFLOODWRULDH &DVWHQKRO] GXH WR D VLPLODU MXQFWLRQDO SRUH GLVWULEXWLRQ *XJOLHOPL DQG &RKHQ %D]LUH HPSKDVL]H WKH QHHG WR ILQG DGGLWLRQDO SKHQRW\SLFIHDWXUHVWRGLVWLQJXLVKArthrospiru. $FFRUGLQJWRWKH%RWDQLFDO&RGHA. jenneri +DVV 6WL]HQEHUJHULVWKHW\SHVSHFLHVDQGA. platensis 3&& ZKLFK ZDV LVRODWHG IURP D VDOLQH PDUVK 'HO
0DU6ORXJK &DOLIRUQLD 86$ LV WKHUHIHUHQFH VWUDLQ IRU WKH %DFWHULRORJLFDO &RGH 5LSSND DQG +HUGPDQ 7DEOH +LQGILN DVVLJQHGDOOWKHSODQNWRQLFIRUPV RI Arthrospira LQFOXGLQJ WKRVH SUHYLRXVO\ GHVFULEHG DV 6 platensis) LQ DONDOLQH VDOLQH ODNHV WR "S. fusiformis" 9RURQLFKLQ V\QRQ\PV A. maxima 6HWFKHOO HW *DUGQHU DQG A. geitleri 'H 7RQL VLQFH LQ DJUHHPHQW ZLWK )RWW DQG .DULP KH FRQVLGHUHG 6 platensis" WR EH D EHQWKLF VSHFLHV 2QWKHEDVLVRIWKLV DVVXPSWLRQ PRVWRIWKH $IULFDQ SRSXODWLRQV ZHUH LQFOXGHG XQGHU A. fusiformis. 7KLV RSLQLRQ DOVR VKDUHG E\ .RPILUHN DQG /XQG KDV EHHQ GHEDWHG E\ 7RPDVHOOL .RPILUHNDQG /XQG LQWKHLUUHYLVLRQ RI WKH WD[RQRP\ DQG QRPHQFODWXUH RI WKH WKUHH UHODWHG VSHFLHV S platensis", “S. maxima" DQG "S. fusiformis", VXJJHVWHG WKDW ZLWKLQ WKH SODQNWRQLF IRUPV WKHUH DUH WZR WD[D "maxima" DQG “fusiformis" DQG IXUWKHUPRUH WKDW WKHVH WZR WD[D ZKLFK FRXOG UHSUHVHQW WZR VSHFLHV (A. fusiformis .RPiUHN DQG $ maxima), KDYH GLIIHUHQW JHRJUDSKLFDO GLVWULEXWLRQV +HQFH WKH\ FRQVLGHU A. maxima WR EH SDQWURSLFDOA. fusiformis WR EH OLPLWHG WR $IULFD DQG WURSLFDO DQG FHQWUDO $VLD DQG FRQVLGHU WKH EHQWKLF A. platensis WR EH HVVHQWLDOO\ UHVWULFWHG WR6RXWK$PHULFD 0RUH UHFHQWO\ 'HVLNDFKDU\ DQG -HHML %DL SURSRVHG WKDWDOO WKHFDO\SWUDWHIRUPV RIA. fusiformis EHLQFOXGHGLQWKHQHZVSHFLHVA. indica 'HVLNDFKDU\
$9RQVKDNDQG/7RPDVHOOL
Table 2. 6SHFLHVRIWKHJHQXVArthrospiru 6WL]HQEHUJHU Species
First description
References
A. fusiformis .RPiUHN
6LEHULDQ VWHSSH 5XVVLD /DNH 7XQDWDQ
9RURQLFKLQ
A. gomontiana 6HWFKHOO
1RUWK $PHULFD VWDJQDQW ZDWHU
*HLWOHU
A. indica 'HVLNDFKDU\ -HHML %DL
0DGXUDL ,QGLD QDWXUDO SRQG
'HVLNDFKDU\ -HHML %DL
A. jenneri 6WL]HQEHUJHU
(XURSH VWDJQDQW ZDWHU
*RPRQW
A. khannae 'URXHW 6WULFNODQG
5DQJRRQ 0\DQPDU QDWXUDO SRQG
'HVLNDFKDU\
A. massartii .XIIHUDWK
/X[HPEXUJ VSULQJ ZDWHU
*HLWOHU
A. maxima 6HWFKHOO *DUGQHU
2DNODQG &DOLIRUQLD ZDUP VDOLQH SRQG
*DUGQHU
A. platensis *RPRQW
0RQWHYLGHR 8UXJXD\ VWDJQDQW ZDWHU
:LWWURFN 1RUGVWHGW
A. spirulinoides &KRVH
/DKRUH 3DNLVWDQ VWDQGLQJ UDLQ ZDWHU
*HLWOHU
A. tenuis %ULLKO %LVZDV
%HQJDO ,QGLD DUWLILFLDO EDVLQ
*HLWOHU
DQG-HHML%DL WKLVLPSOLHVWKDWWKHWKLFNHQLQJRI WKH DSLFDO FHOO ZDOO FDO\SWUD LV D VLJQLILFDQW WD[RQRPLFIHDWXUH 7KHSURSRVDOLVEDVHGRQWKHIDFW WKDW WKH FDO\SWUD ZDV QHYHUUHSRUWHG LQA. fusiformis. ,IWKH H[LVWHQFH RI WKH FDO\SWUD FDQ EH GHPRQVWUDWHG LQWKH RULJLQDOPDWHULDOA. indica ZLOOEHFDPHDODWHU V\QRQ\PRIA. fusiformis 'HVLNDFKDU\ DQG-HHML %DL 1HYHUWKHOHVV WKH SURSRVDO IRU D QHZ VSHFLHV A. indica LVTXHVWLRQDEOHVLQFHRWKHUDXWKRUV)RWWDQG .DULP FRQVLGHU ³S. geitleri” ZKLFK LV D V\QRQ\PRIWKHFDO\SWUDWH“S. maxima“ WREHLGHQWLFDO WR “S. fusiformis" 7KH WUDGLWLRQDO VHSDUDWLRQ RI WKH FRPPRQ VSHFLHV LQWR A. maxima DQG A. platensis VKRXOGEHPDLQWDLQHG XQWLO PRUH GHWDLOHG VWXGLHV KDYH EHHQ FRPSOHWHG 7RPDVHOOL &RQVLGHULQJ WKH JUHDW YDULDWLRQV LQVL]HDQGVKDSHRIWULFKRPHJHRPHWU\LQQDWXUDODQG FXOWXUHG Arthrospira SRSXODWLRQV WD[D EDVHG RQ PRUSKRORJLFDO IHDWXUHV VKRXOG EH FRQILUPHG WKURXJK XVH RI PROHFXODU ELRORJ\ WHFKQLTXHV 7RWDO '1$ UHVWULFWLRQ SURILOH DQDO\VLV SHUIRUPHG RQ WHQ Arthrospira VWUDLQV OHG WR WKH UHFRJQLWLRQ RI WZR ZHOOVHSDUDWHG JHQRW\SLF JURXSV 9LWL HW DO FRLQFLGLQJ ZLWKWKHERWDQLFDO VSHFLHVA. maxima DQG A. platensis, WRZKLFKWKHVWUDLQVKDGSUHYLRXVO\EHHQ DVVLJQHG7RPDVHOOLHWDO D $ IXUWKHU SK\ORJHQHWLF VWXG\ XVLQJ $5'5$ $PSOLILHG 5LERVRPDO '1$ 5HVWULFWLRQ $QDO\VLV RI WKH ,76 ,QWHUQDO 7UDQVFULEHG 6SDFHU DV DPROHFXODU WD[RQRPLFPDUNHUZDVPDGHRQPRUHWKDQVWUDLQV RI Arthrospira, ZKLFK GLIIHUHG ZLWK RQH H[FHSWLRQ IURP WKH SUHYLRXV RQHV 6FKHOGHPDQ HW DO 7KHUHVXOWVRIWKLVVWXG\OHGWKHVWUDLQVEHLQJJURXSHG
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rthrospiru H[WHQGV WR WKH FRPPHUFLDO SURGXFWV DQG KXPDQKHDOWK VLQFHArthrospira ELRPDVV LV VROG DV D KHDOWKIRRG7KHZLGHVSUHDGXVHRIWKHWHUP6SLUXOLQD IRU VXFK PDWHULDO FOHDUO\ FDXVHV SUREOHPV LQ DFFHSWLQJ WKH FRUUHFW WHUPLQRORJ\ EXW LW LV KLJKO\ LPSRUWDQW WR EH DZDUH RIWKH W\SH RIRUJDQLVP EHLQJ FXOWLYDWHG 6RPHSpirulina VSS HJS. subsalsa) DUH LQ IDFW FXOWLYDWHG IRU FRPPHUFLDO SXUSRVHV LQ PLVWDNHQ EHOLHIWKDW WKH\ KDYH WKH QXWULWLRQDO TXDOLW\ DQG WR[LFRORJLFDO VDIHW\ RI WKH WUDGLWLRQDO HGLEOH VSHFLHV RIArthrospira 6KLPDGD HW DO %ULHED DQG0HULQRV
,9 2FFXUUHQFHDQG 'LVWULEXWLRQ $ *HQHUDO 6SHFLHV RI Arthrospira KDYH EHHQ LVRODWHG PDLQO\ IURP DONDOLQH EUDFNLVK DQG VDOLQH ZDWHUV LQ WURSLFDO DQG VHPLWURSLFDO UHJLRQV &DVWHQKRO] A. jenneri DQG A. platensis ZHUH GHVFULEHG E\ *HLWOHU
&KDSWHU $UWKURVSLUD 6SLUXOLQD DV FRVPRSROLWDQ 7KRXJK PRVW RI WKH Arthrospira VSHFLHV KDYH EHHQ IRXQG LQ DONDOLQH ZDWHU ZKHUH WKH\ IRUP PDVVLYH EORRPV VRPH RWKHU VSHFLHV KDYH EHHQ UHSRUWHG WR RFFXU LQ IUHVK ZDWHUV EXW ZLWKRXW VXFK SUHGRPLQDQFH 2WKHU VSHFLHV ZLWK PRUH UHVWULFWHG GLVWULEXWLRQV RFFXU LQ UXQQLQJ IUHVK ZDWHUV A. okensis DQG VSULQJV A. massartii A. spirulinoides, A. gomontiana DQG A. tenuis DUH UHSRUWHG IRU VWDJQDQW IUHVK ZDWHUV 'HVLNDFKDU\ FRQVLGHUVA. jenneri DQG A. platensis DVW\SLFDO RIWURSLFDO VDOLQHHQYLURQPHQWV $PRQJ WKH YDULRXV VSHFLHV LQFOXGHG LQ WKH JHQXV Arthrospira, WKHPRVWZLGHO\GLVWULEXWHGA. platensis, LV PDLQO\ LQ $IULFD &KDG .HQ\D (J\SW (WKLRSLD 6XGDQ /LE\D $OJHULD &RQJR =DLUH =DPELD $VLD 3DNLVWDQ ,QGLD 6UL/DQND 0\DQPDU 7KDLODQG DQG 6RXWK$PHULFD8UXJXD\3HUX A. maxima V\Q A. geitleri) DSSHDUV WR EH FRQILQHG WR &HQWUDO $PHULFD 0H[LFR &DOLIRUQLD86$ 7KLV VSHFLHV LV WKH PDLQ FRPSRQHQW RI WKH SK\WRSODQNWRQ RI WKH VRODU HYDSRUDWLRQ FKDQQHOV LQ /DNH 7H[FRFR 0H[LFR ZKLOH A. platensis GRPLQDWHVWKHDONDOLQHVDOLQH ODNHV RIWKH VHPLGHVHUWLF 6XGDQ6DKHO DUHD &KDG DQG RI WKH 5LIW 9DOOH\V .HQ\D 7KH DEXQGDQW RFFXUUHQFH RIWKH VSHFLHV A. platensis DQG A. maxima LQ VDOLQH DONDOLQH WURSLFDO DQG VXEWURSLFDO ZDWHUV KDV OHG WR UHFRJQLWLRQ RIWKHVH HQYLURQPHQWV DV W\SLFDO IRU WKH HQWLUHJHQXV 7KH JHRJUDSKLFDO GLVWULEXWLRQ RI WKH PDLQ SRSXODWLRQV RI Arthrospira LV JLYHQ LQ 7DEOH Arthrospira VSHFLHV KDYH DSSDUHQWO\ QRW EHHQ UHSRUWHG IURPPDULQH HQYLURQPHQWV +RZHYHU ZKHQ DQDGHTXDWH VXSSO\RIELFDUERQDWH 1DQG 3 WRJHWKHU ZLWK VXLWDEOH S+ DQG VDOLQLW\ YDOXHV DUH SURYLGHG Arthrospira VSHFLHV FDQ EH KLJKO\ SURGXFWLYH LQ VHD ZDWHU7UHGLFLHWDO 7KH PRVW GHWDLOHG VWXGLHV RQ WKH HFRORJ\ RI $ platensis DUHWKRVHRI,OWLV DE D E DE IRUWURSLFDO$IULFDQODNHVDQGWKDWRI %XVVRQ ZKR GHVFULEHG SDVWDQG UHFHQWXVH RI "S. geitleri" V\Q A. maxima DV D IRRG VRXUFH LQ 0H[LFR LQ DGGLWLRQ WR LWV HFRORJ\ 2WKHU VWXGLHV LQFOXGH WKRVH RI -HHML %DL DQG 6HVKDGUL +LQGDN .RPiUHN DQG /XQG DQG 'HVLNDFKDU\DQG-HHML%DL 7KHODVWGHDOVZLWK WKH GLVWULEXWLRQ RI ,QGLDQ LVRODWHV DQG UHYLVHV WKH WD[RQRP\ RI VRPH FXOWXUH FROOHFWLRQ VWUDLQV DVVLJQLQJWKHPWRWKHFDO\SWUDWHVSHFLHVA. indica.
% :DWHU%DVLQVRIWKH6XGDQ6DKHO=RQH 7KH H[WHQVLYH VXUYH\ RI WKH SK\WRSODQNWRQ RI WKH SHUPDQHQW DQG WHPSRUDU\ ODNHV QHDU /DNH &KDG
SHUIRUPHG E\ ,OWLV IRU 256720 2IILFH GH OD 5HFKHUFKH 6FLHQWLILTXHHW7HFKQLTXH 2XWUHPHUV KDV FRQWULEXWHG PXFK WR RXU NQRZOHGJH RIA. platensis. ,WV PDLQ GLVWULEXWLRQ LV LQ WKH ZDWHU EDVLQ RI WKH 6XGDQ6DKHO ]RQH LQ &HQWUDO $IULFD QRUWKHDVW RI /DNH &KDG WRZDUGV WKH UHJLRQ RI 2XQLDQJD .HELU DQG WKH EDVLQV RI WKH (DVW $IULFDQ 5LIW 9DOOH\V ,W DOVR RFFXUV LQ (WKLRSLD /DNH &KLWX DQG /DNH $UDQJXDGL DQG=DLUH/DNH.LYX 7DEOH 0RVW RI WKH ZDWHU ERGLHV LQ ZKLFK Arthrospira WKULYHV DUH SHUPDQHQW RU WHPSRUDU\ VDOLQH DONDOLQH ODNHVQDWURQODNHV WKHVHDUHVFDWWHUHGLQWKHKROORZV RI WKH IRVVLO GXQH V\VWHP RI WKH 6DKDUD DQG WKH 6XGDQ6DKHODUHDV DQGIHG E\DTXLIHUVDQGWKHVFDQW\ UDLQ DERXW PPSHU \HDU 7KH VDOLQLW\ RIWKHVH ODNHV LV KLJKO\ YDULDEOH ZKHQ HYDSRUDWLRQ H[FHHGV LQIORZWKH VDOLQLW\ULVHV DQG OHDGV WRDQ DFFXPXODWLRQ RI VDOW GHSRVLWV RQ WKH VKRUHV ,OWLV E 7KH PHDQZDWHUWHPSHUDWXUHLV a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platensis LV SUHVHQW DV QHDUO\ PRQRVSHFLILF SRSXODWLRQV LQ WKH KLJKO\ VDOLQH ODNHV ,OWLV D RQO\ D IHZ RWKHU SKRWRWURSKV EHLQJ IRXQG ,Q FRQWUDVW LQ WKH PHVRKDOLQH ODNHV Arthrospira FRH[LVWV ZLWK D QXPEHU RI RWKHU SKRWRWURSKV HVSHFLDOO\GLDWRPVDQGGLQRIODJHOODWHV LQDGGLWLRQWR WKH GRPLQDQW F\DQREDFWHULD Anabaenopsis arnoldii DQG PDQ\ Oscillatoria VSS DUH RIWHQ LPSRUWDQW FRPSRQHQWV RI WKH F\DQREDFWHULDO SRSXODWLRQ 6HYHUDO Spirulina VSHFLHV PD\ EH SUHVHQW HVSHFLDOO\ S. laxissima, S. major DQG S. subsalsa. $ VPDOOHU IRUP RI A. platensis, YDU minor 5LFK ZDV DOVR SUHVHQW ,OWLV E ,Q WKH VWXGLHV SHUIRUPHG RQ VDPSOHV RIWZR $IULFDQ QDWURQ ODNHV /DNH 0RPEROR DQG /DNH 5RPERX RI WKH UHJLRQ RI .DQHP O\LQJ QRUWKHDVWRI/DNH&KDG0DUJKHULHWDO IRXQG
$9RQVKDNDQG/7RPDVHOOL
Table 3. *HRJUDSKLFDOGLVWULEXWLRQRIWKHPDLQSRSXODWLRQVRIArthrospira VSS COUNTRY
ENVIRONMENT
SPECIES
REFERENCES
QDWURQ ODNHV %RGRX 0RPEROR 5RPERX
A. platensis, A. platensis /pRQDUG &RPSpUH I minor ,OWLV 5LFK
$)5,&$ &KDG
.HQ\D
A.
5LFK 9DUHVFKL
platensis
A. platensis, A. platensis I minor, A. fusiformis A. platensis, A. platensis A. fusiformis A. platensis, A. platensis $ fusiformis A. platensis A. geitleri A. platensis Arthrospira VS Arthrospira VS Arthrospira VS
7XLWH +LQGiN
SRQGV /RQDU /DNH SRQGV WDQN 0DGXUDL 0&5& LVRODWH &UDWHU ODNHV ILVK SRQG /DKRUH /DNH %HULD ILVK SRQGV 1DQNLQJ WDSLRFD IDFWRU\ HIIOXHQW ODNHV %DQJNRN 7XQDWDQ ODNH 6LEHULDQ VWHSSH SRQG /DKRUH
A. maxima A. indica
'HVLNDFKDU\ -HHML %DL 'HVLNDFKDU\ -HHML %DL
Arthrospira VS Arthrospira VS Arthrospira VS A. platensis Arthrospira VS A. fusiformis Arthrospira VS
0LQ 7KHLQ )R[ )R[ 7VHQ &KDQJ )R[ 9RURQLFKLQ )R[
0H[LFR &DOLIRUQLD 3HUX
/DNH 7H[FRFR VRODU HYDSRUDWRU SRQG 2DNODQG FRDVWDO ODJRRQ 'HO 0DU /DNH +XDFKDFKLQD
A. maxima A. maxima, A. platensis" A. platensis, A. maxima
8UXJXD\
0RQWHYLGHR
A.
%XVVRQ *DUGQHU /HZLQ %XVVRQ 'HVLNDFKDU\ -HHML %DL *RPRQW
/DNH 6DQWD 2ODOOD WLQ\ ODNH &DPDUJXH $GDV]WHYHO2URVKD] ZDWHU EDVLQ .KXPEDVKD
Arthrospira Arthrospira Arthrospira Arthrospira
/DNH 6LPEL (WKLRSLD
/DNH $UDQJXDGL /DNH &KLWX *UHHQ /DNH
(J\SW 6XGDQ $OJHULD &RQJR =DLUH =DPELD
/DNH 0DU\XW /DNH 'DULED -HEHO 0DUUD SRQG 7DPDQUDVVHW /DND 0RXJRXQJD /DNH .LYX /DNH %DQJZHRORX
0HODFN .HEHGH $KOJUHQ 0HODFN .HEHGH $KOJUHQ (O%HVWDZ\ HW DO )RWW .DULP )R[ )R[ )R[ )R[
$6,$ ,QGLD
0\DQPDU 3DNLVWDQ 6UL /DQND &KLQD 7KDLODQG 5XVVLD 3DNLVWDQ $0(5,&$
platensis"
(8523( 6SDLQ )UDQFH +XQJDU\ $]HUEDLMDQ
UHIHUHQFH VWUDLQ 3&& KRORW\SH
VS VS VS VS
5LSSND +HUGPDQ )R[ %XVVRQ )R[
&KDSWHU $UWKURVSLUD 6SLUXOLQD WKH DERYHPHQWLRQHG Spirulina VSHFLHV DQG VHYHUDO Arthrospiru VSS 7KH RYHUZKHOPLQJ PDMRULW\ RI WKH LVRODWHG VWUDLQV RIArthrospira ZHUH LGHQWLILHG DV A. platensis; RQH VWUDLQ ZDV LGHQWLILHG DV A. platensis YDU minor DQG RQH RWKHUDVA. maxima 7RPDVHOOLHW DO D :LWKLQWKHSKRWRV\QWKHVL]LQJSRSXODWLRQ RIWKHSHUPDQHQWDQGWHPSRUDU\ODNHV,OWLVDQG5LRX 'XZDW F REVHUYHG D YHU\ KLJK DOEHLW VHDVRQDOO\ YDULDEOH GHQVLW\ RI URWLIHUV PDLQO\ Brachionus VSS XS WR VHYHUDO KXQGUHG LQGLYLGXDOV SHU OLWHU ,Q VRPH ODNHV Arthrospira FRQVWLWXWHV DQ H[FHOOHQW IHHG IRU WKH FLFKOLG ILVK Tilapia. $QRWKHU FRQVXPHU RIArthrospira DQG RWKHU ODUJH ILODPHQWRXV F\DQREDFWHULD VXFK DV Anabaenopsis LV WKH IODPLQJR (Phoenicopterus minor). 7KH SLQN FRORU RI WKHVH ELUGVFRPHV IURPWKHFDURWHQRLGSLJPHQWVRULJLQDWLQJ LQF\DQREDFWHULD 7KH GHWDLOHG VWXGLHV SHUIRUPHG E\ ,OWLV RYHU WKH FRXUVH RI VHYHUDO \HDUV KDYH HVWDEOLVKHG D GLUHFW FRUUHODWLRQ EHWZHHQ WKH ELRPDVV GHQVLW\ RI Arthrospiru DQG WKH VDOLQLW\ RI WKH ZDWHU 7KH PDVVLYH Arthrospira EORRPV RFFXU DW VDOLQLW\ OHYHOV IURP WR J / KLJK FDUERQDWHELFDUERQDWH FRQFHQWUDWLRQVS+ DQGWHPSHUDWXUHVIURP & *HQHUDOO\ WKH ZDWHUV SRSXODWHG E\ Arthrospiru KDYH D PHDQ VDOLQLW\ RI J / 1HYHUWKHOHVV Arthrospiru KDV EHHQIRXQG WR RFFXUDW VDOLQLW\OHYHOVUDQJLQJIURP WRJ/ LQVRPH FDVHV XS WR J / ,OWLV %LRPDVV GHQVLW\ FDQH[FHHGJ/ 7XLWH IRXQG WKHGHQVLW\RIArthrospira WREH KLJKO\ YDULDEOH 7KH GHFOLQH LQ WKH Arthrospira SRSXODWLRQ ZDV SUHFHGHG E\ D SHULRG RI LQFUHDVHG ZDWHU FRQGXFWLYLW\ ,Q PDQ\ RI WKH QDWURQ ODNHV Arthrospira ZDV DEVHQW RU SUHVHQW RQO\ DW ORZ GHQVLWLHV 0DUNHGIOXFWXDWLRQVZHUHUHFRUGHGLQRWKHU ODNHV ,Q VRPH FDVHV Arthrospira PDLQWDLQHG KLJK GHQVLWLHV IRU SHULRGV XS WR VHYHUDO \HDUV DQG DSSHDUHG WR EH WKH RQO\ VLJQLILFDQW F\DQREDFWHULDO VSHFLHVLQWKHSK\WRSODQNWRQ Arthrospira ELRPDVV GHQVLW\ ZDV ORZHU LQ WKH WHPSRUDU\ ODNHV WKDQ LQ WKH SHUPDQHQW ODNHV EXW UHDFKHG YHU\KLJKOHYHOV XSWR J/ LQWKH IRUPHU LPPHGLDWHO\ SULRU WR WKHLU GHVLFFDWLRQ 7KH SK\WRSODQNWRQ RI WKH WHPSRUDU\ ZDWHU ERGLHV LV XVXDOO\ GRPLQDWHG E\ F\DQREDFWHULD DQG GLDWRPV Arthrospiru EORRPVWHQG WR GHYHORS DIWHU D EORRPRI JUHHQ IODJHOODWHV DQG RWKHU PLFURDOJDH IROORZLQJ WKH UHILOOLQJ RIDODNHE\UDLQZDWHU ,OWLV E D Arthrospira IORXULVKHV DOO \HDU URXQG ZLWK D VPDOO GHFOLQH GXULQJ WKH VKRUW DQG VFDUFH UDLQ\ VHDVRQ LQ WKH SHUPDQHQW ODNHV EXW ODVWV QR PRUH WKDQ WZR PRQWKVLQWKHWHPSRUDU\ ZDWHUERGLHV ,QWKHVHODNHV
WKH GHQVLW\ RI WKH SRSXODWLRQV GHSHQGV QRW RQO\ RQ WKH PHDQ VDOLQLW\ EXWDOVR RQ WKH WLPH SHULRG GXULQJ ZKLFKWKHEDVLQLV IORRGHG Arthrospira EORRPV ZHUH QRW REVHUYHG LQ ZDWHU ERGLHV IORRGHG IRU OHVV WKDQ IRXUPRQWKV
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platensis LV GRPLQDQW DV LW LV LQ WKH &KDG DUHD ,Q /DNH(OPHQWHLWDDQG/DNH1DNXUX *UDQWDQG 7LQGDOO IRXQG A. platensis WR FRQVWLWXWH WKH PDMRU EORRP DOWKRXJK WKH\ DOVR REVHUYHG VLJQLILFDQW QXPEHUV RI RWKHU F\DQREDFWHULD DV ZHOO DV GLDWRPV ZKLFKEHFDPHGRPLQDQWDVWKHF\DQREDFWHULDO EORRPV GHFUHDVHG ,Q KLV UHYLHZ RQ "Spirulina", &LIHUUL VXPPDUL]HG PRVW RI WKH VWXGLHV RQ WKH GLVWULEXWLRQ RI WKLV F\DQREDFWHULXP LQ $IULFD ZLWK SDUWLFXODU HPSKDVLV RQ WKH DONDOLQH ODNHV RI WKH 5LIW 9DOOH\ Arthrospira LV DOVR IRXQG LQ VRPH ODNHV RI WKH QRUWKHUQ $OJHULDQ 6DKDUD KDYLQJ VDOLQH ZDWHUV DQG S+ OHYHOV UDQJLQJ IURP WR 7KHVH ZDWHUV ZKLFK KDYH ORZHU DONDOLQLW\ DERXW PHT / +&2 SOXV &2 EXW 1D OHYHOV VLPLODU WR WKRVH RIWKH KLJKO\ DONDOLQH VDOLQH ZDWHUV RIWKH (DVW $IULFDQ5LIW9DOOH\ PD\EHFRQVLGHUHGWREH VXOIDWH
FKORULGHULFK ZDWHUV ZLWK& WKHPDMRUDQLRQ )RWW DQG .DULP IRXQG WKDW WKH SK\WRSODQNWRQ SRSXODWLRQ RIDVDOLQH DONDOLQH ODNH S+ DERXW LQ -HEHO 0DUUD 6XGDQ FRPSULVHG RQO\ RQH F\DQREDFWHULXP ZKLFK WKH\ QDPHG DV Spirulina geitleri. /DWHU )RWW LGHQWLILHG WKLV VSHFLHV DV S. fusiformis WRGD\ A. fusiformis). +LQGiN DVVLJQHG WKH Arthrospira SRSXODWLRQV RFFXUULQJ LQ /DNH%RJRULD.HQ\D WREHS. fusiformis.
' :DWHU%DVLQVRIWKH 6RODU(YDSRUDWRUDW /DNH7H[FRFR0H[LFR $V SUHYLRXVO\ PHQWLRQHG D PDVVLYH SRSXODWLRQ RI Arthrospiru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maxima LV SUHVHQW LQ DOPRVW PRQRVSHFLILFSRSXODWLRQVEHLQJDVVRFLDWHG ZLWKRQO\ D IHZ RWKHU F\DQREDFWHULD VXFK DV Synechocystis aquatilis.
9 3K\VLRORJ\RIArthrospira (FRORJLFDO VWXGLHV SHUIRUPHG GXULQJ WKH ODVW WKUHH GHFDGHV KDYH OHG WR XQGHUVWDQGLQJ RIWKH FRQGLWLRQV WKDW FDQ IDYRU LWV DEXQGDQW JURZWK LQ SK\WRSODQNWRQ SRSXODWLRQV 7KLV LQIRUPDWLRQ KDV EHHQ XVHIXO IRU GHYHORSLQJ WKH WHFKQRORJ\ IRU PDVV FXOWLYDWLRQ RI Arthrospiru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
$9RQVKDNDQG/7RPDVHOOL FKDQJHV
$ *URZWK
ZKHUH[LVWKHLQLWLDOELRPDVVFRQFHQWUDWLRQ 2JDZDDQG 7HUXL PDGH WKH ILUVWDVVHVVPHQW RITXDQWXP\LHOG
&KDSWHU $UWKURVSLUD 6SLUXOLQD 7KH UHODWLRQVKLS EHWZHHQ WKH VSHFLILF JURZWK UDWH DQGWKHVSHFLILFOLJKWHQHUJ\DEVRUSWLRQUDWHZDVXVHG WR HVWDEOLVK D PDWKHPDWLFDO HTXDWLRQ GHVFULELQJ WKH JURZWK RI Arthrospira LQ D EDWFK FXOWXUH ,HKDQD 7KHHTXDWLRQLQGLFDWHVWKDWLQFXOWXUHV ZLWKD KLJK FHOO FRQFHQWUDWLRQV WKH VSHFLILF JURZWK UDWH LQFUHDVHV OLQHDUO\ ZLWK WKH LQFUHDVH LQ WKH VSHFLILF DEVRUSWLRQ UDWH RI OLJKW HQHUJ\ 7ZR RWKHU JURXSV /HH HW DO DQG &RUQHW HW DO D E KDYH SXEOLVKHG GHWDLOHG VWXGLHV RQ DWWHPSWV WR HVWDEOLVK D PDWKHPDWLFDO PRGHO IRU WKH JURZWK RI Arthrospira XQGHU D YDULHW\ RI JURZWK FRQGLWLRQV ,W VHHPV WKDW DOO RI WKH PRGHOV ILW WKH H[SHULPHQWDO JURZWK GDWD ZHOO XQGHU QRUPDO VWHDG\ VWDWH FRQGLWLRQV ZKHUH OLJKWLVHLWKHUOLPLWLQJRUDWLWVVDWXUDWLRQOHYHO 7KHVH PRGHOVUHTXLUH PRGLILFDWLRQLIVWUHVV FRQGLWLRQV VXFK DV SKRWRLQKLELWLRQ RU HQYLURQPHQWDO VWUHVV LH WHPSHUDWXUHRUVDOLQLW\ DUHLQWURGXFHG % 5HVSRQVH WR (QYLURQPHQWDO )DFWRUV (IIHFWV RI /LJKW D (IIHFWV RQ *URZWK DQG 3KRWRV\QWKHVLV =DUURXN ZDV WKH ILUVWWR VWXG\ WKHUHVSRQVH RI “Spirulina maxima” WR OLJKW KH UHDFKHG WKH FRQFOXVLRQ WKDW JURZWK RIWKLV RUJDQLVP VDWXUDWHV DW OHYHOV RI .OX[
Fig. 2. 7\SLFDO FXUYH RI VSHFLILF SKRWRV\QWKHWLF UDWH 3 DV D IXQFWLRQ RI OLJKW LQWHQVLW\ , LOOXVWUDWLQJ WKH PD[LPXP SKRWRV\QWKHWLFUDWH3PD[ WKHVDWXUDWLRQRQVHWSDUDPHWHU,N DQG WKHVORSH RIWKHOLQHDUSDUWRIWKHFXUYH3,FXUYH
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rthrospira FXOWXUHV WR KLJK SKRWRQ IOX[ GHQVLWLHV DERYH WKH VDWXUDWLRQ SRLQWPD\ UHVXOW LQ D UHGXFHG UDWH RI SKRWRV\QWKHVLV WKLV SKHQRPHQRQ LV GHILQHG DV SKRWRLQKLELWLRQ 7KH FODVVLFDOYLHZWKDWSKRWRLQKLELWLRQLVREVHUYHG RQO\DW KLJK LUUDGLDQFH YDOXHV VHHPV WR EH D YHU\ VLPSOLVWLF RQH 3KRWRLQKLELWLRQ PD\ EH REVHUYHG HYHQ DW UHODWLYHO\ ORZ LUUDGLDQFH OHYHOV ZKHQ RWKHU HQYLURQPHQWDO VWUHVVHV DUH LQYROYHG 7KLV ZLOO EH GLVFXVVHGODWHU 7KH 3 DQG ,N OHYHOV DUH KLJKO\ GHSHQGHQW RQ JURZWK FRQGLWLRQV Arthrospira FXOWXUHV JURZQ DW KLJK RU ORZ OLJKW LQWHQVLWLHV ZLOO KDYH GLIIHUHQW 3 DQG,NYDOXHV &KDQJHVLQWKHVHYDOXHVPD\UHSUHVHQW WKH RUJDQLVP¶V DELOLW\ WR SKRWRDGDSW RU DFFOLPDWH WR GLIIHUHQW OLJKW HQYLURQPHQWV DV GHPRQVWUDWHG LQ DQRWKHU F\DQREDFWHULXP S. subsalsa JURZQ DW WZR SKRWRQIOX[ GHQVLWLHV 7RPDVHOOL HWDO 7DEOH VXPPDUL]HVH[SHULPHQWV FDUULHG RXWE\9RQVKDNHW DO D LQGLFDWLQJ GLIIHUHQW YDOXHV RI PHDVXUHG SKRWRV\QWKHWLF SDUDPHWHUV LQ WKUHH GLIIHUHQW Arthrospira VWUDLQV $OWKRXJKWKHVWUDLQV ZHUH JURZQ
$9RQVKDNDQG/7RPDVHOOL
XQGHUWKHVDPHWHPSHUDWXUHDQGOLJKWFRQGLWLRQV WKH\ KDYH GLIIHUHQW DQG ,N YDOXHV 6LPLODU REVHUYDWLRQV ZHUHUHSRUWHGE\7RPDVHOOLHWDOD
(IIHFWVRI7HPSHUDWXUH D (IIHFWRI7HPSHUDWXUHRQ *URZWK
E /LJKW6WUHVV 3KRWRLQKLELWLRQ 3KRWRLQKLELWLRQ DV PHQWLRQHG HDUOLHU LV GHILQHG DV D ORVVRISKRWRV\QWKHWLFFDSDFLW\GXHWRGDPDJHFDXVHG E\ SKRWRQ IOX[ GHQVLWLHV 3)'V LQ H[FHVV RI WKRVH UHTXLUHG WR VDWXUDWH SKRWRV\QWKHVLV 7KH SKHQRPHQRQ RI SKRWRLQKLELWLRQ KDV EHHQ VWXGLHG H[WHQVLYHO\ DQG LV ZHOO GRFXPHQWHG LQ DOJDH DQG KLJKHUSODQWV 3RZOHV .\OH DQG 2KDG 3KRWRLQKLELWLRQ LQ ODERUDWRU\ Arthrospiru FXOWXUHV ZDV ILUVW VWXGLHG E\ .DSODQ ZKR REVHUYHG D UHGXFWLRQ LQ WKH SKRWRV\QWKHWLF DFWLYLW\ ZKHQ WKH FHOOVZHUHH[SRVHGWRVWURQJOLJKWXQGHU&2GHSOHWHG FRQGLWLRQV ,W ZDV VXJJHVWHG WKDW WKH UHGXFWLRQ LQ SKRWRV\QWKHWLF DFWLYLW\ ZDV GXH WR WKH DFFXPXODWLRQ RI+2 'LIIHUHQW VWUDLQV RIArthrospiru PD\ GLIIHU LQ WKHLU VHQVLWLYLW\ WROLJKWVWUHVV 9RQVKDNHWDO E ,Q DWOHDVWRQHVWUDLQVXFKDGLIIHUHQFHZDVSUREDEO\GXH WR D GLIIHUHQFH LQ WKH WXUQRYHU UDWH RI D VSHFLILF SURWHLQ ' ZKLFK LV SDUW RI 36,, 7KH GLIIHUHQW UHVSRQVHV RI GLIIHUHQW Arthrospiru VWUDLQV WR SKRWRLQKLELWRU\ VWUHVV PD\ EH JHQRW\SLF FKDUDFWHULVWLFV DV ZHOO EHLQJ GHSHQGHQW RQ JURZWK FRQGLWLRQV &XOWXUHV JURZQ DW KLJK OLJKW LQWHQVLWLHV H[KLELW D KLJKHU UHVLVWDQFH WR SKRWRLQKLELWLRQ 9RQVKDNHWDO D ,WVKRXOGEHHPSKDVL]HGWKDW SKRWRLQKLELWLRQ DIIHFWV QRW RQO\ WKH 3 OHYHO EXW DFWXDOO\ KDV D VWURQJHU HIIHFW RQ OLJKWOLPLWHG SKRWRV\QWKHWLF DFWLYLW\ 7KXV D VLJQLILFDQW UHGXFWLRQ LQ LVREVHUYHGLQSKRWRLQKLELWHGFXOWXUHV Table 4. 6SHFLILFJURZWKUDWH DQGSKRWRV\QWKHWLFSDUDPHWHUVRI WKUHH Arthrospira VWUDLQV K , LUUDGLDQFHDWRQVHWRIOLJKW VDWXUDWLRQ SPRO SKRWRQ P V 3PD[ PD[LPDO OLJKWVDWXUDWHG UDWH RI 2 HYROXWLRQ PRO K PJ &KO D OLJKWOLPLWHG VORSH RI WKH 3, FXUYH PRO PJ &KO DPRO SKRWRQ P 9DOXHVDUHPHDQ± 6(Q
3$5$0(7(56 675$,16 %3
33
=
,N
3PD[
Fig. 3. 7HPSHUDWXUHUHVSRQVH RI Spirulina LVRODWHV PHDVXUHG DV WKHLQFUHDVHLQ&KORURSK\OOFRQFHQWUDWLRQ DIWHULQFXEDWLQJWKH FHOOV IRU GD\V RQ DWHPSHUDWXUHEORFN XQGHUFRQVWDQW OLJKW SPRO P V
,Q QDWXUH Arthrospira LV IRXQG LQ SHUPDQHQW RU WHPSRUDU\ ZDWHU ERGLHV DW UHODWLYHO\ KLJK WHPSHUDWXUHV 7KHRSWLPDOWHPSHUDWXUHIRUODERUDWRU\ FXOWLYDWLRQ RI WKLV RUJDQLVP UDQJHV IURP & +RZHYHU PDQ\ Arthrospira VWUDLQV GLIIHU LQ WKHLU RSWLPDO JURZWK WHPSHUDWXUH DV ZHOO DV LQ WKHLU VHQVLWLYLW\ WR H[WUHPH YDOXHV )RU LQVWDQFH LQ WKH FRPSDULVRQRIVWUDLQVVKRZQLQ)LJ VWUDLQ6$KDV D UHODWLYHO\ ORZ RSWLPDO JURZWK WHPSHUDWXUH & ZKLOH WKH (< VWUDLQ JURZV ZHOO XS WR & 7KH LVRODWH PDUNHG . KDV D UHODWLYHO\ ZLGH UDQJHRIRSWLPDOJURZWKWHPSHUDWXUHV & *URZWK WHPSHUDWXUH DOVR DIIHFWV WKH ELRFKHPLFDO FRPSRVLWLRQ RI WKH FHOOV 7RPDVHOOL HW DO :KHQ ³S. platensis” 0 LV JURZQ XQGHU OLJKWOLPLWHG WXUELGRVWDWLF FRQGLWLRQV DW LWV PD[LPXP JURZWK WHPSHUDWXUH & D PDUNHG GHFUHDVH LQ WKH SKRWRV\QWKHWLF SLJPHQWV DQG SURWHLQV OHYHO LV REVHUYHG 8QGHU WKHVH FRQGLWLRQV DQ LQFUHDVH LQ FDUERK\GUDWH FHOO FRQWHQW DVVRFLDWHG ZLWK FKDQJHV LQ WKH GHJUHH RI IDWW\ DFLG VDWXUDWLRQ LH UHGXFWLRQ RI OLQROHQLF V\QWKHVLV LQ IDYRU RI OLQROHLF DFLG DFFXPXODWLRQ LVDOVRREVHUYHG
E (IIHFWRI7HPSHUDWXUHRQ 3KRWRV\QWKHVLV DQG 5HVSLUDWLRQ 7KH QHW SURGXFWLYLW\ RI DQ DOJDO FXOWXUH LV GLUHFWO\ FRUUHODWHG ZLWK WKH JURVV UDWH RI&2 IL[DWLRQ RU 2 HYROXWLRQ SKRWRV\QWKHVLV DQG ZLWK WKH UDWH RI
&KDSWHU $UWKURVSLUD 6SLUXOLQD UHVSLUDWLRQ 7KHVH WZR PHWDEROLF DFWLYLWLHV DUH WHPSHUDWXUH GHSHQGHQW ZKLOH RQO\ &2 IL[DWLRQ RU 2 HYROXWLRQ LV ERWK OLJKW DQG WHPSHUDWXUH GHSHQGHQW $ GHWDLOHG VWXG\ RI WKH UHVSRQVH RI DQ Arthrospira VWUDLQPDUNHG0 SHUIRUPHGE\7RU]LOOR DQG 9RQVKDN GHWHUPLQHG WKH RSWLPDO WHPSHUDWXUH IRU SKRWRV\QWKHVLV & DQG DOVR PHDVXUHG WKH HIIHFW RI WHPSHUDWXUH RQ WKH GDUN UHVSLUDWLRQ UDWH RI Arthrospira E\ IROORZLQJ WKH 2 XSWDNH UDWH LQ WKH GDUN $ WHPSHUDWXUHGHSHQGHQW H[SRQHQWLDO UHODWLRQVKLS ZDV REWDLQHG ZLWK WKH UHVSLUDWLRQ UDWH LQFUHDVLQJ DV WHPSHUDWXUH LQFUHDVHG 7KH WHPSHUDWXUHGHSHQGHQW GDUN UHVSLUDWLRQ UDWH LV JLYHQE\ T R H ZKHUH5LVWKHUHVSLUDWLRQUDWHPRO2 PJ FK K DQG 7 LV WKH WHPSHUDWXUH & $W & DQG & GDUN UHVSLUDWLRQ UDWHV GURSSHG DOPRVW WR ]HUR $Q $UUKHQLXV SORW IRU UHVSLUDWLRQ VKRZHG DQ DFWLYDWLRQ HQHUJ\ RI N- PRO IRU Arthrospira. 7KH WHPSHUDWXUH FRHIILFLHQW 4 FDOFXODWHG IRU WKH & UDQJH ZDV 7KH UHVSLUDWLRQ WR SKRWR V\QWKHVLV UDWLR LQ Arthrospira ZDV DW & DQG DW & 7RU]LOOR DQG 9RQVKDN 7KHVH ORZ YDOXHV FRQILUP WKH JHQHUDO DVVXPSWLRQ WKDW F\DQREDFWHULD KDYH ORZ UHVSLUDWLRQ UDWHV YDQ /LHUH DQG 0XU 7KH UHVSLUDWLRQWRSKRWRV\QWKHVLV UDWHV PHDVXUHG LQ WKHVH H[SHULPHQWV ZHUH PXFK ORZHU WKDQ WKRVH UHSRUWHG IRU RXWGRRU FXOWXUHV RI Arthrospira, ZKHUH XS WR RI WKH ELRPDVV SURGXFHG GXULQJ GD\OLJKW PD\ EH ORVW WKURXJK UHVSLUDWLRQDWQLJKW*XWHUPDQHWDO 7RU]LOORHW DO ,QWKHArthrospira VWUDLQ0WKHRSWLPXP WHPSHUDWXUH IRU UHVSLUDWRU\ DFWLYLW\ ZDV PXFK KLJKHU WKDQ IRU SKRWRV\QWKHWLF DFWLYLW\ 1HYHUWKHOHVV WKH SKRWRV\QWKHWLF DFWLYLW\RIWKHFHOOV ZDVPRUHUHVLVWDQW WRWHPSHUDWXUHH[WUHPHV WKDQ ZDV GDUNUHVSLUDWLRQ DW WKHPLQLPXPDQGPD[LPXPWHPSHUDWXUHVWHVWHG
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
WHPSHUDWXUH LQ WKH OLJKW ,W LV WKXV VXJJHVWHG WKDW Arthrospiru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rthrospira VSHFLHV KDYH EHHQ LVRODWHG IURP D YDULHW\ RIDONDOLQH HQYLURQPHQWV :H ZLOOGHVFULEHWKH ZRUNGRQHXVLQJ VWUDLQVLVRODWHGIURPDONDOLQHDQGEUDFNLVKZDWHUV
D (IIHFWRI6DOLQLW\RQ *URZWK 7KHRFFXUUHQFHRIArthrospira VSS LQPDULQHKDELWDWV KDV QRW EHHQ GRFXPHQWHG DW OHDVW QRW DV GRPLQDQW SRSXODWLRQVOLNHWKRVHIRXQGLQWKHQDWURQODNHV 7KLV LV PRVW OLNHO\ GXH PRUH WR WKH YHU\ ORZ FRQWHQW RI ELFDUERQDWH WKDQ WR WKH KLJK FRQWHQW RI 1D&O Arthrospiru, XQOLNH PDULQH RUJDQLVPV GRHV QRW UHTXLUH KLJK 1D&O FRQFHQWUDWLRQ IRU JURZWK UDWKHU LW RQO\ WROHUDWHV WKH VDOW 7KH H[SRVXUH RI Arthrospira FXOWXUHVWRKLJK1D&OFRQFHQWUDWLRQV DWILUVWUHVXOWVLQ DQ LPPHGLDWH FHVVDWLRQ RIJURZWK DIWHU DODJ SHULRG DQHZ VWHDG\ VWDWH RIJURZWK LV HVWDEOLVKHG 9RQVKDN HW DO E 7KH OHQJWK RI WKH WLPH ODJ LV H[SRQHQWLDOO\ FRUUHODWHG WR WKH GHJUHH RI VDOLQLW\ VWUHVVLPSRVHGRQWKHFHOOV ,QPDQ\FDVHVWKLVODJLV DVVRFLDWHG ZLWK DGHFOLQH LQ FKORURSK\OO DQG ELRPDVV FRQFHQWUDWLRQVLQWKHFXOWXUH9RQVKDNHWDO E 7KH UHVSRQVH RI Arthrospira WR VDOLQLW\ LQ WHUPV RI WKH GHJUHH WR ZKLFK LWV JURZWK LV LQKLELWHG LWV DGDSWDWLRQ WR VDOW OHYHOV DQG WKH UDWH RI VXFK DGDSWDWLRQ YDULHV ZLGHO\ GHSHQGLQJ RQ WKH VWUDLQ XVHGLQWKHVWXG\ $GHFUHDVHLQWKH JURZWK UDWH GXH WR VDOW VWUHVV KDV DOVR EHHQ GHPRQVWUDWHG LQ RWKHU F\DQREDFWHULD VXFK DV Anacystis 9RQVKDN DQG 5LFKPRQG DQGNostoc %OXPZDOGDQG7HO2U
E (IIHFWRI6DOLQLW\RQ 3KRWRV\QWKHVLVDQG 5HVSLUDWLRQ ,WKDVEHHQVXJJHVWHGWKDWH[SRVXUHWRKLJKVDOLQLW\LV DFFRPSDQLHG E\ D KLJKHU GHPDQG IRU HQHUJ\ E\ WKH VWUHVVHG FHOOV %OXPZDOG DQG 7HO2U &KDQJHV LQ WKH SKRWRV\QWKHWLF DQG UHVSLUDWRU\ DFWLYLWLHV RI DQ Arthrospira VWUDLQ ZHUH PHDVXUHG RYHUDSHULRGRIKEHJLQQLQJPLQDIWHUH[SRVXUH WR DQG 0 1D&O $ PDUNHG GHFUHDVH LQ WKH SKRWRV\QWKHWLF R[\JHQ HYROXWLRQ UDWH ZDV REVHUYHG PLQ DIWHUH[SRVXUHWR WKH VDOW 7KLV GHFOLQH ZDV IROORZHG E\ D UHFRYHU\ SHULRG FKDUDFWHUL]HG E\ D ORZHU VWHDG\ VWDWH UDWH RI SKRWRV\QWKHVLV 9RQVKDN E 7KH LPPHGLDWHLQKLELWLRQRIWKHSKRWRV\QWKHWLF DQG UHVSLUDWRU\ V\VWHPV DIWHU H[SRVXUH WR VDOW VWUHVV KDYH EHHQ H[SODLQHG E\ (KUHQIHOG DQG &RXVLQ DQG 5HHG HW DO UHVSHFWLYHO\ LQ Dunaliella DQG LQ Synechocystis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
F 2VPRUHJXODWLRQDQG6WUDLQ6SHFLILF 5HVSRQVH RI$UWKURVSLUD WR 6DOLQLW\ 'XULQJ WKH FRXUVH RI DGDSWDWLRQ WR VDOLQLW\ DQ RVPRWLF DGMXVWPHQW LV UHTXLUHG ,Q Arthrospira, ORZ PROHFXODU ZHLJKW FDUERK\GUDWHV DFFXPXODWH 7KHVH KDYH EHHQ LGHQWLILHG DV D QLQH FDUERQ KHWHURVLGH FDOOHG JOXFRV\OJO\FHURO SOXV D WUHKDORVH 0DUWHO HW DO :KHQ WKH ELRPDVV FRPSRVLWLRQV RI WZR Arthrospira VWUDLQV JURZQ XQGHU GLIIHUHQW VDOW VWUHVV FRQGLWLRQV ZHUH FRPSDUHG VLJQLILFDQW FKDQJHV PDLQO\DQLQFUHDVHLQFDUERK\GUDWHVDQGDGHFUHDVHLQ WKH SURWHLQ OHYHO ZHUH REVHUYHG 7KHVH FKDQJHV FRUUHODWHG ZLWK WKH GHJUHH RI VWUHVV LPSRVHG LH D KLJKHU OHYHO RI FDUERK\GUDWHV DW D KLJKHU VDOW FRQFHQWUDWLRQ 7KH GLIIHUHQFH LQ WKH OHYHO RI FDUERK\GUDWHV DFFXPXODWHG E\ WKH WZR VWUDLQV PD\ DOVRUHIOHFWDGLIIHUHQFHLQWKHLUDELOLW\WRDGDSWWRVDOW
$9RQVKDNDQG/7RPDVHOOL VWUHVV &KDQJHV LQ OLSLG V\QWKHVLV KDYH DOVR EHHQ GHPRQVWUDWHG LQ VDOWVWUHVVHG FHOOV DQ LQFUHDVH LQ OLSLGFRQWHQWDQGLQWKHGHJUHHRIIDWW\DFLG VDWXUDWLRQ LV REVHUYHG ZLWK VSHFLILF PRGLILFDWLRQV LQ WKH ORQJ FKDLQIDWW\DFLGV & ,QArthrospira VWUDLQ0 WKH ROHLF DFLG FRQWHQW GRXEOHG ZKHQ WKH RUJDQLVP ZDV JURZQ LQ SUHVHQFH RI 0 1D&O 7RPDVHOOL HW DO D
& 7KH$ONDOLSKLOLF1DWXUH RI$UWKURVSLUD $V SUHYLRXVO\ LQGLFDWHG PRVW Arthrospira VSHFLHV ZHUH LVRODWHG IURP DONDOLQH DQG VDOLQH RU EUDFNLVK ZDWHUV FKDUDFWHUL]HG E\ KLJK OHYHOV RI FDUERQDWH ELFDUERQDWH DQG KLJK S+ OHYHOV 7KH SK\VLRORJLFDO FKDUDFWHULVWLFV ZKLFK HQDEOH D PLFURRUJDQLVP WR WKULYH LQ VXFK DQ H[WUHPH HQYLURQPHQW DUH VWLOO QRW IXOO\ XQGHUVWRRG ,W VKRXOG EH SRLQWHG RXW WKDW Arthrospira QRW RQO\ VXUYLYHV DW KLJK S+ YDOXHV EXW DFWXDOO\ WKULYHV LQ VXFK FRQGLWLRQV %HONLQ DQG %RXVVLED ZKR FRPSDUHG WKH JURZWK S+ RSWLPD IRU WKH WZR F\DQREDFWHULD Anabaena DQG Arthrospira, GHPRQVWUDWHG WKDW ZKLOH WKH RSWLPDO S+ IRU Anabaena ZDV LQ WKH UDQJH RI WKH PD[LPDO JURZWKUDWHIRUArthrospira ZDV REWDLQHG LQ WKH UDQJH :KHQ LQFXEDWHG DW S+ WKH JURZWKUDWHRIArthrospira ZDV VHYHUHO\ LQKLELWHG DQG ZDV RQO\ RI WKDW REWDLQHG XQGHU WKH RSWLPDO FRQGLWLRQV 7KLV KLJK S+ ! UHTXLUHPHQW FOHDUO\ GHILQHV Arthrospira DV DQ REOLJDWRU\ DONDOLSKLOH *UDQWHWDO 2QH RI WKH PDMRU SUREOHPV IDFHG E\ FHOOV LQ DQ DONDOLSKLOLF HQYLURQPHQW LV WKDW RI UHJXODWLQJ WKHLU LQWHUQDO S+ %HONLQ DQG %RXVVLED ZHUH WKH ILUVWWRPHDVXUH WKH DELOLW\ RIArthrospira WR PDLQWDLQ DS+ JUDGLHQWDFURVV LWV F\WRSODVPDWLF PHPEUDQH ,W ZDV GHPRQVWUDWHG WKDW DW H[WHUQDO S+ YDOXHV RI DQG WKH LQWUDFHOOXODU S+ YDOXHV ZHUH RQO\ DQGUHVSHFWLYHO\ 0DQ\ DONDOLSKLOHV UHTXLUH VRGLXP LQ RUGHU WR VXUYLYH LQ H[WUHPH DONDOLQH HQYLURQPHQWV +RULNRVKL DQG$NLED 3DGDQ HW DO GHPRQVWUDWHG WKDWDQ DFWLYH VRGLXPSURWRQ DQWLSRUWHULVUHTXLUHG LQ RUGHUWRPDLQWDLQDORZLQWHUQDOS+ ,QDPRUHUHFHQW VWXG\ 6FKOHVLQJHU HW DO GHPRQVWUDWHG WKDW "Spirulina platensis" UHTXLUHV VRGLXP LQ RUGHU WR PDLQWDLQ RSWLPDO JURZWK DW S+ 7KH DXWKRUV GHPRQVWUDWH WKDW XQGHU VRGLXP GHSULYDWLRQ WKH S+ JUDGLHQW FROODSVHV DQG WKH FHOOV XQGHUJR IDVW O\VLV 'HSULYLQJ WKH FHOOV RI VRGLXP FDXVHV LQDFWLYDWLRQ RI WKH VRGLXPSURWRQ DQWLSRUWHU WKDW LV UHVSRQVLEOH IRU WKH LQZDUG SXPSLQJ RI SURWRQV DQG WKH RXWZDUG SXPSLQJ RIVRGLXP 7KLV PHFKDQLVP LV RQH ZD\ WR
&KDSWHU $UWKURVSLUD 6SLUXOLQD PDLQWDLQ ORZHU LQWHUQDO S+ OHYHOV LQ FHOOV XQGHU DONDOLSKLOLFFRQGLWLRQV )XUWKHUVWXGLHVRQWKHHQHUJ\ UHTXLUHPHQW DQG LQYROYHPHQW RI OLJKW LQ WKH UHJXODWRU\SURFHVVDUH\HWWREHFDUULHGRXW
' +RZ'RHV$UWKURVSLUD &RPSHWHLQ &XOWXUH " 6WXGLHV SHUIRUPHG RQ GLIIHUHQW Arthrospira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rthrospira SOD\V D UROH LQ F\FOLQJ QXWULWLRQDO HOHPHQWV DQG LQ WKHLU SDVVDJH LQWR GLIIHUHQW IRRG FKDLQV PRUHRYHULWSURYLGHVDQLPSRUWDQWGLUHFWIRRG VRXUFH IRU ]RRSODQNWRQ ILVKHV ELUGV DQG KXPDQV 7KH VDPH IDFWRUV WKDW LQ WKH RULJLQDO KDELWDW UHJXODWH WKH RFFXUUHQFH RI Arthrospira LQ PRQRVSHFLILF SRSXODWLRQV DUH WKRVH UHTXLUHG IRU ERWK VXFFHVVIXOO\ PDLQWDLQLQJ D PRQRFXOWXUH LQ LQWHQVLYH SURGXFWLRQ SRQGV RXWGRRUV DQG IRU DFKLHYLQJ KLJK SURGXFWLYLW\ )DFWRUV VXFK DV KLJK DONDOLQLW\ DQG VDOLQLW\ DUH WKH NH\ IHDWXUHV WKDWVHOHFWLYHO\ OLPLWWKH JURZWK RIRWKHU RUJDQLVPVDQGSUHGDWRUV (YHQ LI KLJK S+ DQG VDOW FRQFHQWUDWLRQ UHGXFH FRQWDPLQDWLRQ IURP ERWK DLU DQG ZDWHU Arthrospira FXOWXUHV PD\ ZKHQ HIIHFWLYH SRQG PDQDJHPHQW LV ODFNLQJ EH JUHDWO\ FRQWDPLQDWHG E\ RWKHU RUJDQLVPV VXFK DV KHWHURWURSKLF EDFWHULD IXQJL SURWR]RD URWLIHUV RWKHU F\DQREDFWHULD DQG PLFURDOJDH &RQWDPLQDWLRQ RI RXWGRRU Arthrospira FXOWXUHV E\ RWKHU PLFURDOJDH LV RI IUHTXHQW RFFXUUHQFH WKHVH FRQWDPLQDQWV DUHXVXDOO\ JUHHQ DOJDH (Chlorella) DQG GLDWRPV $V WKH GLDWRPV OLYH WLJKWO\ DWWDFKHG WR
VXUIDFHVVXFKDVSDGGOHZKHHOV WKH\ DUHGLIILFXOWWR ZDVK RXWRIWKH FXOWXUH V\VWHP 0RUHSUREOHPDWLF LV WKH SUHVHQFH RI RWKHU ILODPHQWRXV Oscillatoria-like F\DQREDFWHULD ZKLFK KDYH VLPLODU HFRORJLFDO GHPDQGV DQGPD\LQYDGHWKHFXOWXUH $QLQFUHDVHLQ WKHRUJDQLFORDGVXEVHTXHQWWRGHWHULRUDWLRQSURFHVVHV FHOO O\VLV DOVR VWLPXODWHV WKH GHYHORSPHQW RI IXQJL DQG EDFWHULD 1HYHUWKHOHVV RQFH WKH RSWLPDO FRQGLWLRQV IRU WKH JURZWK RI Arthrospira DUH PDLQWDLQHG LQ SRQGV WKLV RUJDQLVP VXFFHVVIXOO\ RXW FRPSHWHV RWKHU RFFDVLRQDOO\ RFFXUULQJ SKRWRWURSKV IRU WKH SULPDU\ UHVRXUFHV DQG PD\ WKHQ UHDFK KLJK SRSXODWLRQ GHQVLWLHV 7KLV LV DWWULEXWDEOH WR LWV IDVW UDWHV RI JURZWK DQG SKRWRDFFOLPDWLRQ DQG LWV KLJK WROHUDQFH WR HQYLURQPHQWDO VWUHVV LUUDGLDQFH DONDOLQLW\VDOLQLW\WHPSHUDWXUH
9, &RQFOXGLQJ 5HPDUNV Arthrospira LV D FRVPRSROLWDQ F\DQREDFWHULXP \HW RFFXSLHVYHU\VHOHFWLYHKDELWDWV 7KHPDLQHFRORJLFDO GHPDQGV DUH VWDEOH DONDOLQH VDOLQH ZDWHUV ZDUP JURZWK WHPSHUDWXUHV KLJK LUUDGLDQFH DQG HXWURSKLF FRQGLWLRQV LH DEXQGDQFH RI ELFDUERQDWH DQG RWKHU PDFURQXWULHQWV &RQVHTXHQWO\ SULPDU\ SURGXFWLYLW\ RI Arthrospira LQ VRPH WURSLFDO DQG VXEWURSLFDO ODNHV FDQEHYHU\KLJK 7KH XVH RI Arthrospira DV IRRG E\ LQGLJHQRXV SRSXODWLRQV LQ GLIIHUHQW SDUWV RI WKH ZRUOG LV ZHOO GRFXPHQWHG ,Q UHFHQW \HDUV WKLV F\DQREDFWHULXP KDV JDLQHG DFRQVLGHUDEOHUHSXWDWLRQ DV DKHDOWK IRRG DGGLWLYH DQG LV PDUNHWHG FRPPHUFLDOO\ ,Q PDQ\ FDVHV WKH WHUP 6SLUXOLQD KDV EHFRPH D SUDFWLFDO V\QRQ\P IRU FXOWLYDWHG PLFURDOJDH $W SUHVHQW Arthrospira UHSUHVHQWV WKH VHFRQG PRVW LPSRUWDQW FRPPHUFLDO PLFURDOJD IRU WKH SURGXFWLRQ RIELRPDVV DVDKHDOWKIRRG DQGDQDQLPDO IHHGDIWHU Chlorella). :LWK WKH FXUUHQW UDSLG LQFUHDVH LQ SURGXFWLRQ DQG D GHPDQG UHDFKLQJ W LQ SUHGLFWHG WR UHDFK W E\ WKH EHJLQQLQJ RI WKH QHZ PLOOHQQLXP WKHUH LV QR GRXEW WKDW Arthrospira ZLOO EH WKH PRVW LPSRUWDQW PDVVSURGXFHG PLFURDOJD IRU FRPPHUFLDO SXUSRVHVLQWKHQHDUIXWXUH 7KH DFFXPXODWHG NQRZOHGJH RI WKH HFRORJ\ SK\VLRORJ\ DQG ELRFKHPLVWU\ RI Arthrospira KDV PDGH VXFK UHVXOWV IHDVLEOH \HW PXFK PRUH ZRUN LV UHTXLUHG LQ WHUPV RI VWUDLQ VHOHFWLRQ DQG EHWWHU XQGHUVWDQGLQJ RI WKH IDFWRUV ZKLFK JRYHUQ JURZWK DQG OLJKWXWLOL]DWLRQ LQ GHQVHFXOWXUHV WR VXSSRUW WKLV JURZLQJDJURLQGXVWU\
5HIHUHQFHV $QDJQRVWLGLV . DQG .RPiUHN - 0RGHP DSSURDFK WR WKH FODVVLILFDWLRQ V\VWHP RI F\DQRSK\WHV 2VFLOODWRULDOHV $UFK +\GURELRO6XSSO$OJRORJLFDO6WXGLHV %HDGOH /& 7KH ,QODQG :DWHUV RI 7URSLFDO $IULFD $Q ,QWURGXFWLRQWR7URSLFDO/LPQRORJ\/RQJPDQ/RQGRQ %HONLQ 6 DQG%RXVVLED 6 5HVLVWDQFHRISpirulina platensis WRDPPRQLDDWKLJKS+YDOXHV3ODQW&HOO3K\VLRO %OXPZDOG ( 0HKOKRUQ 5- DQG 3DFNHU / 6DOW DGDSWDWLRQ PHFKDQLVPV LQ WKH F\DQREDFWHULXP Synechococcus ,Q 6\EHVPD6 HG $GYDQFHVLQ3KRWRV\QWKHVLV 5HVHDUFKSS 0DUWLQXV1LMKRII'U:-XQN3XEOV7KH+DJXH %OXPZDOG ( DQG 7HO2U ( 2VPRUHJXODWLRQ DQG FHOO FRPSRVLWLRQ LQ VDOW DGDSWDWLRQ RI Nostoc muscorum. $UFK 0LFURELRO %RURZLW]ND /- 2VPRUHJXODWLRQ LQ EOXHJUHHQ DOJDH ,Q 5RXQG )( DQG &KDSPDQ ' HGV 3URJUHVV LQ 3K\FRORJLFDO 5HVHDUFK9ROSS%LRSUHVV/WG%ULVWRO %UDQGLO\ 0< 'HSXLV GHV OXVWUHV XQH WULEX SULPLWLYH GX 7FKDGH[SORLWHODQRXPWXUH GHO DQ 6FL $YHQLU %ULHED & DQG 0HULQRV & 6RPH DGYDQFHV LQ WKH Spirulina FXOWXUH LQ VHDZDWHU ,Q 3URJUHVV LQ %LRWHFKQRORJ\ RI 3KRWRDXWRWURSKLF0LFURRUJDQLVPV$EVWUWK,&$$S&HVNp %XGHMRYLFH&]HFK5HSXEOLF %XVVRQ ) Spirulina platensis *RP *HLWOHU HW Spirulina geitleri - GH 7RQL &\DQRSK\FpHV $OLPHQWDLUHV 6HUYLFH GH 6DQWp0DUVHLOOH &DUU1*DQG:KLWWRQ%$HGV 7KH%LRORJ\RI%OXH*UHHQ $OJDH%ODFNZHOO6FLHQWLILF2[IRUG &DQ 1* DQG :KLWWRQ %$ 7KH %LRORJ\ RI&\DQREDFWHULD %ODFNZHOO6FLHQWLILF2[IRUG &DVWHQKRO] 5: 6XEVHFWLRQ ,,, 2UGHU 2VFLOODWRULDOHV ,Q 6WDOH\ -7 %U\DQW 03 3IHQQLQJ 1 DQG +ROW -* HGV %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ 9RO SS :LOOLDPVDQG:LONLQV&R%DOWLPRUH &LIHUUL 2 Spirulina, WKH HGLEOH PLFURRUJDQLVP 0LFURELRO 5HY &OpPHQW* $QHZ W\SHRIIRRG DOJDH ,Q 0DWHOHV 5- DQG 7DQQHQEDXP 65 HGV 6LQJOHFHOO 3URWHLQ 0,7 3UHVV &DPEULGJH0DVV &RKHQ = DQG 9RQVKDN $ )DWW\ DFLG FRPSRVLWLRQ RI Spirulina DQG 6SLUXOLQDOLNH VWUDLQV LQ UHODWLRQ WR WKHLU FKHPRWD[RQRP\3K\WRFKHPLVWU\ &RPHW -) 'XVVDS &* &OX]HO 3 DQG 'XEHUWUHW * E $ VWUXFWXUHG PRGHO IRU VLPXODWLRQ RI FXOWXUHV RI WKH F\DQREDFWHULXP Spirulinu platensis LQ SKRWRELRUHDFWRUV ,, ,GHQWLILFDWLRQ RI NLQHWLF SDUDPHWHUV XQGHU OLJKW DQG PLQHUDO OLPLWDWLRQV%LRWHFKQRO%LRHQJLQ &RPHW -) 'XVVDS &* DQG 'XEHUWUHW * D $ VWUXFWXUHG PRGHOIRUVLPXODWLRQRIFXOWXUHVRIWKHF\DQREDFWHULXPSpirulina platensis LQSKRWRELRUHDFWRUV , &RXSOLQJEHWZHHQOLJKWWUDQVIHU DQGJURZWKNLQHWLFV%LRWHFKQRO%LRHQJLQ 'DQJHDUG3 6XUXQHDOJXHEOHXHDOLPHQWDLUHSRXUO KRPPH Arthrospira platensis 1RUGVW *RPRQW $FWHV 6RF /LQQ %RUGHDX[ 'H7RQL*% 1RWHUHOOHGLQRPHQFODWXUDDOJRORJLFD 'HVLNDFKDU\ 79 &\DQRSK\WD ,QGLDQ &RXQFLO IRU $JULFXOWXUDO5HVHDUFKSS1HZ'HOKL 'HVLNDFKDU\ 79 DQG -HHML %DL 1 7D[RQRPLF VWXGLHV LQ Spirulina ,Q 6HVKUDGL &9 DQG -HHML %DL 1 (GV Spirulina (77$1DWLRQDO6\PSRVLXPSS0&5&0DGUDV 'HVLNDFKDU\ 79 DQG -HHML %DL 1 7D[RQRPLF VWXGLHV LQ Spirulinu ,, 7KH LGHQWLILFDWLRQ RI Arthrospira ("Spirulina")
$9RQVKDNDQG/7RPDVHOOL VWUDLQV DQG QDWXUDO VDPSOHV RI GLIIHUHQW JHRJUDSKLFDO RULJLQV $UFK+\GURELRO6XSSO$OJRORJLFDO6WXGLHV (KUHQIHOG - DQG &RXVLQ -/ ,RQLF UHJXODWLRQ RI WKH XQLFHOOXODU JUHHQ DOJD Dunaliella tertiolecta: UHVSRQVH WR K\SHUWRQLFVKRFN -0HPEU%LRO (O%HVWDZ\ ( %HOOLQJHU (* DQG 6LJHH '& (OHPHQWDO FRPSRVLWLRQ RI SK\WRSODQNWRQ LQ D VXEWURSLFDO ODNH ;UD\ PLFURDQDO\WLFDOVWXGLHVRQWKHGRPLQDQWDOJDHSpirulina platensis &\DQRSK\WD DQG Cyclotella meneghiniana %DFLOODULRSK\FHDH (XU-3K\FRO )RJJ*(6WHZDUW:'3)D\3DQG:DOVE\$( 7KH%OXH *UHHQ$OJDH$FDGHPLF3UHVV/RQGRQ )RWW% 7D[RQRP\RISODQNWRQLF6SLUXOLQDVLQVDOLQHWURSLFDO ODNHV6FKZHL]=+\GURELRO )RWW% DQG.DULP*$$ Spirulina SODQNWRQFRPPXQLW\LQD /DNH LQ-HEHO0DUUD6XGDQ$UFK3URWLVWHQN )R[ 5' Spirulina 3URGXFWLRQDQG 3RWHQWLDO(GLVXG $L[ HQ3URYHQFH)UDQFH )U\,9+XIOHMW0(UEHU::$3HVFKHN*$DQG3DFNHU/ 7KH UROH RI UHVSLUDWLRQ GXULQJ DGDSWDWLRQ RI WKH IUHVKZDWHU F\DQREDFWHULXP Synechococcus WR VDOLQLW\ $UFK%LRFKHP %LRSK\V *DOOHJRV $- 7KH SDVW SUHVHQW DQG IXWXUH RI DOJDH LQ 0H[LFR%XOO,QVWRFpDQRJUDSK0RQDFR *DUGQHU 1/ 1HZ 3DFLILF FRDVW PDULQH DOJDH , 8QLY &DOLI 3XEO%RW *HLWOHU / &\DQRSK\FHDH ,Q 5DEHQKRUVW V .U\SWRJDPHQIORUD YRQ 'HXWVFKODQG gVWHUUHLFK XQG GHU 6FKZHL]9RO SS $NDG9HUODJ/HLS]LJ *LRYDQQRQL 6- 7XUQHU 6 2OVHQ *- %DUQV 6 /DQH '- DQG 3DFH 15 (YROXWLRQDU\UHODWLRQVKLSV DPRQJ F\DQREDFWHULDDQG JUHHQFKORURSODVWV-%DFWHULRO *RPRQW0 0RQRJUDSKLHGHV2VFLOODULpHV$QQ6FL1DW%RW 6p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pULHV3URWLVWRORJLFD *XWHUPDQ+9RQVKDN$DQG%HQ
&KDSWHU $UWKURVSLUD 6SLUXOLQD ,OWLV $ D 3K\WRSODQFWRQ GHV HDX[ QDWURQpHV GX .DQHP 7FKDG ,/HVODFVSHUPDQHQWVD6SLUXOLQHV&DK256720VHU +\GURELRO ,OWLV $ E 3K\WRSODQFWRQ GHV HDX[ QDWURQpHV GX .DQHP 7FKDG ,, /HV PDUHV WHPSRUDLUHV &DK 256720 VpU +\GURELRO ,OWLV $ D 3K\WRSODQFWRQ GHV HDX[ QDWURQpHV GX .DQHP 7FKDG ,,, 9DULDWLRQV DQQXHOOHV GX SODQFWRQ GXQH PDUH WHPSRUDLUH&DK256720VpU+\GURELRO ,OWLV $ E 3K\WRSODQFWRQ GHV HDX[ QDWURQpHV GX .DQHP 7FKDG ,9 1RWH VXU OHV HVSqFHV GX JHQUH Oscillatoria, VRXV JHQUH Spirulina &\DQRSK\WD &DK256720VpU+\GURELRO ,OWLV $ D 1RWH VXU Oscillatoria VRXVJHQUH Spirulina) platensis 1RUGVW %RXUUHOO\ &\DQRSK\WD DX 7FKDG &DK 256720VpU+\GURELRO ,OWLV $ E 3K\WRSODQFWRQ GHV HDX[ QDWURQpHV GX .DQHP 7FKDG 9/HVODFVPHVRKDOLQV&DK256720VpU+\GURELRO ,OWLV $ $OJXHV GHV HDX[ QDWURQpHV GX .DQHP 7FKDG , 3UHPLpUHSDUWLF&DK256720VHU+\GURELRO ,OWLV $ DQG 5LRX'XZDW 6 F 9DULDWLRQV VDLVRQQLpUHV GX SHXSOHPHQWHQ URWLIHUHV GHV HDX[QDWURQpHV GX .DQHP7FKDG &DK256720VpU+\GURELRO -HHML %DL 1 &RPSHWLWLYH H[FOXVLRQ RU PRUSKRORJLFDO WUDQVIRUPDWLRQ" $ FDVH VWXG\ ZLWK Spirulina fusiformis. $UFK +\GURELRO6XSSO$OJRORJLFDO6WXGLHV -HHML %DL 1 DQG 6HVKDGUL &9 2QFRLOLQJDQG XQFRLOLQJ RI WULFKRPHV LQ WKH JHQXV Spirulina. $UFK +\GURELRO 6XSSO $OJRORJLFDO6WXGLHV .DSODQ$ 3KRWRLQKLELWLRQLQ Spirulina platensis: UHVSRQVH RISKRWRV\QWKHVLV DQG +&2 XSWDNH FDSDELOLW\ WR &2 GHSOHWHG FRQGLWLRQV-([SHU%RW .HEHGH(DQG$KOJUHQ * 2SWLPXPJURZWKFRQGLWLRQV DQG OLJKW XWLOLVDWLRQ HIILFLHQF\ RI Spirulina platensis HTXDO Arthrospirafusiformis) &\DQRSK\WD IURP/DNH&KLWX(WKLRSLD +\GURELRORJLD .RPiUHN-DQG/XQG-:* :KDWLV"Spirulina platensis" LQ IDFW"$UFK+\GURELRO6XSSO$OJRORJLFDO6WXGLHV .\OH '- DQG 2KDG , 7KH PHFKDQLVP RISKRWRLQKLELWLRQ LQ KLJKHUSODQWV DQGJUHHQ DOJDH ,Q 6WDHKHOLQ/$DQG $UQW]HQ&- HGV (QF\FORSHGLD RI 3ODQW 3K\VLRORJ\ 1HZ 6HULHV 9RO 3KRWRV\QWKHVLV,,,SS6SULQJHU9HUODJ%HUOLQ /HH +< (ULFNVRQ /( DQG
0HODFN -0 3KRWRV\QWKHVLV DQG JURZWK RI Spirulina platensis &\DQRSK\WD LQ DQ HTXDWRULDO ODNH /DNH 6LPEL .HQ\D /LPQRO2FHDQRJU 0LQ 7KHLQ 3URGXFWLRQ RI6SLUXOLQD LQ 0\DQPDU %XOO ,QVW RFpDQRJUDSK0RQDFR 0ROLWRU 9 (UEHU : DQG 3HVFKHN *$ ,QFUHDVHG OHYHOV RI F\WRFKURPH R[LGDVH DQG VRGLXPSURWRQ DQWLSRUWHULQ WKH SODVPD PHPEUDQH RI Anacystis nidulans DIWHU JURZWK LQ VRGLXP HQULFKHGPHGLD)(%6/HWW 1HOLVVHQ % :LOPRWWH $ 1HHIV -0 DQG 'H :DFKWHU 5 3K\ORJHQHWLF UHODWLRQVKLS DPRQJ ILODPHQWRXV KHOLFDO F\DQREDFWHULDLQYHVWLJDWHG RQ WKHEDVLV RI 6 ULERVRPDO 51$ JHQHVHTXHQFHDQDO\VLV6\VWHP$SSO0LFURELRO 2JDZD 7 DQG $LED 6 &2 DVVLPLODWLRQ DQG JURZWK RI D EOXHJUHHQ DOJD Spirulina platensis, LQ FRQWLQXRXV FXOWXUH - $SSO&KHP%LRWHFKQRO 2JDZD 7DQG 7HUXL * 6WXGLHV RQ WKH JURZWK RISpirulina platensis , 2QWKHSXUHFXOWXUHRISpirulina platensis. -)HUPHQW 7HFKQRO 3DGDQ(=LOEHUVWHLQ'DQG6FKXOGLQHU6 S++RPHRVWDVLVLQ EDFWHULD%LRFKHP%LRSK\V$FWD 3RZOHV 6% 3KRWRLQKLELWLRQ RI SKRWRV\QWKHVLV LQGXFHG E\ YLVLEOHOLJKW$QQX5HY3ODQW3K\VLRO 5HHG 5+ 5LFKDUGVRQ '/ DQG 6WHZDUW :'3 1D XSWDNH DQGH[WUXVLRQLQWKH F\DQREDFWHULXP Synechocystis 3&&LQ UHVSRQVH WR K\SHUVDOLQH WUHDWPHQW (YLGHQFH IRU WUDQVLHQW FKDQJHV LQ SODVPDOHPPD 1D SHUPHDELOLW\ %LRFKLP %LRSK\V $FWD 5LFK) 1RWHVRQArthrospira platensis. 5HY$OJRO 5LFKPRQG$ Spirulina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rthrospira ('Spirulina') VWUDLQV IURP IRXU FRQWLQHQWV DUH UHVROYHG LQWR WZR RQO\ WZR FOXVWHUV EDVHG RQ $5'5$ $PSOLILHG 5LERVRPDO '1$ 5HVWULFWLRQ$QDO\VLV RIWKH ,76 ,QWHUQDOO\7UDQVFULEHG6SDFHU )(060LFURELRO/HWW 6FKOHVLQJHU3%HONLQ6DQG%RXVVLED6 6RGLXPGHSULYDWLRQ XQGHU DONDOLQH FRQGLWLRQV FDXVHV UDSLG GHDWK RIWKH ILODPHQWRXV F\DQREDFWHULXP Spirulina SODWHQVLV-3K\FRO 6KLPDGD$2JXFKL0 2WVXER.1LWWD. .R\DQR7DQG0LNL . $SSOLFDWLRQRIWXEXODUSKRWRELRUHDFWRUV\VWHP WR FXOWXUH Spirulina IRUIRRGSURGXFWLRQ DQG JDV H[FKDQJH ,Q 0L\DFKL 6 .DUXEH , DQG ,VKLGD < HGV &XUUHQW 7RSLFV LQ 0DULQH %LRWHFKQRORJ\SS-DS62& 0DU%LRWHFKQRO7RN\R 6WL]HQEHUJHU ( Spirulina XQG Arthrospira QRY JHQ +HGZLJLD 7RPDVHOOL / 6\VWHPDWLFDQG(FRORJ\ ,Q 9RQVKDN$ HG Spirulina platensis (Arthrospira): 3K\VLRORJ\ &HOOELRORJ\ DQG %LRWHFKQRORJ\SS7D\ORU )UDQFLV/WG/RQGRQ8. 7RPDVHOOL / %ROGULQL * DQG 0DUJKHUL 0& 3K\VLRORJLFDO EHKDYLRXURIArthrospira (Spirulina) maxima GXULQJ DFFOLPDWLRQ WRFKDQJHVLQLUUDGLDQFH-$SSO3K\FRO 7RPDVHOOL )HURFL / 0DUJKHUL 0& DQG 3HORVL ( 'LH 8OWUDVWUXNWXU YRQ Spirulina LP 9HUJOHLFK ]X Oscillatoria. =EO %DNW$EW,,
7RPDVHOOL / *LRYDQQHWWL / DQG 0DUJKHUL 0& 2Q WKH PHFKDQLVP RI WULFKRPH EUHDNDJH LQ Spirulina platensis DQG S. maxim. $QQ0LFURELRO 7RPDVHOOL / *LRYDQQHWWL / 6DFFKL$DQG%RFFL) (IIHFWV RI WHPSHUDWXUH RQ JURZWK DQG ELRFKHPLFDO FRPSRVLWLRQ LQ 6SLUXOLQDSODWHQVLV VWUDLQ0 ,Q 6WDHGOHU7 0ROOLRQ- 9HUGXV 0& .DUDPDQRV < 0RUYDQ + &KULVWLDHQ ' HGV $OJDO %LRWHFKQRORJ\ SS (OVHYLHU$SSOLHG 6FLHQFHV /RQGRQ DQG1HZ
$9RQVKDNDQG/7RPDVHOOL 9RQVKDN$E 6WUDLQVHOHFWLRQRISpirulina VXLWDEOHIRUPDVV SURGXFWLRQ+\GURELRORJLD 9RQVKDN $ HG D Spirulinu platensis (Arthrospira): 3K\VLRORJ\ &HOO%LRORJ\ DQG %LRWHFKQRORJ\ 7D\ORU )UDQFLV /WG/RQGRQ8. 9RQVKDN $ E 6SLPOLQD JURZWK SK\VLRORJ\ DQG ELRFKHPLVWU\ ,Q 9RQVKDN $ HG Spirulina platensis (Arthrospira): 3K\VLRORJ\ &HOO%LRORJ\ DQG %LRWHFKQRORJ\ SS 7D\ORU )UDQFLV/WG/RQGRQ8. 9RQVKDN$ $EHOLRYLFK$ %RXVVLED 6 $UDG 6 DQG5LFKPRQG $ 3URGXFWLRQ RI Spirulina ELRPDVV HIIHFWV RI HQYLURQPHQWDO IDFWRUV DQG SRSXODWLRQ GHQVLW\ %LRPDVV 9RQVKDN $ %RXVVLED6 $EHOLRYLFK $ DQG 5LFKPRQG $ 3URGXFWLRQ RI Spirulina ELRPDVV PDLQWHQDQFH RI SXUH FXOWXUH RXWGRRUV %LRWHFKQRO%LRHQJ 9RQVKDN $ &KDQDZRQJVH / %XQQDQJ % DQG 7DQWLFKDURHQ 0 D /LJKWDFFOLPDWLRQDQGSKRWRLQKLELWLRQLQWKUHH Spirulina platensis F\DQREDFWHULD LVRODWHV-$SSO3K\FRO 9RQVKDN $ DQG *X\ 5 3KRWRDGDSWDWLRQ SKRWRLQKLELWLRQ DQG SURGXFWLYLW\ LQ RXWGRRU JURZQ Spirulina platensis VWUDLQV 3ODQW&HOO(QYLURQP 9RQVKDN $ *X\ 5 DQG *X\ 0 D 7KH UHVSRQVH RI WKH ILODPHQWRXV F\DQREDFWHULXP Spirulina platensis WR VDOW VWUHVV $UFK0LFURELRO 9RQVKDN $ *X\ 5 3RSODZVN\ 5 DQG 2KDG E 3KRWRLQKLELWLRQ DQG LWV UHFRYHU\ LQ WZR GLIIHUHQW VWUDLQV RI Spirulina. 3ODQW&HOO3K\VLRO 9RQVKDN $ .DQFKDUDNVD 1 %XQQDQJ % DQG 7DQWLFKDURHQ 0 E 5ROH RI OLJKW DQG SKRWRV\QWKHVLV RQ WKH DFFOLPDWLRQ SURFHVV RI WKH F\DQREDFWHULXP Spimlina platensis WR VDOLQLW\ VWUHVV-$SSO3K\FRO 9RQVKDN$DQG5LFKPRQG$ 3KRWRV\QWKHWLFDQGUHVSLUDWRU\ DFWLYLW\ LQ Anacystis nidulans DGDSWHG WR RVPRWLF VWUHVV 3ODQW 3K\VLRO 9RQVKDN$DQG5LFKPRQG $ 0DVVSURGXFWLRQ RISpirulina DQRYHUYLHZ%LRPDVV 9RURQLFKLQ 11 %LRORJLH GHV EDVVLQV PLQHUDO GHV VWHSSHV .RXORQGLQH ([SHGLWLRQ GH , $FDGHPLH GHV 6FLHQFHV G 8566 , /HQLQJUDG 7UDY 6RYLHW SRXU VHUY O pWXGH UHVVRXU QDWXU6pU6LEHULH :D\QH /* %UHQQHU '- &ROZHOO 55 *ULPRQW 3$' .DQGOHU .ULFKHYVN\ 0, 0RRUH /+ 0RRUH :(& 0XUUD\ 5*( 6WDFNHEUDQGW( 6WDUU03 7USHU+* 5HSRUWRIWKH ad hoc FRPPLWWHH RQ UHFRQFLOLDWLRQ RI DSSURDFKHV WR EDFWHULDO V\VWHPDWLFV ,QW-6\VWHP%DFWHULRO :LWWURFN DQG 1RUGVWHGW $OJDH DTXDH GXOFLD H[LFFDWD IDVFLFXOH;,9Q,Q'HVFULSWLRQHVV\VWHPDWLFHGLVSRVLWDHS =DUURXN & &RQWULEXWLRQ j , pWXGH GXQH &\DQRSK\FWH ,QIOXHQFH GH GLYHUV IDFWHXUV SK\VLTXHV HW FKLPLTXHV VXU OD FURLVVDQFHHW ODSKRWRV\QWKHVH GH Spirulina maxima 6HWFKHOO HW *DUGQHU *HLWOHU3K'7KHVLV8QLYHUVLW\RI3DULV)UDQFH
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hitton and M . Potts (eds), The Ecology of Cyanobacteria, . © KIuwer Academic Publishers . Printed in the Netherlands
'*$GDPV
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
,9$ +RUPRJRQLD IRUPDWLRQ DQG KRUPRJRQLD WKHPVHOYHV DUH UHVSRQVLYH WR SODQW H[WUDFHOOXODU VLJQDOV 6HFWLRQ ,9% DQG LW PD\ EH IRU WKHVH UHDVRQV WKDW SODQW F\DQRELRQWV DUH DOPRVW DOZD\V PHPEHUVRIWKHJHQXVNostoc 6HFWLRQ,,, ,Q PRVW RIWKH V\PELRVHV GHVFULEHG LQ WKLV UHYLHZ WKH V\PELRQWV DUH UHFRJQLVLEOH DV F\DQREDFWHULD DQG DUH FDSDEOH RILQGHSHQGHQW JURZWK DOWKRXJK LQ VRPH FDVHV WKH\ KDYH GHILHG ODERUDWRU\ FXOWLYDWLRQ +RZHYHU WKHUH DUH WZR H[FHSWLRQV WKH SURFKORURSK\WHV DQGWKH F\DQHOOHV 7KHIRUPHU DUH D JURXS RI R[\JHQLF SKRWRV\QWKHWLF SURNDU\RWHV WKDW FORVHO\ UHVHPEOH F\DQREDFWHULD EXW GLIIHU LQ WKH SRVVHVVLRQ RIERWK FKORURSK\OOV a DQG b DQG WKH ODFN RISK\FRELOLQ SLJPHQWV 6HFWLRQ ,,( $ GHVFULSWLRQ RI RQH SURFKORURSK\WH Prochloron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
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV F\DQREDFWHULDOV\PELRVHVPRVWUHVHDUFKKDVIRFXVVHG RQ WKH H[SHULPHQWDOO\ DPHQDEOH SODQW DVVRFLDWLRQV DQGWKH OLWHUDWXUH UHIOHFWV WKLV +RZHYHU WKLVUHYLHZ ZLOODOVRDGGUHVVUHFHQWDGYDQFHVLQNQRZOHGJHRIWKH PDQ\ RWKHU F\DQREDFWHULDO DVVRFLDWLRQV DQG WKHLU HQYLURQPHQWDOVLJQLILFDQFH
,, 7KH6\PELRVHVDQG7KHLU (QYLURQPHQWDO ,PSDFW
GHSHQGHQW RQ WKH VSHFLILF FRPELQDWLRQ RI SODQW FXOWLYDU DQG F\DQREDFWHULDO VWUDLQ EXW FORVH DVVRFLDWLRQ RI WKH WZR LV QRW HVVHQWLDO 7KH QLWURJHQDVH DFWLYLW\ RIVRPH RIWKH F\DQREDFWHULD FDQ EHVXVWDLQHGLQWKHGDUNZKHQWKH\DUHDVVRFLDWHGZLWK WKH URRWV RI ZKHDW SODQWV SUHVXPDEO\ VXSSRUWHG E\ FDUERQFRPSRXQGVSUHVHQWLQURRWH[XGDWHV *DQWDUHW DOE
$ 3ODQWV /RRVH$VVRFLDWLRQV /RRVH DVVRFLDWLRQV EHWZHHQ SODQWV DQG F\DQREDFWHULD DUHOLNHO\WREHZLGHVSUHDG )RUH[DPSOHLWKDV ORQJ EHHQ NQRZQ WKDW WKH SUHVHQFH RI ERWK IUHHOLYLQJ LQFOXGLQJ HSLSK\WLF DQG V\PELRWLF (AzollaDVVRFLDWHG6HFWLRQ,,$ F\DQREDFWHULDLQULFHILHOGV LPSURYHV ERWK JURZWK DQG FURS \LHOG RIWKH ULFH VHH &KDSWHU (SLSK\WLF JURZWK RI Gloeotrichia pisum RQ VXEPHUJHG URRWV DQG VWHPV RIGHHSZDWHU ULFH KDV DOVR EHHQ REVHUYHG :KLWWRQ HW DO 3ODWH F G DQG LV DVVRFLDWHG ZLWK QLWURJHQIL[LQJ DFWLYLW\ 5RWKHU HW DO 6LPLODUO\ FRORQLHV RI WKH KHWHURF\VWRXV JHQHUD Nostoc, Gloeotrichia, Anabaena, Calothrix DQG Cylindrospermum KDYH EHHQ IRXQG JURZLQJ HSLSK\WLFDOO\ RQ WKH ORZHU HSLGHUPLV DQG WKH UHSURGXFWLYH SRFNHWV RIGXFNZHHG (Lemna) OHDYHV DQG DUH SUHVXPDEO\ UHVSRQVLEOH IRU WKHQLWURJHQIL[LQJDFWLYLW\DVVRFLDWHG ZLWKGXFNZHHG EORRPV'XRQJDQG7LHGMH $OWKRXJKNQRZOHGJHRIWKHHIIHFWVRIF\DQREDFWHULD RQ FURS SODQWV RWKHU WKDQ ULFH LV YHU\ OLPLWHG WKH LPSRUWDQFH RI ZKHDW DV D WHPSHUDWH IRRG FURS KDV SURPSWHGLQYHVWLJDWLRQVLQWKLVDUHDDQGKDVSURYLGHG HYLGHQFH WKDW F\DQREDFWHULD DVVRFLDWHG ZLWK ZKHDW URRWV FDQ VWLPXODWH SODQW JURZWK )RU H[DPSOH DQ DPPRQLDH[FUHWLQJ PXWDQW RI Anabaena variabilis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
Fig. 1. 6FDQQLQJHOHFWURQPLFURJUDSKVRIZKHDWURRWV $ 7KLFN OD\HU RI F\DQREDFWHULD RQ WKH URRW VXUIDFH % 3DFNDJHV RI F\DQREDFWHULDO ILODPHQWV WLJKWO\ DVVRFLDWHG ZLWK WKH URRW VXUIDFH DIWHU UHPRYDO RI ORRVHO\ DVVRFLDWHG F\DQREDFWHULD E\ ZDVKLQJ %DUV P 5HSURGXFHG ZLWK SHUPLVVLRQ IURP *DQWDU HW DO 1HZ3K\WRO &DPEULGJH8QLYHUVLW\3UHVV
' * $GDPV
%U\RSK\WHV 0RVVHV +RUQZRUWVDQG /LYHUZRUWV $PRQJ EU\RSK\WHV WKH +HSDWLFDH OLYHUZRUWV $QWKRFHURWDH KRUQZRUWV DQG 0XVFL PRVVHV DOO VKRZ H[DPSOHV RI HSLSK\WLF RU HQGRSK\WLF DVVRFLDWLRQV ZLWK F\DQREDFWHULD SULPDULO\ Nostoc. $VVRFLDWLRQV ZLWK PRVVHV DUH PRVWO\ HSLSK\WLF 0HHNV WKH H[FHSWLRQV EHLQJ VSHFLHV RI Sphagnum LQZKLFKWKHNostoc RFFXSLHVK\DOLQHFHOOV WKH IXQFWLRQ RI ZKLFK LV XQNQRZQ Hapalosiphon VRPHWLPHV DOVR JURZV FORVHO\ DVVRFLDWHG ZLWK Sphagnum, EXWFULWLFDO VWXGLHV RQKRZIDUWKLVFDQEH HQGRSK\WLF DSSHDU WR EH ODFNLQJ ,W KDV EHHQ VXJJHVWHG WKDW EXIIHULQJ ZLWKLQ K\DOLQH FHOOV KHOSV Nostoc WR JURZ DW S+ YDOXHV 0HHNV 3HWHUV +RZHYHU VSHFLHV RI WKH 1IL[LQJ JHQHUD Hapalosiphon DQG Tolypothrix DUH VRPHWLPHV IUHTXHQW LQ SRROV DW S+ YDOXHV DV ORZ DV %$ :KLWWRQSHUVFRPP 7KHVHHQGRSK\WLFDVVRFLDWLRQVDUHODUJHO\OLPLWHGWR SODQWVJURZLQJDERYHWKHZDWHUOLQHDQGKDYHQLWURJHQ IL[DWLRQ UDWHV EHORZ WKRVH RI HSLSK\WLF DVVRFLDWLRQV ZKLFK DUH XVXDOO\ FRQILQHG WR VXEPHUJHG SODQWV 7KHUHLV VRPHHYLGHQFHIRU WUDQVIHUWR WKH Sphagnum RI QLWURJHQ IL[HG E\ WKH F\DQREDFWHULD DQG IRU WKH HOHYDWLRQ RI QLWURJHQ IL[DWLRQ UDWHV RI HSLSK\WLFDOO\ DVVRFLDWHG Nostoc VSS DERYH WKH VDPH IUHHOLYLQJ VWUDLQV0HHNV 7KH QLWURJHQIL[LQJHSLSK\WLF DQG HQGRSK\WLF F\DQREDFWHULDO DVVRFLDWLRQV ZLWK PRVVHV VXFK DV Sphagnum DQG Drepanocladus PD\ FRQWULEXWH WR WKH QLWURJHQ HFRQRP\ RIERJ DQG PLUH KDELWDWV SDUWLFXODUO\ LQ VXEDUFWLF UHJLRQV 0HHNV (SLSK\WLF DVVRFLDWLRQV DOVR RFFXU ZLWK JUDVVODQG DQG IRUHVW PRVVHV VXFK DV Ceratodon DQG Funaria, DOWKRXJK WKH UDWHV RI QLWURJHQ IL[DWLRQ DUH PXFKORZHUWKDQWKRVHRISphagnum DVVRFLDWLRQV &\DQREDFWHULDO DVVRFLDWLRQV ZLWK OLYHUZRUWV DUH YHU\UDUHZLWKRQO\IRXURIRYHUJHQHUDNQRZQWR EH LQYROYHG WZR IRUPLQJ HSLSK\WLF DQG WZR HQGRSK\WLF DVVRFLDWLRQV )RXU RI WKH VL[ KRUQZRUW JHQHUD IRUP HQGRSK\WLF DVVRFLDWLRQV 7KH HSLSK\WLF DVVRFLDWLRQV DUH SRRUO\ XQGHUVWRRG DOWKRXJK WKH\ PD\EHPRUHFRPPRQWKDQRQFHWKRXJKW'DOWRQDQG &KDWILHOG %UDVHOOHWDO 7KHHQGRSK\WLF DVVRFLDWLRQV KDYH EHHQ VWXGLHG LQ PXFK PRUH GHWDLO EHFDXVH RI WKHLU DGDSWDELOLW\ WR ODERUDWRU\ H[SHULPHQWDWLRQ +RUQZRUWV VXFK DVAnthoceros DQG Phaeoceros, DQG OLYHUZRUWV VXFK Blasia, FDQ EH JURZQ D[HQLFDOO\ LQ VKDNHQ OLTXLG FXOWXUH ZLWK RU ZLWKRXW WKHLU V\PELRWLF F\DQREDFWHULDO SDUWQHUV (QGHUOLQ DQG 0HHNV 0HHNV .LPXUD
DQG1DNDQR 0HHNV 7KHV\PELRQWIUHH SODQWV FDQ EH UHDGLO\ UHLQIHFWHG ZLWK D[HQLF F\DQREDFWHULD 3ODWH H PDNLQJ WKLV DQ H[FHOOHQW PRGHO V\VWHP IRU WKH VWXG\ RI F\DQREDFWHULDSODQW V\PELRVLV ,Q WKHLU QDWXUDO KDELWDW WKHVH OLYHUZRUWV DQG KRUQZRUWV JURZ DV D SURVWUDWH JDPHWRSK\WH WKDOOXV DIHZFHQWLPHWHUV LQ OHQJWK EHLQJ DWWDFKHG WR WKH VXEVWUDWXP E\ UKL]RLGV 6\PELRWLF FRORQLHV DUH YLVLEOH DV GDUN VSRWV DSSUR[LPDWHO\ PP LQ GLDPHWHUZLWKLQWKHSODQWWLVVXHG'6HFWLRQ9
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ucalyptus IRUHVWVRI$XVWUDOLDWKHF\FDGV FDQ PDNH VLJQLILFDQW FRQWULEXWLRQV WR WKH ORFDO QLWURJHQHFRQRP\/LQGEODGDQG%HUJPDQ
$QJLRVSHUPV *XQQHUD 2I DOO WKH DQJLRVSHUPV IORZHULQJ SODQWV RQO\ WKH JHQXV Gunnera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unnera LVWKHRQO\RQHRIWKH
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Fig. 2. &\DQREDFWHULDO V\PELRVLV ZLWK WKH F\FDG Zamia skinneri. $ &RUDOORLG URRW FROOHFWHG IURP FP EHORZ WKH VXUIDFH % 7UDQVYHUVHVHFWLRQRIWKHURRWVKRZLQJWKHGDUNF\DQREDFWHULDO ]RQH & 6FDQQLQJHOHFWURQPLFURJUDSKRIWKHF\DQREDFWHULDO ]RQH 0RVW RI WKH F\DQREDFWHULD DQG PXFLODJH KDYH EHHQ UHPRYHG E\ ZDVKLQJ WR UHYHDO WKH HORQJDWHG Zamia FHOOV WUDYHUVLQJ WKH ]RQH $ IHZ F\DQREDFWHULDO FHOOV FDQ VWLOO EH VHHQ DUURZV ' 6FDQQLQJ HOHFWURQ PLFURJUDSK RI WKH F\DQREDFWHULDO ]RQH VKRZLQJ F\DQREDFWHULDO ILODPHQWV ZLWK KHWHURF\VWV D IHZ RIZKLFK DUH LQGLFDWHG E\ DUURZKHDGV %DUV PP $ PP % P & DQG P ' 5HSURGXFHGZLWKSHUPLVVLRQIURP/LQGEODGHWDO1HZ3K\WRO © &DPEULGJH8QLYHUVLW\3UHVV
SODQWF\DQREDFWHULD V\PELRVHV LQ ZKLFK WKH F\DQRELRQW (Nostoc VS LV IRXQG LQWUDFHOOXODUO\ ZLWKLQ VSHFLILF PXFXVVHFUHWLQJ JODQGV RQ WKH SODQW VWHPDW WKH EDVH RIHDFK SHWLROH )LJ 6HFWLRQ 9 (YHQ WKHODUJHVW Gunnera SODQWVFRORQLVHGE\Nostoc DUHDEOHWRIL[VXIILFLHQWQLWURJHQIRUWKHLURZQQHHGV DQG WKLV EHFRPHV DYDLODEOH WR WKH ORFDO QLWURJHQ HFRQRP\ %RQQHWW 2VERUQH HW DO Gunnera LVRQO\ DQJLRVSHUPIRUPLQJ V\PELRVHVZLWK
F\DQREDFWHULD DQG WKH F\DQRELRQWV RFFXU LQWUDFHOOXODUO\ 6LQFH DOO PDMRU FURS SODQWV DUH DQJLRVSHUPV DQ LPSURYHG XQGHUVWDQGLQJ RI WKH Nostoc-Gunnera V\VWHP PD\ LPSURYH WKH FKDQFHV RI GHYHORSLQJ QRYHO QLWURJHQIL[LQJ V\PELRVHV ZLWK HFRQRPLFDOO\LPSRUWDQWSODQWV
'*$GDPV
3WHULGRSK\WHV$]ROOD :LWKLQ WKH 3WHULGRSK\WD RQO\ Azolla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nabaena azollae, DOWKRXJK WKHLU WD[RQRPLF DVVLJQPHQW LV FRQWURYHUVLDO 6HFWLRQ ,,,$ Azolla FDQ EH IUHHG RILWV F\DQRELRQWV EXW QR ORQJHU IL[HV QLWURJHQ DQG EHFRPHV UHOLDQW RQ WKH SUHVHQFH RI FRPELQHGQLWURJHQ/LQDQG:DWDQDEH 5HFRUGV IURP WK FHQWXU\ 9LHWQDP DQG WK FHQWXU\ &KLQD GHVFULEH WKH XVH RIAzolla DV D JUHHQ PDQXUH LQ ULFH FXOWLYDWLRQ DQG LW LV VWLOO LQ XVH WKURXJKRXW WKH ZRUOG &KDSWHU 7KH IHUQ FDQ EH JURZQRQLWV RZQDQGLQFRUSRUDWHGLQWRWKH VRLOSULRU WR WUDQVSODQWLQJWKHULFH RUDV DQLQWHUFURS DORQJVLGH WKHULFH 1LHU]ZLFNL%DXHU ,QFRUSRUDWLRQ RI WKH Azolla, XVXDOO\ E\PHFKDQLFDOWXUQLQJ RIWKH VRLO RU E\ LQGXFHG FKHPLFDO RU QDWXUDO GHFRPSRVLWLRQ UHVXOWVLQWKHUHOHDVHRIFRPELQHGQLWURJHQXVXDOO\DV DPPRQLD 1LHU]ZLFNL%DXHU 7KH XVH RI Azolla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zolla LV XVHG DV DQ DQLPDO IRGGHU DV D ZHHG VXSSUHVVRU ZKHQ LQ GXDO FXOWLYDWLRQ ZLWK ULFH DQG KDV EHHQ VXJJHVWHG DV D PHDQV RILPSURYLQJ ZDWHU TXDOLW\ E\ WKH UHPRYDO RI H[FHVV TXDQWLWLHV RI QLWUDWH DQG SKRVSKRUXV 1LHU]ZLFNL%DXHU :DWDQDEH DQG /LX 7KH ZDWHU IHUQ FDQ DOVR EH XVHG DV IRRG IRU ILVK UHDUHGDORQJVLGHULFHLQIORRGHGULFHILHOGVDQGLWPD\ KHOS UHGXFH ORVV RIQLWURJHQ IURP WKH ILHOGV WKURXJK
DPPRQLD YRODWLOL]DWLRQ :DWDQDEH DQG /LX Azolla KDV DOVR EHHQ XVHG DV D UHSODFHPHQW IRU LQRUJDQLF QLWURJHQ IRU WKH JURZWK RI WRPDWR SODQWV 0LOLFLDDQG)DYLOOL
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ostoc, DOWKRXJK VSHFLHV RI Calothrix, Scytonema DQG Fischerella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
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV
Fig. 3. 7KHZDWHUIHUQAzolla. $ Azolla VSRURSK\WHYLHZHG IURPDERYHVKRZLQJWKHDSLFDOPHULVWHPDP DQGWKHGRUVDO G DQG YHQWUDO Y OHDIOREHV% ' 6FDQQLQJHOHFWURQPLFURJUDSKVRIOHDIFDYLWLHV RIAzolla VSS % 'HQVHO\SDFNHGF\DQREDFWHULDOILODPHQWVZLWKLQD OHDIFDYLW\ & $ILOLFXORLGHV VKRZLQJ WKH GLVWULEXWLRQ RI F\DQREDFWHULDO ILODPHQWV FRQWDLQLQJ KHWHURF\VWV WKH ODUJHU FHOOV DURXQG WKH SHULSKHU\RIWKHOHDIFDYLW\ $QAzolla KDLUFHOODUURZ LVDOVRYLVLEOH ' &\DQREDFWHULDO ILODPHQWV DVVRFLDWHGZLWKDSODQWKDLUFHOOKF %DUV PP$ P% DQG& DQG P' 5HSURGXFHG ZLWKSHUPLVVLRQ IURP % DQG ' 6KL HW DO 3ODQWD 6SULQJHU9HUODJ *PE+ &R .* & 5HSULQWHG IURP )(06 0LFURELRO /HWW 5RELQV HW DO ZLWK NLQG SHUPLVVLRQ IURP(OVHYLHU6FLHQFH 1/ 6DUD%XUJHUKDUWVWUDDW .9 $PVWHUGDP 7KH 1HWKHUODQGV $ SKRWRJUDSK FRXUWHV\ RI6 1LHU]ZLFNL%DXHU
QLWURJHQ IURP WKDOOL RU E\ WKH GHFD\ RI ROGHU WKDOOL WKDW IDOO WR WKH JURXQG $KPDGMLDQ &\DQROLFKHQVFDQDOVREHLPSRUWDQWSURYLGHUVRIIL[HG QLWURJHQ LQ WKH VXEDUFWLF DQG DUFWLFDQWDUFWLF UHJLRQV ZKHUH WKH\ IL[ QLWURJHQ DW D VXUIDFH WHPSHUDWXUH DV ORZDV& 3HWHUVHWDO DQGSKRWRV\QWKHVLVH DW & /DQJH HW DO )RU H[DPSOH WKH QLWURJHQIL[LQJ ELRWD RI WKH VXEDQWDUFWLF 0DF4XDULH ,VODQG LV GRPLQDWHG E\ F\DQREDFWHULD JURZLQJ ERWK HSLSK\WLFDOO\ DQG V\PELRWLFDOO\ ZLWK SODQWV VXFK DV
OLYHUZRUWV DQG PRVVHV DQG OLFKHQV VXFK DV Peltigera VS/LQH 0XFKRIWKHFDUERQIL[HGE\WKHSKRWRELRQWPD\EH XVHGLQWKHIRUPRISRO\ROV QRWDVDQXWULHQWEXWDV D SK\VLRORJLFDO EXIIHU KHOSLQJ WKH OLFKHQ WR VXUYLYH HQYLURQPHQWDO VWUHVVHV )DUUDU $KPDGMLDQ WKH PDMRU RQH EHLQJ WKH GDLO\ F\FOH RIGU\LQJ DQG ZHWWLQJ H[SHULHQFHG E\ PDQ\ OLFKHQV $KPDGMLDQ ,QGHHG OLFKHQV VKRZ UDSLG UHFRYHU\ RI QLWURJHQ IL[DWLRQ DIWHU SHULRGV RI
'*$GDPV
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cytonema, WKDW IDOO VKRUW RIWKHGHILQLWLRQRIDOLFKHQ JLYHQDERYH$KPDGMLDQ Geosiphon pyriforme, DQ DVVRFLDWLRQ RI Nostoc sphaericum ZLWK D SK\FRP\FHWH LV XQLTXH EHFDXVH WKH F\DQREDFWHULXP OLYHV HQGRV\PELRWLFDOO\ ZLWKLQ YHVLFOHV LQVLGH WKH IXQJDO K\SKDH 6HFWLRQV ,,%DQG9
*HRVLSKRQS\ULIRUPH ,QFRPPRQZLWKWKHOLFKHQV Geosiphom pyriforme LV D V\PELRWLF DVVRFLDWLRQ EHWZHHQ D IXQJXV DQG D F\DQREDFWHULXP 0ROOHQKDXHU .OXJH HW DO 0ROOHQKDXHU DQG .OXJH 6FKOHU HW DO 0ROOHQKDXHU HW DO +RZHYHU LQPDQ\ RWKHUUHVSHFWV WKHVH WZR IRUPV RI V\PELRVLV DUH YHU\ GLIIHUHQW 7KH IXQJXV LV D SK\FRP\FHWH WKRXJKW WR EH UHODWHG WR Glomus 6FKOHUHWDO *HKULJHWDO ZKHUHDV WKH F\DQREDFWHULXP LV Nostoc, WKRXJKW WR EH N. punctiforme RU N. sphaericum .OXJH HW DO %LOJHUHWDO DOWKRXJKRWKHUNostoc VSSFDQEH LQFRUSRUDWHGE\WKHIXQJXV.OXJHHWDO 7KH F\DQRELRQWV OLYH LQWUDFHOOXODUO\ ZLWKLQ VSHFLDOLVHG EODGGHUV SURGXFHG E\ WKH IXQJDO K\SKDH 3ODWH I DQG)LJ6HFWLRQ9
Geosiphon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eosiphon EODGGHUV JHUPLQDWH DQG FRQWLQXH WR OLYH LQ WKH VRLO 0ROOHQKDXHU ,QWKHILHOGGeosiphon LVIRXQG ZLWK WKH OLYHUZRUW Blasia DQG WKH KRUQZRUW Anthoceros, DQG WKHUH LV HYLGHQFH WKDW WKH Nostoc F\DQRELRQWV RIGeosiphon FDQ LQIHFW WKH EU\RSK\WHV DQG YLFH YHUVD 0ROOHQKDXHU .OXJH HW DO
& 'LDWRPV 7KH PDULQH SHQQDWH GLDWRP Rhizosolenia LV DEXQGDQW LQWKH1RUWK3DFLILF DQGIUHTXHQWO\IRUPVDV\PELRWLF DVVRFLDWLRQ ZLWK WKH ILODPHQWRXV KHWHURF\VWRXV F\DQREDFWHULXP Richelia intracelluluris, ZKLFK IL[HV ERWK &2 DQG 1 IRU WKH KRVW 5DL D (DFK F\DQREDFWHULDOILODPHQWRIZKLFKWKHUHFDQEHVHYHUDO LQ HDFK GLDWRP FHOO XVXDOO\ FRQWDLQV D VLQJOH EDVDO KHWHURF\VW DOWKRXJK RFFDVLRQDOO\ RQH LV SUHVHQW DW HDFK HQG 7KH DSH[ RI Rhizosolenia clevei FRQWDLQV F\WRSODVP RQO\ LQ WKH UHJLRQ RFFXSLHG E\ WKH F\DQREDFWHULXP DOWKRXJK WKH F\DQRELRQW LV DOZD\V H[WHUQDO WR WKH KRVW F\WRSODVP DQG LWV FHOO ZDOO UHPDLQV LQWDFW -DQVRQ HW DO E 7KH OHYHO RI JOXWDPLQH V\QWKHWDVH SURWHLQ LV YHU\ ORZ LQ WKH F\DQRELRQW DQG WKLV PD\ UHVXOW LQ UHOHDVH RI IL[HG QLWURJHQWRWKHKRVW-DQVRQHWDO E +RZHYHU WKHSUHVHQFHRIRichelia LV QRWHVVHQWLDOWRWKHJURZWK DQG VXUYLYDO RIRhizosoleniu, EHFDXVH WKH GLDWRPFDQ EHIRXQGZLWKRXWWKHF\DQRELRQW9LOODUHDO
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Fig. 4. Geosiphon pyriforme. $ 0DWXUHEODGGHU% VKRZLQJDGMDFHQWIXQJDOK\SKDH+\ % *URZWKRQSODQNWRQJDX]HVKRZLQJWKH FKDUDFWHULVWLF EODGGHUV FRQWDLQLQJ WKH F\DQREDFWHULXP Nostoc. & (DUO\ VWDJH LQ WKH LQFRUSRUDWLRQ RINostoc E\ WKH IXQJXV $IXQJDO K\SKD +\ KDV WUDQVIRUPHG LQWR D GXPEEHOO VKDSHG VWUXFWXUH LQ ZKLFK D EOXQW SURMHFWLRQ RI IXQJDO F\WRSODVP VXUURXQGV WKH F\DQREDFWHULXP&\ ' $ODWHUVWDJHRILQFRUSRUDWLRQLQZKLFKWKHIXQJDOZDOOKDV VRIWHQHGDQG DGGLWLRQDOF\WRSODVPLFEXOJHV VXUURXQG WKHF\DQREDFWHULXP %DUV PP$ PP% DQG P&DQG' 5HSURGXFHGZLWKSHUPLVVLRQIURP $ %LOJHUHWDO-3K\FRO 3K\FRORJLFDO6RFLHW\RI$PHULFD% .OXJHHWDO%RW$FWD *HRUJ7KLHPH9HUODJ& DQG ' 0ROOHQKDXHUHWDO3URWRSODVPD 6SULQJHU9HUODJ:LHQ
Richelia intracellularis LV DOVR IRXQG LQ GLDWRPV RI WKHJHQXVHemiaulus, ZKLFKZDVRQFHWKRXJKWWREHD UHODWLYHO\ XQFRPPRQ DQG XQVWDEOH V\PELRVLV 5DL D +RZHYHULWKDVUHFHQWO\EHHQVKRZQWKDW RIWKUHHHemiaulus VSS LQWKH%DKDPD,VODQGV DQG WKH &DULEEHDQ 6HD FRQWDLQ Richelia
intracellularis, WKLV V\PELRVLV EHLQJ XS WR WLPHV DV FRPPRQ KHUH DV WKH Rhizosolenia-Richelia DVVRFLDWLRQ 9LOODUHDO 7KH Hemiaulus V\PELRVHVKDYHEHHQIRXQGDVIDUUDQJLQJ DV1LQ WKH $WODQWLF DQG 6 LQ WKH 3DFLILF (SLIOXRUHVFHQW PLFURVFRS\LVUHTXLUHGIRUUHOLDEOHYLVXDOLVDWLRQRIWKH
'*$GDPV
F\DQRELRQW ZKLFK LV XVXDOO\ QRW YLVLEOH LQ OLYLQJ PDWHULDO XVLQJ WUDQVPLWWHG OLJKW SRVVLEO\ H[SODLQLQJ WKHSUHYLRXVXQGHUHVWLPDWHV IRUWKHRFFXUUHQFHRIWKLV V\PELRVLV9LOODUHDO 7ZR GLDWRPV RI WKH (SLWKHPLDFHDH Rhopalodia gibba DQG Epithemia turgida, SRVVHVV XQLFHOOXODU F\DQREDFWHULDO V\PELRQWV LQ WKHLU F\WRSODVP 5DL D 'H
' 'LQRIODJHOODWHV &\DQREDFWHULDO V\PELRQWV NQRZQ DV SKDHRVRPHV 7D\ORU DUH IRXQG LQ D YDULHW\ RI WURSLFDO PDULQH GLQRIODJHOODWHV /XFDV *RUGRQ HW DO -DQVRQHWDO D ,QWKHVXEWURSLFDOZDWHUV RIWKH *XOIRI$TDED KHWHURWURSKLF GLQRIODJHOODWHV RI JHQHUD Ornithocercus, Histioneis DQG Citharistes KDYH EHHQ VKRZQ WR FRQWDLQ F\DQREDFWHULDO V\PELRQWV WKRXJKWWR EH PHPEHUV RIWKH XQLFHOOXODU JHQHUD Synechococcus DQG Synechocystis *RUGRQ HW DO 7KHF\DQREDFWHULDDUHORFDWHGEHWZHHQWKH XSSHUDQGORZHUOLVWVRIWKHKRUL]RQWDOJURRYHRIFHOOV RI Ornithocercus DQG Histioneis, DQG ZLWKLQ VSHFLDO FKDPEHUVLQVLGHWKHFHOOVRICitharistes )LJ 7KH F\DQREDFWHULD ZHUH WKRXJKW WR EHQHILW E\ WKH SURYLVLRQ RID VXLWDEOH PLFURDHURELF HQYLURQPHQW IRU WKHHIILFLHQWIXQFWLRQLQJRIQLWURJHQDVH*RUGRQHWDO +RZHYHU UHFHQW LPPXQRORFDOLVDWLRQ VWXGLHV ZLWK WKH SKDHRVRPHV RI Ornithocercus magnifcus DQG O. steinii, ZKLFK DUH F\DQREDFWHULD RI WKH XQLFHOOXODU JHQXV Synechococcus, IDLOHG WR GHWHFW QLWURJHQDVHHYHQ LQVDPSOHVFROOHFWHG DWQLJKW ZKHQ VRPH XQLFHOOXODU F\DQREDFWHULD DUH NQRZQ WR IL[ QLWURJHQ WR DYRLG WKH LQLKLELWRU\ HIIHFWV RI SKRWRV\QWKHWLFDOO\ JHQHUDWHG R[\JHQ GXULQJ WKH GD\ -DQVRQ HW DO D 7KH F\DQRELRQWV VHHP WR SURYLGH IL[HG FDUERQ UDWKHU WKDQ QLWURJHQ IRU WKHLU QRQSKRWRV\QWKHWLF KRVW DOWKRXJK WKH\ PD\ DOVR EH SKDJRWURSKLFDOO\FRQVXPHGZKHQWKH\VHQHVFH
( 3URFKORURSK\WHV 7KH SURFKORURSK\WHV DUH D JURXS RI R[\JHQLF SKRWRV\QWKHWLF SURNDU\RWHV WKDW UHVHPEOH F\DQREDFWHULD LQ PDQ\ ZD\V EXW GLIIHU LQ WKH SRVVHVVLRQ RIERWK FKORURSK\OOV a DQG b DQG WKH ODFN RI SK\FRELOLQ SLJPHQWV %XUJHU:LHUVPD DQG 0DWWKLMV 0XU DQG %XUJHU:LHUVPD %XOOHUMDKQDQG3RVW 3RVWDQG%XOOHUMDKQ 0DWWKLMVHWDO 2QO\WKUHHSURFKORURSK\WHVDUH NQRZQ DQG WKHVH KDYH EHHQ SODFHG LQ WKUHH GLIIHUHQW JHQHUD Prochloron didemni, Prochlorothrix hollandica DQG Prochlorococcus marina. 7KH SK\ORJHQHWLF SRVLWLRQ RIWKHVH SURNDU\RWHV DQG WKHLU UHODWLRQVKLS WR WKH HYROXWLRQ RI SODVWLGV LV VWLOO FRQWURYHUVLDO %XOOHUMDKQ DQG 3RVW 3RVW DQG %XOOHUMDKQ 0DWWKLMV HWDO %KDWWDFKDU\D DQG 0HGOLQ EXWUHFHQWHYLGHQFH LQGLFDWHV WKDW Prochloron LV FORVHO\ UHODWHG WR Prochlorothrix DQG WKH F\DQREDFWHULD ZKHUHDV Prochlorococcus LV PRUH FORVHO\ UHODWHG WR WKH \SXUSOH EDFWHULD 6KLPDGD HW DO 7KH ILUVW RI WKH SURFKORURSK\WHV WR EH GLVFRYHUHGZDVProchloron didemni ZKLFKLVIRXQGLQ D V\PELRWLF UHODWLRQVKLS ZLWK PDULQH FRORQLDO DVFLGLDQV FKLHIO\ GLGHPQLGV LQ VXE WURSLFDO FRDVWDO ZDWHUV 7KHRWKHUWZRDUHIUHHOLYLQJVWUDLQVWKDWZLOO QRW EH GLVFXVVHG KHUH EXW KDYH EHHQ H[WHQVLYHO\ UHYLHZHG LQ UHFHQW \HDUV %XUJHU:LHUVPD DQG 0DWWKLMV 0XU DQG %XUJHU:LHUVPD %XOOHUMDKQDQG3RVW 3RVWDQG%XOOHUMDKQ 0DWWKLMVHWDO Prochloron LVXQLFHOOXODUZLWKFRFFRLGFHOOVWR P LQ GLDPHWHU DQG LV IRXQG DV D V\PELRQW LQ DVFLGLDQV RIWKH 'LGHPQLGDH ZKHUH LW XVXDOO\ RFFXUV H[WUDFHOOXODUO\ HLWKHU RQ WKH VXUIDFH RU ZLWKLQ WKH FORDFDO FDYLW\ RI WKH KRVW /HZLQ /HZLQ DQG &KHQJ *ULIILWKV DQG 7KLQK 6ZLIW DQG 5REHUWVRQ +RZHYHU Prochloron KDV EHHQ IRXQG LQ WKH DVFLGLDQ Lissoclinum punctatum ERWK H[WUDFHOOXODUO\ DQG LQWUDFHOOXODUO\ LQ WKH WXQLF FHOOV +LURVH HW DO 7KH LQWUDFHOOXODU Prochloron DUH PRUSKRORJLFDOO\ VLPLODU WR WKRVH IRXQG H[WUDFHOOXODUO\ DQG VKRZ QR VLJQV RI UHMHFWLRQ RU GHJHQHUDWLRQ LPSO\LQJ D VWDEOH V\PELRVLV &HOOV RI Prochloron FRQWDLQ FDUER[\VRPHV DQG DUH ERXQGHG E\DF\WRSODVPLFPHPEUDQHSHULSODVPLF VSDFH RXWHU PHPEUDQH DQG RXWHU VKHDWK 6ZLIW DQG /HVHU %XOOHUMDKQ DQG 3RVW 3RVW DQG %XOOHUMDKQ DQG WKRVH FHOOV IURP VRPH GLGHPQLGV FRQWDLQ RQHRUPRUHFKDUDFWHULVWLFYDFXROHVWKDWRFFXS\PXFK RIWKHFHOOYROXPH6ZLIW
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Fig. 5. &\DQREDFWHULDOV\PELRVLVZLWKGLQRIODJHOODWHV $ 6FDQQLQJHOHFWURQPLFURJUDSKRIOrnithocercus magnificus GDXJKWHUFHOOVKRUWO\ DIWHU GLYLVLRQ VKRZLQJ F\DQREDFWHULD LQ WKH FLQJXOXP DUURZ % Histioneis highleyi; & Citharistes apsteinii; ' Omirhocercus quadratus; ( Histioneis elongata. $UURZV LQ % ' LQGLFDWH WKH ORFDWLRQ RI F\DQREDFWHULD %DUV P 5HSURGXFHG ZLWK SHUPLVVLRQIURP/XFDV2SKHOLD 2SKHOLD3XEOLFDWLRQV
%LRFKHPLFDO DQG SK\VLRORJLFDO VWXGLHV ZLWK Prochloron :KDWOH\ DQG $OEHUWH KDYH EHHQ KDPSHUHG E\ WKH LQDELOLW\ WR JURZ LW LQ FXOWXUH +RZHYHU LWLV NQRZQWR IL[&2 IRULWVKRVW6HFWLRQ 9,' DQG WKHUH KDV EHHQ RQH UHSRUW RI DFHW\OHQH UHGXFWLRQ LQGLFDWLQJ QLWURJHQ IL[DWLRQ LQ RQH DVVRFLDWLRQ 3DHUO DOWKRXJK WKH 1IL[LQJ DFWLYLW\ RI VRPH 3URFKORURQDVFLGLDQ DVVRFLDWLRQV LQ WKH 6H\FKHOOHV KDV EHHQ VKRZQ QRW WR EH DVVRFLDWHG ZLWK Prochloron LWVHOI2GLQWVRY &KORURSK\OO IOXRUHVFHQFH TXHQFKLQJ DQDO\VLV KDV UHFHQWO\ EHHQ XVHG WR DVVHVV WKH SKRWRV\QWKHWLF SHUIRUPDQFH RI Prochloron ZLWKLQ WKH KRVW Lissoclinum patella 6FKUHLEHU HW DO 7KH V\PELRQW GLVSOD\HG H[FHSWLRQDOO\ KLJK TXDQWXP \LHOGV XQGHU ERWK OLJKW OLPLWLQJDQGOLJKWVDWXUDWHGFRQGLWLRQV 7KH UHVXOWV RI ROLJRQXFOHRWLGH FDWDORJXLQJ 6WDFNHEUDQGW HW DO DQG '1$'1$ UHDVVRFLDWLRQVWXGLHV6WDPHWDO +ROWRQHWDO KDYH LPSOLHG WKDW Prochloron LVRODWHG IURP GLIIHUHQW KRVWV DW RQH VLWH RU HYHQ KRVWV DW ZLGHO\ GLIIHUHQW ORFDWLRQV DSSHDU WR EH FORVHO\ UHODWHG RU SRVVLEO\ WKH VDPH VSHFLHV +RZHYHU DQ DQDO\VLV RI WKH PRUSKRORJLFDO FKDUDFWHULVWLFV RI ERWK V\PELRQW DQGKRVWOHG&R[ WRGLYLGHProchloron LVRODWHV LQWR WKUHHJURXSV 7KHVH UHVXOWV KDYH EHHQ SDUWLDOO\ VXSSRUWHG E\ FRPSDULVRQ RIQXFOHRWLGH VHTXHQFHV RI WKH '1$GHSHQGHQW 51$ SRO\PHUDVH (rpoC) JHQHV FRQILUPLQJ WKDWDWOHDVWWZR FORVHO\UHODWHG JURXSV RI Prochloron H[LVW LQ WKH VDPH DUHD RI 3DODX :HVW &DUROLQH ,VODQG DOWKRXJK VRPH PRUSKRORJLFDOO\ GLVWLQFWLVRODWHVDUHYHU\FORVHO\UHODWHGLPSO\LQJWKDW GLIIHUHQW SK\VLRORJLFDO VWDWHV RI Prochloron LQ GLIIHUHQW KRVWV PD\ DIIHFW WKHLU PRUSKRORJLFDO DSSHDUDQFH3DOHQLNDQG6ZLIW
) &\DQHOOHV &\DQHOOHV DUHRQH RIDUDQJH RISODVWLGV IRXQG LQ WKH FHOOV RI SKRWRV\QWKHWLF HXNDU\RWHV DQG ZHUH RQFH WKRXJKW WR EH F\DQREDFWHULDO HQGRV\PELRQWV RI WKHLU HXNDU\RWLFKRVWV3DVFKHU 7KH\SRVVHVVPDQ\ VLPLODULWLHV ZLWK F\DQREDFWHULD LQFOXGLQJ D SHSWLGRJO\FDQ ZDOO FKORURSK\OO a, SK\FRELOLSURWHLQV DQG FDUER[\VRPHOLNH VWUXFWXUHV 7UHQFK :DVPDQQ HW DO /RIIHOKDUGW DQG %RKQHUW DE3IDQ]DJOHWDO DE \HWWKHLUJHQRPHV DUH RQO\ WR NES VLPLODU LQ VL]H WR WKRVH RI FKORURSODVWVEXWRQO\DERXW RIWKRVHRIIUHHOLYLQJ F\DQREDFWHULD 7KH SK\ORJHQHWLF UHODWLRQVKLS EHWZHHQ F\DQREDFWHULD DQG SODVWLGV KDV UHFHLYHG FRQVLGHUDEOH DWWHQWLRQ LQ UHFHQW \HDUV DQG UHPDLQV
'*$GDPV FRQWURYHUVLDO DOWKRXJK WKH HPHUJLQJ YLHZ VHHPV WR EH WKDW DOO SODVWLGV LQFOXGLQJ F\DQHOOHV DURVH E\ DOPRVW VLPXOWDQHRXV GLYHUJHQFHV IURP D VLQJOH FRPPRQ DQFHVWRU LWVHOI WKH UHVXOW RI DQ HQGRV\PELRVLV EHWZHHQ D SKDJRWURSKLF KRVW DQG D F\DQREDFWHULXP 'RXJODV /RIIHOKDUGW DQG %RKQHUW D E +HOPFKHQ HW DO %KDWWDFKDU\D HW DO &\DQHOOHV KDYH EHHQ IRXQG LQ WKH FHOOV RI YHU\ GLIIHUHQW HXNDU\RWHV WKDW VKRZQHLWKHUV\VWHPDWLF QRUHYROXWLRQDU\ UHODWHGQHVV DOWKRXJK WKH WD[RQRPLF SRVLWLRQ RI WKHVH HXNDU\RWHV LV RIWHQ FRQIXVHG VHH :DVPDQQ HW DO IRU D GLVFXVVLRQ RIWKLV 'HWHUPLQLQJ WKH DQFHVWU\ RI WKH VHFRQGDU\ SODVWLGV SUHVHQW LQ PDQ\ SKRWRV\QWKHWLF HXNDU\RWHV LV FRPSOH[ EHFDXVH PDQ\ DUH OLNHO\ WR KDYHDFTXLUHGWKHVHE\LQMHVWLRQ RIDQRWKHUHXNDU\RWH DOUHDG\ FRQWDLQLQJ D SODVWLG 'HOZLFKH DQG 3DOPHU 7KH REOLJDWHO\ SKRWRDXWRWURSKLF DOJD Cyanophora paradoxa FDQ EH JURZQ HDVLO\ DQG LV FRQVHTXHQWO\ ZHOO VWXGLHG :DVPDQQHW DO /RIIHOKDUGWDQG %RKQHUW DE WKHFRPSOHWHQXFOHRWLGHVHTXHQFH RI WKH F\DQHOOH JHQRPH KDYLQJ EHHQ SXEOLVKHG 6WLUHZDOWHW DO 7KH FHQWUH RI Cyanophora F\DQHOOHV FRQWDLQV D FKDUDFWHULVWLF SURWHLQ LQFOXVLRQ ERG\ UHVHPEOLQJ WKH F\DQREDFWHULDO FDUER[\VRPH %ODQN DQG FRQWDLQLQJ PRVW RI WKH F\DQHOOH ULEXORVH ELVSKRVSKDWH FDUER[\ODVH 0DQJHQH\ DQG *LEEV ZKHUHDV SKRVSKRULEXORNLQDVH LV IRXQG LQVROXEOHIRUPLQWKHVWURPD0DQJHQH\HWDO 7KLVUHVHPEOHV WKH VLWXDWLRQ LQF\DQREDFWHULD \HWWKH F\DQHOOH LQFOXVLRQ ERGLHV ODFN WKH SURWHLQDFHRXV PHPEUDQH FKDUDFWHULVWLF RI WKH F\DQREDFWHULDO FDUER[\VRPH DQG LW LV XQOLNHO\ WKDW WKH LQFOXVLRQ ERGLHV DUH IXQFWLRQDO FDUER[\VRPHV /RIIHOKDUGW DQG %RKQHUWD &\DQHOOHV DUH XQLTXH DPRQJ SODVWLGV LQ EHLQJ VXUURXQGHGE\DSHSWLGRJO\FDQZDOORIWKHVDPH W\SH DV F\DQREDFWHULD /RIIHOKDUGW DQG %RKQHUW D E 3IDQ]DJO HW DO D E SURYLGLQJ DQ H[SODQDWLRQ IRU WKH OHWKDO HIIHFW RI ODFWDP DQWLELRWLFV RQ WKH REOLJDWHO\ DXWRWURSKLF RUJDQLVPV SRVVHVVLQJ F\DQHOOHV %HUHQJXHU HW DO 7KH JHQHVIRUWKHHQ]\PHVRISHSWLGRJO\FDQV\QWKHVLVDQG PHWDEROLVP KDYH QRW EHHQ LGHQWLILHG ZLWKLQ WKH FRPSOHWH VHTXHQFH RI WKH Cyanophora paradoxa JHQRPH DQG DUH SUHVXPDEO\ HQFRGHG E\ WKH QXFOHXV 6WLUHZDOWHWDO 7KHSHSWLGRJO\FDQLV ORFDWHG EHWZHHQ WKH F\DQHOOH PHPEUDQH DQG DQ RXWHU HQYHORSHPHPEUDQH/RIIHOKDUGWDQG%RKQHUW D E 7KHRULJLQRIWKHODWWHULV XQNQRZQEXWPD\EH D
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV UHPQDQW RIWKH RXWHU PHPEUDQH RIWKH F\DQREDFWHULDO DQFHVWRURIWKHF\DQHOOH
* $QLPDOV6SRQJHV$VFLGLDQV (FKLXURLG :RUPVDQG0LGJH/DUYDH $OPRVW JHQHUD RI PDULQH VSRQJH DUH NQRZQ WR FRQWDLQ V\PELRWLF F\DQREDFWHULD IRXQG ERWK LQWUD DQG LQWHUFHOOXODUO\ DQG GLVWULEXWHG HLWKHU WKURXJKRXW WKH VSRQJH RUMXVW LQ WKH VXSHUILFLDO WLVVXHV )LJ 5DL D:LONLQVRQ 7KH F\DQRELRQWV DUH XVXDOO\ PHPEHUV RI WKH XQLFHOOXODU JHQXV Aphanocapsa, RU WKH ILODPHQWRXV JHQXV Phormidium 5DL D DOWKRXJK WKH F\DQRELRQWRIWKH WURSLFDO PDULQH VSRQJH Dysidea herbacea KDV EHHQ LGHQWLILHG DV Oscillatoria spongelia 8QVRQDQG)DXONQHU +LQGHHWDO 8QVRQHWDO $OO RIWKHVH JHQHUD ODFN KHWHURF\VWV DOWKRXJK WKH F\DQRELRQWV RI DWOHDVWWZR VSRQJHVKDYHEHHQ VKRZQWRIL[ QLWURJHQ 5DL D &\DQRELRQWV DUH SKRWRV\QWKHWLFDOO\ DFWLYH RQO\ EHLQJ IRXQG LQ VSRQJHV LQ WKH SKRWLF ]RQHDQGWUDQVIHUXSWR RIWKHFDUERQWKH\IL[WR WKH KRVW SUREDEO\ LQ WKH IRUP RI JO\FHURO 5DL D 7KH GHJUHH RIPHWDEROLF UHOLDQFH RIVRPH VSRQJHV RQ WKH SUHVHQFH RIWKH F\DQRELRQWV LV LOOXVWUDWHG E\ Chondrilla nucula ZKLFK FDQQRW FRORQLVH GDUN VLWHV DQG LIWUDQVIHUUHG WR WKHGDUN ORRVHV LWV F\DQRELRQWV DQGGLHV$ULOORHWDO 7KHEHQHILWVWRWKHKRVW KDYH EHHQ GHPRQVWUDWHG LQ -DPDLFDQ FRUDO UHHI VSRQJHV LQ ZKLFK WKRVH FRQWDLQLQJ F\DQREDFWHULDO V\PELRQWV JURZ PRUH UDSLGO\ :LONLQVRQ DQG &KHVKLUH 7KHF\DQRELRQWVDUHDOVRWKRXJKWWR KHOS SURWHFW WKH VSRQJH IURP GDPDJH E\ KLJK OLJKW LQWHQVLW\ 5DL D 2Q UDUH RFFDVLRQV WKH UHODWLRQVKLS EHWZHHQ JURZWK RI KRVW DQG F\DQRELRQW FDQ EHFRPH XQEDODQFHG $Q H[DPSOH RFFXUV LQ WKH VSRQJH Geodia papyraceae IURP D PDQJURYH LVODQG RII %HOL]H WKDW FRQWDLQV LQWHUFHOOXODU FRFFRLG F\DQREDFWHULDHTXDO LQYROXPHWR DOPRVWKDOILWV RZQ FHOOXODU WLVVXH 5XHW]OHU 8QGHU IDYRXUDEOH JURZWKFRQGLWLRQVWKHF\DQRELRQWFHOOVPXOWLSO\PRUH UDSLGO\ WKDQ WKH VSRQJH FDQ HOLPLQDWH WKHP FDXVLQJ KRVW WLVVXH GDPDJH VRFDOOHG PDQJURYH VSRQJH GLVHDVH 6SRQJHV VXFK DV Theonella swinhoei DQG Siphonochalina tabernacula )LJ WKDW FRQWDLQ QLWURJHQIL[LQJ F\DQRELRQWV PRVWOLNHO\ EHQHILWIURP WKHSURYLVLRQ RIFRPELQHG QLWURJHQ 5DL D DQG WKLV SUHVXPDEO\ EHQHILWV WKH ORFDO QLWURJHQ HFRQRP\ ,QGHHG WKH VSRQJH Chondrilla nucula, ZKLFK LV XELTXLWRXVLQFRUDOUHHIVRII3XHUWR5LFRFRQWDLQV1
IL[LQJ F\DQREDFWHULDO V\PELRQWV DQG UHOHDVHV VXIILFLHQW QLWUDWH WR DFFRXQW IRU RI WKH QLWURJHQ UHTXLUHG WR VXVWDLQ UHHI SURGXFWLYLW\ &RUUHGRUHWDO 0DQ\ VSRQJHV DOVR SURGXFH ELRDFWLYH QDWXUDO SURGXFWV DW OHDVW VRPH RI ZKLFK PD\ KHOS SURWHFW DJDLQVW SRWHQWLDO SUHGDWRUV DQG EDFWHULDO LQYDVLRQ )DXONQHU HW DO Oscillatoria spongelia, WKH F\DQREDFWHULDO V\PELRQW RI WKH PDULQH VSRQJH Dysidea herbacea, SURGXFHV D JURXSRISRO\FKORULQDWHG 8QVRQDQG)DXONQHU )DXONQHU HW DO +DQGD\DQL HW DO DQG SRO\EURPLQDWHG 8QVRQ HW DO VHFRQGDU\ PHWDEROLWHV 2QH RI WKH ODWWHU LV LQKLELWRU\ WR WKH JURZWK RI D UDQJH RI *UDPSRVLWLYH DQG *UDP QHJDWLYH EDFWHULD DQG GHWHUV IHHGLQJ E\ JHQHUDOLVW ILVKHV )DXONQHU HW DO $ QRYHO XOWUDYLROHW DEVRUELQJ P\FRVSRULQHOLNH DPLQR DFLG LV DOVR SURGXFHG E\ WKH F\DQRELRQW RI Dysidea herbacea %DQGDUDQD\DNH HW DO %DQGDUDQD\DNH 7KH SURGXFWLRQ RI VXFK VHFRQGDU\ PHWDEROLWHV ZDV RQFHWKRXJKWWREHOLPLWHGWR WKH VSRQJHWLVVXH RUWR DVVRFLDWHGKHWHURWURSKLFEDFWHULD8QVRQHWDO )DXONQHUHWDO ,QGHHG WKH VSRQJH Theonella swinhoei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ermocarpa, Xenococcus DQG Synechococcus. 5HSUHVHQWDWLYHVRIWKHDGGLWLRQDO F\DQREDFWHULDO JHQHUD Oscillatoria, Spirulina DQG Anabaena KDYHEHHQIRXQGLQWKHLQWHUWXQLFFDYLW\RI WKH GRXEOHWXQLF DVFLGLDQV Pyura cancellata DQG P. carnea /DPEHUW HW DO &\DQREDFWHULD KDYH EHHQ UHSRUWHG LQ WKH VXEHSLGHUPDO FRQQHFWLYH WLVVXH RI WZR VSHFLHV RI HFKLXURLG ZRUP Ikedosoma gogoshimense DQG Bonellia fuliginosa, EXW OLWWOH LV NQRZQRIWKHVHDVVRFLDWLRQV5DL D $Q DSSDUHQWO\ PXWXDOLVWLF DVVRFLDWLRQ RFFXUV EHWZHHQ ODUYDH RIWKH FKLURQRPLG PLGJH Cricotopus nostocicola OLYLQJ LQVLGH FRORQLHV RI Nostoc parmelioides LQ PRGHUDWHO\ IDVWIORZLQJ VWUHDPV 6KHDWK DQG &ROH 'RGGV HW DO 7KH ODUYDH HDW WKH Nostoc DQG DUH SURWHFWHG IURP SUHGDWLRQ ZKLOH Nostoc FRORQLHV FRQWDLQLQJ
'*$GDPV
Fig. 6. 6\PELRWLF F\DQREDFWHQDLQWKHVSRQJH Siphonochalina tabernacula. $ &\DQREDFWHULD&\ DQGEDFWHULD %D ZLWKLQVSRQJHFHOO YDFXROHV % 6HFWLRQWKURXJKVSRQJHYDFXROHVKRZLQJF\DQREDFWHULXPZLWKVSLUDOWK\ODNRLGVDQGDGMDFHQWVPDOOHUEDFWHULD %DUV P $ DQG , P% 5HSURGXFHGZLWKSHUPLVVLRQIURP:LONLQVRQDQG)D\1DWXUH 0DFPLOODQ0DJD]LQHV/LPLWHG
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
,,, 7KH6\PELRQWV $ &\DQREDFWHULD ,Q WKH SODQWF\DQREDFWHULD V\PELRVHV WKH F\DQRELRQW LV DOPRVW DOZD\V Nostoc 7DEOH 7KH DELOLW\ WR IRUP KRUPRJRQLD LV WKH NH\ WR VXFFHVV DV D V\PELRQW EHFDXVH KRUPRJRQLD DUH WKH LQIHFWLYH DJHQWV LQ PRVW LI QRW DOO RI WKHVH V\PELRVHV 7KH LPPRWLOH SDUHQW ILODPHQWV ZRXOG EH XQDEOH WR JDLQ HQWU\ WR WKH SODQW VWUXFWXUHV WKDW KRXVH WKH V\PELRWLF FRORQLHV ,W LV FOHDU KRZHYHU WKDW D ZLGH UDQJH RI Nostoc VSHFLHV DQG VWUDLQV DUH V\PELRWLFDOO\ FRPSHWHQW LQ D ZLGH
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Table I. ,GHQWLW\DQGORFDWLRQRIWKHV\PELRQWVLQSODQWF\DQREDFWHULDV\PELRVHV
3ODQW KRVW
3ODQWVWUXFWXUHLQIHFWHG´
&\DQRELRQW
&\DQRELRQW ORFDWLRQ
&\FDGV
&RUDOORLG URRWV )LJ
Nostoc Calothrix)b
,QWHUFHOOXODU
Gunnera
6WHP JODQGV )LJ
Nostoc
,QWUDFHOOXODUG
/LYHUZRUWV
$XULFOHV 3ODWH DH DQG )LJ
Nostoc
,QWHUFHOOXODU b
,QWHUFHOOXODU ,QWHUFHOOXODU
+RUQZRUWV
6OLPH FDYLWLHV
Nostoc Calothrix)
Azolla
'RUVDO OHDIFDYLWLHV )LJ
Anabaena (Nostoc " F
7KHILJXUHVLQWKHWH[WWKDWLOOXVWUDWHHDFKVWUXFWXUHDUHJLYHQLQSDUHQWKHVHV 7KHV\PELRQWVDUHNostoc VSSLQDOPRVWDOOFDVHVWKHUHKDYHEHHQUDUHUHSRUWVRICalothrix VSS DVV\PELRQWV6HFWLRQ,,,$ F 7KH Azolla V\PELRQW LV XVXDOO\ UHIHUUHG WR DV Anabaena VS EXW WKH DOWHUQDWLYH JHQHUD Nostoc DQG Trichormus KDYH EHHQ VXJJHVWHG 6HFWLRQ,,,$ G Gunnera LVWKHRQO\SODQWV\PELRVLVLQZKLFKWKHF\DQREDFWHULDDUHIRXQGLQWUDFHOOXODUO\VHH6HFWLRQ9
D
E
UDQJHRISODQWV/HL]HURYLWFKHWDO =LPPHUPDQ DQG&XOOH\ %HUJPDQHWDO E/RWWLHWDO 0HPEHUVRIJHQHUDRWKHUWKDQNostoc FDQDOVR IRUPV\PELRVHVZLWKSODQWV )RUH[DPSOH WKHF\FDG Encephalartos KDVEHHQUHSRUWHGWRFRQWDLQ Calothrix VSS DVF\DQRELRQWV *UREEHODDUHWDO DV KDV WKH KRUQZRUW Phaeoceros :HVW DQG $GDPV ,Q FRPPRQ ZLWK Nostoc, Calothrix LV KHWHURF\VWRXV DQGFDSDEOHRIKRUPRJRQLDIRUPDWLRQ 1RWZLWKVWDQGLQJ WKH SUHYLRXV H[DPSOH F\FDG V\PELRQWV DUH XVXDOO\ Nostoc VSS 3ODWH I DQG PRUSKRORJLFDOFRPSDULVRQV *UREEHODDUHW DO DQG UHVWULFWLRQ IUDJPHQW SRO\PRUSKLVP 5)/3 DQDO\VLV RIF\DQRELRQW '1$ /LQGEODG HW DO KDYH VKRZQ WKDWDYDULHW\ RIVWUDLQV FDQEH LQYROYHG DOWKRXJK LW VHHPV WKDW D VLQJOH Nostoc VWUDLQ LV SUHGRPLQDQWLQ DSDUWLFXODUURRW 6LPLODUFRQFOXVLRQV KDYHEHHQGUDZQ IURPFRPSDULVRQV RIOHFWLQELQGLQJ HOHFWURSKRUHWLF SURWHLQ SURILOHV DQG ]\PRJUDPV RI Nostoc VSS IURPF\FDGVDWDVLQJOHVLWH=LPPHUPDQ DQG 5RVHQ $V ZLWK Gunnera, DOWKRXJK VHYHUDO Nostoc VSS FDQ LQIHFWWKHSODQW =LPPHUPDQ DQG %HUJPDQ 6RGHUEDFN %HUJPDQ HW DO D LW LV OLNHO\ WKDW RQO\ RQH RU D IHZ DUH SUHVHQW LQ DQ LQGLYLGXDO SODQW %HUJPDQ HW DO E 7KH F\DQRELRQWV RI OLFKHQV VKRZ PXFK JUHDWHU GLYHUVLW\ ZLWK ERWK ILODPHQWRXV DQG XQLFHOOXODU JHQHUD UHSRUWHG +RZHYHU WKH SLFWXUH LV OHVV WKDQ FOHDU EHFDXVH PRVW LGHQWLILFDWLRQV RI F\DQRELRQWV KDYH UHOLHG RQ GLUHFW REVHUYDWLRQ UDWKHU WKDQ FXOWXULQJ DQG F\DQREDFWHULD RIWHQ GLVSOD\ YHU\ GLIIHUHQWPRUSKRORJLHVZKHQV\PELRWLFDOO\DVVRFLDWHG
UDWKHU WKDQ IUHHOLYLQJ 5DL E 6HFWLRQ 9,% 0HPEHUV RI DW OHDVW IRXU JHQHUD RI ILODPHQWRXV KHWHURF\VWRXV F\DQREDFWHULD DUH NQRZQ WR EH LQYROYHG Nostoc, Calothrix, Scytonema DQG Fischerella 5DL E 2I WKH XQLFHOOXODU F\DQREDFWHULD WKH PRVW FRPPRQ DUH Gloeocapsa, Gloeothece DQG Synechocystis, DOWKRXJK WKHLU H[DFW WD[RQRPLF VWDWXV LV RIWHQ XQFHUWDLQ 5DL E $ UHFHQW FRPSDUVLRQ RI3&5 DPSOLILHG W51$/HX 8$$ LQWURQV IURP WKH Nostoc V\PELRQWV RI WKH OLFKHQV Nephroma arcticum, Peltigera aphthosa, P. membranacea DQG P. canina UHYHDOHG WKDW D VLQJOH V\PELRQW ZDV SUHVHQW LQ DQ LQGLYLGXDO WKDOOXV 3DXOVUXG DQG /LQGEODG &ORVHO\ UHODWHG F\DQRELRQWV ZHUH DOVR SUHVHQW LQ WKH VDPH OLFKHQ FROOHFWHG IURP GLIIHUHQW ORFDWLRQV LQ 6ZHGHQ DQG LQ )LQODQG LPSO\LQJ DKLJK GHJUHH RIVSHFLILFLW\ LQ WKH FKRLFH RI Nostoc V\PELRQW 3DXOVUXG HW DO 7KH VDPH WHFKQLTXH KDV EHHQ XVHG WR VKRZ WKDW WKH VDPH Nostoc V\PELRQW RFFXUV LQ WKH ELSDUWLWH DQG WULSDUWLWHOREHV RIDSKRWRV\PELRGHPH RIP. aphthosa 3DXOVUXGHWDO ,Q WKH FDVH RI WKH XQLTXH IXQJDO DVVRFLDWLRQ Geosiphon pyriforme, RQO\ Nostoc VSS VHHP WR EH F\DQRELRQWV ,WLVLQWHUHVWLQJWKDWLQQDWXUHGeosiphon RFFXUV DW WKH VDPH VLWHV DV Blasia DQG Anthoceros, DQGWKHUHLV HYLGHQFHWKDWWKH Geosiphon F\DQRELRQWV FDQ LQIHFW WKH EU\RSK\WHV DQG YLFH YHUVD 0ROOHQKDXHU .OXJHHWDO 7KH F\DQRELRQW RI Azolla KDV EHHQ WUDGLWLRQDOO\ UHIHUUHG WR DV Anabaena azollae, DOWKRXJK WKLV WD[RQRPLF DVLJQPHQW KDV EHHQ FRQWURYHUVLDO DV WKH GLVFXVVLRQ EHORZ UHYHDOV Azolla LV XQXVXDO LQ
'*$GDPV
FDUU\LQJ D FRORQ\ RI WKH F\DQRELRQW ZLWKLQ WKH VSRURFDUS GXULQJ VH[XDO UHSURGXFWLRQ PDNLQJ UH LQIHFWLRQXQQHFHVVDU\ 3HWHUVHWDO 3HWHUV DQG 0HHNV 3OD]LQVNL %HUJPDQ HW DO E DQGUHPRYLQJWKH QHHGIRUWKHF\DQRELRQWWR SURGXFHKRUPRJRQLDIRULQIHFWLRQWRRFFXU 0HPEHUV RI D QRQKRUPRJRQLXPIRUPLQJ JHQXV VXFK DV Anabaena FRXOG WKHUHIRUH EH FRQVLGHUHG SRWHQWLDO V\PELRQWVLQWKLVV\VWHP +RZHYHU XQGLIIHUHQWLDWHG ILODPHQWV UHVHPEOLQJ KRUPRJRQLD DUH DVVRFLDWHG ZLWK WKH DSLFDO PHULVWHP RI HDFK Azolla VWHP SURYLGLQJ WKH LQRFXOXP IRU QHZ OHDIFDYLWLHV 3HWHUV DQG0HHNV 7KH Azolla F\DQRELRQW ZDV ILUVW GHVFULEHG DV Nostoc E\6WUDVEXUJHU ZKRODWHUDVVLJQHGLWWR WKH JHQXV Anabaena DQG UHQDPHG LW Anabaena azollae 6WUDVEXUJHU 7KLV FODVVLILFDWLRQ KDV RIWHQEHHQTXHVWLRQHGDQGWKHJHQXVNostoc KDVEHHQ SURSRVHG EDVHG RQ FRPSDULVRQV RI WKH DUUDQJHPHQW RIQLIJHQHV 0HHNV HW DO DQG WKH QXPEHUV RI VXSHUR[LGH GLVPXWDVH LVRHQ]\PHV &DQLQL HW DO E LQ V\PELRWLF DQG IUHHOLYLQJ Anabaena DQG Nostoc VWUDLQV 7KH UHVXOWV RI5)/3 DQDO\VLV RIWKH JHQRPLF '1$ RI F\DQRELRQWV IURP VHYHQ Azolla VSHFLHV DOVR LPSO\ WKDWWKH\ DUH PRUH FORVHO\ UHODWHG WR Nostoc WKDQ WRAnabaena 3OD]LQVNL HW DO 7KLV DVVHUWLRQ LV VXSSRUWHG E\ WKH UHVXOWV RI SRO\PHUDVH FKDLQ UHDFWLRQ 3&5 DPSOLILFDWLRQ RI Azolla F\DQRELRQW '1$ .LP HW DO +RZHYHU FRPSDULVRQV RI FHOOXODU IDWW\ DFLGV KDYH LPSOLHGWKDWWKHAzolla V\PELRQWV DUHHTXDOO\GLVWLQFW IURP ERWK Anabaena DQG Nostoc &DXGDOHV HW DO 7KH DOWHUQDWLYH QDPH Trichormus azollae KDV EHHQVXJJHVWHG *ULOOL&DLRODSHUV FRPP TXRWHG LQ %HUJPDQ HW DO DQG VXSSRUWHG E\ VXEVHTXHQW REVHUYDWLRQV RIWKH FKDUDFWHULVWLFV RIDNLQHWHV SUHVHQW LQ WKH F\DQRELRQW RIAzolla jiliculoides *ULOOL&DLROD HWDO 7KH WUXH LGHQWLW\ RI WKH Azolla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
,PPXQRORJLFDO $UDG HW DO DQG OHFWLQ KDHPDJJOXWLQDWLRQ VWXGLHV 0F&RZHQ HW DO =LPPHUPDQ HW DO 5)/3 DQDO\VHV )UDQFKH DQG &RKHQ%D]LUH 0HHNV HW DO 3OD]LQVNL HW DO *HEKDUGW DQG 1LHU]ZLFNL %DXHU DQG HQ]\PDWLF FRPSDULVRQV =LPPHUPDQ HW DO KDYH FRQILUPHG WKDW WKH UHDGLO\ FXOWXUDEOH F\DQRELRQWV RI Azolla DUH QRW WKH VDPH DV WKH IUHVKO\LVRODWHG V\PELRQW 3&5EDVHG WHFKQLTXHV KDYH EHHQ XVHG VXFFHVVIXOO\ WR GLIIHUHQWLDWHEHWZHHQAzolla V\PELRQWV IURP GLIIHUHQW SODQWVSHFLHV (VNHZHWDO .LP HWDO 7KH JHQHWLF GLYHUVLW\ RI Anabaena azollae VWUDLQV IURPGLIIHUHQWIHUQV KDV DOVR EHHQ DVVHVVHG E\ 5)/3 DQDO\VLVXVLQJQLIJHQHSUREHV 9DQ&RSSHQROOHHWDO D DQG WKH UHVXOWV VXJJHVW D WD[RQRP\ RI A. azollae SDUDOOHO ZLWK Azolla LWVHOI GHWHUPLQHG E\ '1$ DPSOLILFDWLRQ XVLQJ DUELWUDU\ SULPHUV 9DQ &RSSHQROOHHWDO E 6LPLODUO\ DQDO\VLV RIWKH IDWW\ DFLG FRPSRVLWLRQ RIF\DQRELRQWV IURP GLIIHUHQW $PHULFDQ Azolla VSHFLHV LPSOLHV WKDWWKH F\DQRELRQWV IRUP VHYHUDO FOXVWHUV WKDW FRUUHODWH ZHOO ZLWK WKH WD[RQRPLF JURXSLQJ RI WKH SODQWV LPSO\LQJ WKDW WKH F\DQRELRQWV KDYHFRHYROYHG ZLWK WKHSODQW &DXGDOHV HWDO SUHVXPDEO\DVDUHVXOWRIWKHSUHVHQFHRI Anabaena azollae WKURXJKRXWWKH OLIH F\FOH RIAzolla, DYRLGLQJWKHQHHGIRUFRQVWDQWUHLQIHFWLRQ
% %DFWHULD ,Q DGGLWLRQ WR WKH F\DQREDFWHULD WKH OHDI FDYLWLHV RI Azolla FRQWDLQ KHWHURWURSKLF EDFWHULD *ULOOL &DLRODHW DO 1LHU]ZLFNL%DXHU DQG $XOILQJHU 3OD]LQVNL HW DO E LQFOXGLQJ DEXQGDQW Arthrobacter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
&KDSWHU 6\PELRWLF,QWHUDFWLRQV PXFLODJH SURGXFHG E\ WKH F\DQRELRQWV )RUQL HW DO E $V ZLWK WKH F\DQRELRQW WKH EDFWHULD DSSHDU WR UHPDLQ DVVRFLDWHG ZLWK Azolla WKURXJKRXW LWV OLIH F\FOH&DUUDSLoR DQG 7DYDUHV %UDXQ+RZODQG DQG 1LHU]ZLFNL%DXHU &DUUDSLoR ,QGHHG EDFWHULD DUH IRXQG WRJHWKHU ZLWK F\DQREDFWHULDO DNLQHWHV EHQHDWK WKH LQGXVLXP RI Azolla PHJDVSRURFDUSV DQG WKH\ FRPPHQFH FHOO GLYLVLRQ DW WKH VDPH WLPH DV WKH DNLQHWHV JHUPLQDWH $XOILQJHUHWDO /LWWOH LV NQRZQ RI WKH LQYROYHPHQW RI EDFWHULD LQ F\DQREDFWHULDO V\PELRVHV DSDUW IURP Azolla. ,Q WKH FDVH RI Gunnera, DOWKRXJK EDFWHULD DUH SUHVHQW ZLWK WKH F\DQREDFWHULD LQ WKH HDUO\ VWDJHV RI JODQG IRUPDWLRQ WKH\GRQRWHQWHUWKHKRVWFHOOVDORQJZLWK WKHF\DQREDFWHULD%HUJPDQHWDO E
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scillutoria, WKH KRUPRJRQLD DUH IRUPHG E\ VLPSOH IUDJPHQWDWLRQ RIWKH SDUHQW WULFKRPH DQG DSDUWIURP
EHLQJ VKRUWHU LQ OHQJWK UHVHPEOH WKH RULJLQDO WULFKRPH LQ DOO UHVSHFWV LQFOXGLQJ EHLQJ PRWLOH RU LPPRWLOH ,W PD\ EH WKDW WKH FKDUDFWHULVWLF VPDOO FHOOHG KRUPRJRQLDIRUPHG E\UHGXFWLYH FHOO GLYLVLRQ H[HPSOLILHG E\ Nostoc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
% 6LJQDOOLQJEHWZHHQ 3RWHQWLDO3DUWQHUV %HFDXVH KRUPRJRQLD DUH WKH SULPDU\ LQIHFWLYH DJHQWV LQ PRVW LI QRW DOO SODQW V\PELRVHV LW LV QR VXUSULVH WKDW SODQW KRVWV SURGXFH H[WUDFHOOXODU VLJQDOV WKDW WULJJHU KRUPRJRQLD IRUPDWLRQ LQ VXLWDEOH F\DQREDFWHULD Anthoceros punctatus UHOHDVHV D ORZ PROHFXODU PDVV KHDWODELOH SURGXFW WKDW VWLPXODWHV KRUPRJRQLD IRUPDWLRQ LQ Nostoc VWUDLQV &DPSEHOO DQG 0HHNV 7KLV GRHV QRW RFFXU ZKHQ WKH KRUQZRUW LV FXOWXUHG LQ PHGLXP FRQWDLQLQJ H[FHVV 1+ LPSO\LQJ WKDWWKH FRPSRXQG LV SURGXFHG ZKHQ WKHSODQWLV QLWURJHQ VWDUYHG 7KH LPSRUWDQFH RIWKLV SODQW VLJQDO LV LOOXVWUDWHG E\ D WUDQVSRVRQ PXWDQW RI WKH V\PELRWLF VWUDLQ Nostoc ZKLFK VKRZV LQFUHDVHG VHQVLWLYLW\ WR WKH Anthoceros KRUPRJRQLD LQGXFLQJ IDFWRU DQG KDV D PXFK KLJKHU LQLWLDO IUHTXHQF\ RILQIHFWLRQ RIWKH KRUQZRUW WKDQ WKH ZLOG W\SH&RKHQHWDO 7KH DFLGLF PXFLODJH RI WKH Gunnera VWHP JODQG DOVR VHUYHV DV D KRUPRJRQLDLQGXFLQJ VLJQDO WKH DFWLYH FRPSRQHQW RIZKLFK VHHPV WR EH D KHDWODELOH SURWHLQ RI OHVV WKDQ N'D SUHVHQW RQO\ LQ WKH
'*$GDPV
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unnera V\PELRWLF WLVVXH DUH DOVR SRWHQWLDO VLJQDOV LQ WKLV V\VWHP ZLWK SRVVLEOH UROHV LQ DWWUDFWLRQ GHIHQFH RU WKH WUDQVFULSWLRQDO DFWLYDWLRQ RIF\DQREDFWHULDO JHQHV LQYROYHGLQV\PELRVLV%RQQHWW %HUJPDQHWDO D )ODYRQRLGV RI OHJXPHV LQGXFH V\PELRVLV VSHFLILF JHQH H[SUHVVLRQ LQ UKL]RELD YDQ 5KLMQ DQG 9DQGHUOH\GHQ DQG WKHUH LV HYLGHQFH RI D GHJUHH RI KRPRORJ\ EHWZHHQ VRPH RI WKHVH IODYRQRLGUHVSRQVLYH JHQHV DQG FHUWDLQ JHQH VHTXHQFHV RI WKH F\DQREDFWHULD LQIHFWLQJ Azolla 3OD]LQVNL HW DO DQG Gunnera 5DVPXVVHQ HW DO 6LJQDOVWKDWIXQFWLRQLQWKHRSSRVLWHGLUHFWLRQIURP F\DQREDFWHULXP WR Gunneru, KDYH UHFHQWO\ EHHQ SRVWXODWHG EDVHG RQ VHYHUDO OLQHV RI HYLGHQFH %HUJPDQ HW DO )LUVWO\ WKHUH LV VRPH KRPRORJ\ EHWZHHQ WKH UKL]RELD JHQHV nodEF, nodMN DQG WKH nod ER[ DQG JHQRPLF UHJLRQV RI F\DQREDFWHULDWKDW LQIHFW Gunneru 5DVPXVVHQ HW DO DQGAzolla 3OD]LQVNLHWDO ,QUKL]RELD WKHVH nod JHQHV FRGH IRU WKH ELRV\QWKHVLV RI OLSRROLJRVDFFKDULGHV WKDW VHUYH DV QRGXOHLQGXFLQJ IDFWRUVSURGXFHGLQUHVSRQVHWRSODQWIODYRQRLGV YDQ 5KLMQ DQG 9DQGHUOH\GHQ 6HFRQGO\ WDUJHW Gunnera FHOOV DUH WKRXJKW WR XQGHUJR HQKDQFHG PLWRWLF GLYLVLRQ LQ UHVSRQVH WR WKH SUHVHQFH RI V\PELRWLFF\DQREDFWHULDUHVXOWLQJLQWKHIRUPDWLRQRI D V\PELRWLF RUJDQ ZLWKLQ WKH VWHP JODQG EHDULQJ IXQFWLRQDO VLPLODULWLHV WR WKHURRW QRGXOHV RIOHJXPHV -RKDQVVRQ %HUJPDQHWDO 7KHJURZWK RIWKHILQJHUOLNHWUDQVIHUFHOOVLQWRV\PELRWLFFRORQLHV LQEU\RSK\WHVVXFKDVAnthoceros DQG Blasia 6HFWLRQ 9 PD\ DOVR UHVXOW IURP WKH SUHVHQFH RI VLJQDOV GHULYHGIURPWKHF\DQREDFWHULD )LQDOO\FHOOGLYLVLRQ GXULQJ HPEU\RJHQHVLV LQ FDUURW LV VWLPXODWHG E\ H[WUDFWV RIFHUWDLQ F\DQREDFWHULD :DNH HW DO $ SRVVLEOH VRXUFH RIWKH VLJQDOV WKDW LQGXFH VXFK FKDQJHV LQ WKH SODQW KRVW PD\ EH WKH DUDELQRJDODFWDQ SURWHLQV UHOHDVHG E\ PDQ\
F\DQREDFWHULD%HUJPDQHWDO VXFK$*3V DUH WKRXJKW WR KDYH LPSRUWDQW UROHV LQ SODQW JURZWK DQG GHYHORSPHQW3HQQHOO ,QWHUDFWLRQV EHWZHHQ SODQW DQG F\DQREDFWHULXP DUH QRW OLPLWHG WR WKH HQGRSK\WLF DVVRFLDWLRQV +\GURSRQLFDOO\ JURZQ ZKHDW URRWV DOVR UHOHDVH D KRUPRJRQLD LQGXFLQJ IDFWRU WKDW LV SURGXFHG DW D KLJKHU OHYHO LQ QLWURJHQIUHH PHGLXP *DQWDU HW DO 7KHLQLWLDOFRORQL]DWLRQRIWKHURRWVE\Nostoc 6% RFFXUV E\ PRWLOH KRUPRJRQLD HYHQ ZKHQ WKH LQLWLDO F\DQREDFWHULDO LQRFXOXP FRQWDLQV QR KRUPRJRQLD*DQWDUHWDO 7KLVLVEHFDXVHWKH KRUPRJRQLD LQGXFLQJ IDFWRU WULJJHUV KRUPRJRQLD IRUPDWLRQ WKHUHE\ RYHUFRPLQJ DQ DSSDUHQW DXWRLQKLELWRU RI KRUPRJRQLD GHYHORSPHQW SUHVHQW LQ WKH DVHULDWH SDFNDJHV RI ILODPHQWV WKDW DUH FKDUDFWHULVWLFRINostoc 6% )RU HIILFLHQW LQIHFWLRQ RI PRVW SODQWV WKH PHUH SUHVHQFH RI KRUPRJRQLD LV XQOLNHO\ WR EH VXIILFLHQW EHFDXVH RI WKH UHVWULFWHG DFFHVVLELOLW\ RI WKH SODQW VWUXFWXUHV LQYROYHG 7KHKRUPRJRQLDPXVWEHJXLGHG LQWR WKH SODQW E\ FKHPRDWWUDFWLRQ $OWKRXJK WKH PXFLODJH RI Gunnera VWHP JODQGV FDQ LQGXFH KRUPRJRQLDIRUPDWLRQLWGRHVQRWDSSHDUWRFRQWDLQD FKHPRDWWUDFWDQW5DVPXVVHQ HWDO +RZHYHU ZKHQ VWDUYHG RI FRPELQHG QLWURJHQ WKH OLYHUZRUW Blasia UHOHDVHV H[WUDFHOOXODU VLJQDOV WKDW QRW RQO\ WULJJHUKRUPRJRQLDIRUPDWLRQ %DELF EXWDOVR VHUYH DV YHU\ HIIHFWLYH FKHPRDWWUDFWDQWV .QLJKW DQG $GDPV *HUPLQDWLQJZKHDWVHHGVDOVRUHOHDVH KRUPRJRQLD FKHPRDWWUDFWDQWV .QLJKW DQG $GDPV
& 2WKHU,PSRUWDQW)DFWRUV /LWWOH LV NQRZQ DERXW WKH UHFRJQLWLRQ SURFHVVHV LQYROYHG LQ WKH LQIHFWLRQ RI KRVWV E\ F\DQREDFWHULD ,Q WKH OLFKHQ EU\RSK\WH DQG Azolla V\PELRVHV WKH KRVWSODQW LV NQRZQ WR SURGXFH OHFWLQV WKDWUHFRJQLVH VSHFLILF VXJDU UHVLGXHV RQ WKH F\DQREDFWHULDO FHOO VXUIDFH %UDXQ+RZODQG DQG 1LHU]ZLFNL%DXHU .DUGLVK HW DO %HUJPDQ HW DO E +RQHJJHU 7KH XVH RI IOXRUHVFHLQ LVRWKLRF\DQDWHFRQMXJDWHG OHFWLQV DQG FRQIRFDO ODVHU VFDQQLQJPLFURVFRS\KDVUHFHQWO\UHYHDOHGFKDQJHVLQ WKH H[WUDFHOOXODU VOLPH RI WKH Nostoc F\DQRELRQW RI Geosiphon pyriforme GXULQJ LWV OLIH F\FOH ZLWK D' PDQQRV\O RU 'JOXFRV\O UHVLGXHV RU JO\FRFRQMXJDWHV FRQWDLQLQJWKHP EHLQJFKDUDFWHULVWLF RI WKH SRVWKRUPRJRQLD VWDJH WKDW LV FDSDEOH RI IRUPLQJ V\PELRVLV ZLWK WKH IXQJXV 6FKOHU HW DO
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV 7KH IRUPDWLRQ RI ERWK HQGRSK\WLF DQG HSLSK\WLF DVVRFLDWLRQV UHTXLUHV WKH DWWDFKPHQW RIF\DQREDFWHULD WR WKH KRVW VXUIDFH DQG ERWK ILPEULDH SLOL DQG H[WUDFHOOXODUSRO\VDFFKDULGHV KDYHEHHQLPSOLFDWHG LQ WKH SURFHVV )LPEULDH KDYH EHHQ REVHUYHG LQ WKH Nostoc F\DQRELRQW RIPeltigera canina DQG PD\ SOD\ DUROHLQVSHFLILFLW\ 'LFNDQG 6WHZDUW 3ODQW DQG F\DQREDFWHULDO SRO\VDFFKDULGHV DUH LPSRUWDQW IRU WKH DWWDFKPHQW RI F\DQREDFWHULD WR SODQW FHOOV DQG LQHUWVXUIDFHV5RELQVHWDO 7KHLQYROYHPHQW RIILPEULDHDQGSRO\VDFFKDULGHVLQWKHFRORQL]DWLRQRI ZKHDWURRWVE\Nostoc 6%LQYROYHVWZRVWDJHVILUVW KRUPRJRQLD EHFRPH ZHDNO\ DWWDFKHG E\ PHDQV RI ILPEULDH VHFRQG WKH KRUPRJRQLD GHYHORS LQWR KHWHURF\VWRXV ILODPHQWV WKDW EHFRPH ILUPO\ DWWDFKHG E\WKHSURGXFWLRQRIDPXFLODJLQRXVVKHDWK*DQWDUHW DO 7KH FKDUDFWHULVWLFV RIWKH VKHDWK VHHP WR EH LPSRUWDQW IRU WKH DWWDFKPHQW WR ZKHDW URRWV EHFDXVH WKH LVRODWHG VKHDWK RI Nostoc 6% ZKLFK IRUPV DWLJKWDVVRFLDWLRQ ZLWKWKHURRWV LV FDSDEOH RI WLJKW DWWDFKPHQW WR WKH URRW VXUIDFH ZKHUHDV WKDW RI Anabaena & ZKLFK IRUPV RQO\ D ORRVH DVVRFLDWLRQ ZLWKWKHURRWV LVRQO\ORRVHO\DWWDFKHG*DQWDUHWDO D ,QWKHGunnera-Nostoc V\PELRVLVWKHLPSRUWDQFHRI ILPEULDH KDV EHHQTXHVWLRQHG EHFDXVHWKH\ DUH IRXQG LQERWKLQIHFWLYHDQGQRQLQIHFWLYH Nostoc VWUDLQV DUH PRVW DEXQGDQW LQ WZR QRQLQIHFWLQJ VWUDLQV DQG DSSHDU WR EH DEVHQW IURP VRPH LQIHFWLYH VWUDLQV -RKDQVVRQ DQG %HUJPDQ 1HYHUWKHOHVV ILPEULDH PD\ VHUYH D QRQVSHFLILF IXQFWLRQ LQ LQLWLDO DGKHVLRQ RI SRWHQWLDO V\PELRQWV WR WKH Gunnera VXUIDFH SULRU WR WKH LQYROYHPHQW RI IXUWKHU DV \HW XQNQRZQ UHFRJQLWLRQ VLJQDOV WKDW GHWHUPLQH WKH XOWLPDWH VXFFHVV RU IDLOXUH RI WKH LQIHFWLRQ SURFHVV 7KDW WKH SODQW KRVW SOD\V DQ LPSRUWDQW UROH LQ GHWHUPLQLQJ WKH IDWH RI D SRWHQWLDO V\PELRQW LV LOOXVWUDWHGE\WKHREVHUYDWLRQWKDWVRPHNostoc VWUDLQV WKDWVXFFHVVIXOO\LQIHFWERWKGunnera DQGEU\RSK\WHV EHFRPHLQWUDFHOOXODULQWKHIRUPHUEXWQRWLQWKHODWWHU -RKDQVVRQDQG%HUJPDQ
9 +RVW 6WUXFWXUHVDQG7KHLU,QIHFWLRQ 8QOLNH WKH EDFWHULDO V\PELRQWV LQ UKL]RELDO DQG DFWLQRUKL]DOV\PELRVHVF\DQREDFWHULDRFFXS\H[LVWLQJ SODQWVWUXFWXUHV 7KH\ DUH QRWOLPLWHG WR DUHDV RIWKH SODQWUHFHLYLQJOLJKWEXWFDQEHIRXQGJURZLQJSKRWR RU FKHPRKHWHURWURSKLFDOO\ LQ IRU H[DPSOH WKH FRUDOORLG URRWV RI F\FDGV ZKLFK UHFHLYH OLWWOH RU QR OLJKW
7KH F\DQREDFWHULDO V\PELRQWV RI WKH OLYHUZRUW Blasia RFFXS\ DXULFOHV ZKLFK DUH DOPRVW VSKHULFDO VWUXFWXUHV RQ WKH YHQWUDO VXUIDFH RIWKH WKDOOXV 3ODWH DHDQG)LJ ZKHUHDVLQWKHPXFKWKLFNHUWKDOOXV RI WKH KRUQZRUWV Anthoceros DQG Phaeoceros WKH F\DQREDFWHULD DUH IRXQG ZLWKLQ WKH WKDOOXV LQ VOLPH FDYLWLHV WKDW RSHQ WR WKH YHQWUDO VXUIDFH YLD QDUURZ VOLWOLNHSRUHV 3ODWH $ ,Q ERWK FDVHV WKH PDWXUH V\PELRWLF FRORQLHV DUH XS WR PP LQ GLDPHWHU 2[\JHQ PLFURHOHFWURGH PHDVXUHPHQWV KDYH VKRZQ WKDWLQAnthoceros WKHFDYLW\LVPLFURDHURELFDQG DVD UHVXOW FDQVXSSRUWQLWURJHQDVH DFWLYLW\ LQ DPXWDQWRI WKH V\PELRWLF Nostoc WKDW LV XQDEOH WR IL[ QLWURJHQ XQGHU DHURELF FRQGLWLRQV EHFDXVH RI D GHIHFWLYH KHWHURF\VW HQYHORSH &DPSEHOO DQG 0HHNV 7KH F\DQREDFWHULD HQWHU Blasia DXULFOHV DQG SUHVXPDEO\ KRUQZRUW VOLPH FDYLWLHV DV KRUPRJRQLD ZKHUHXSRQ WKH\ ORVH PRWLOLW\ DQG GLIIHUHQWLDWH KHWHURF\VWV.LPXUDDQG1DNDQR ,Q Azolla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zolla VWHP FDUULHV D FRORQ\ RI XQGLIIHUHQWLDWHG F\DQRELRQW ILODPHQWV UHVHPEOLQJ KRUPRJRQLD SURWHFWHG E\ WKH QHZO\ GHYHORSLQJ ELOREHG OHDYHV $V QHZ OHDYHV JURZ WKHLU GHYHORSLQJ OHDIFDYLWLHV EHFRPH LQIHFWHG IURPWKLVDSLFDOAnabaena azollae FRORQ\3HWHUVDQG 0HHNV %UDXQ+RZODQG DQG1LHU]ZLFNL%DXHU 3HUNLQV DQG 3HWHUV 3HWHUV DQG 3HUNLQV &RQWLQXLW\ RI WKH V\PELRVLV LV PDLQWDLQHG WKURXJK WKH VH[XDO F\FOH E\ WKH SDFNDJLQJ RI Anabaena azollae LQWR WKH GHYHORSLQJ PHJDVSRURFDUSVZKHUHWKH\GLIIHUHQWLDWHLQWRDNLQHWHV 3HWHUV DQG 0HHNV %UDXQ+RZODQG DQG 1LHU]ZLFNL%DXHU 3HWHUV $XOILQJHU HW DO 3HUNLQV DQG 3HWHUV 3HWHUV DQG
'*$GDPV
Fig. 7. &\DQREDFWHULDOLYHUZRUWV\PELRVHV $ ' 6FDQQLQJHOHFWURQPLFURJUDSKVRIBlasia pusilla. $ )ODVNVKDSHGJHPPDUHFHSWDFOH RQWKHYHQWUDOVXUIDFHRIWKHWKDOOXV ,QGLYLGXDOHOOLSVRLGDOJHPPDHDIDUP RIDVH[XDOSURSDJXOH FDQEHVHHQZKHUHWKHUHFHSWDFOHZDOOKDV EHHQGDPDJHGDWWKHEDVH % DQG& &URVVVHFWLRQVRIDXULFOHVFRQWDLQLQJF\DQREDFWHULDOFRORQLHV ' 6WHOODWHJHPPDDVHFRQGIRUP RI DVH[XDOSURSDJXOH RQWKHVXUIDFHRIWKHWKDOOXV LQSUR[LPLW\WRWZRDXULFOHV DUURZV ( /LJKWPLFURJUDSKRIIRXUBlasia pusilla DXULFOHV $XULFOH LV XQLQIHFWHG DQG KDYH UHFHQWO\ EHFRPH LQIHFWHG DQG FRQWDLQ VLQJOH F\DQREDFWHULDO ILODPHQWV DQG FRQWDLQV D GHYHORSLQJ V\PELRWLF FRORQ\ ) )LODPHQWRXV SURWUXVLRQV I H[WUXGHG IURP DQ LQIHFWHG DXULFOH VXUURXQGHG E\ F\DQREDFWHULDO FHOOV VKRZLQJWKHODFNRIWKHILODPHQWRXVKDELWWKDWFKDUDFWHULVHVWKHIUHHOLYLQJVWDWH %DUV P$ P%DQG& P' DQG P(DQG) 5HSURGXFHGZLWKSHUPLVVLRQIURP $ ' +RQQHJJHU)ORUD *XVWDY)LVFKHU9HUODJ) 5RGJHUV DQG6WHZDUW1HZ3K\WRO &DPEULGJH8QLYHUVLW\3UHVV( SKRWRJUDSKFRXUWHV\6%DELF
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unnera-Nostoc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ostoc, IXQJL DQG EDFWHULD 7KLV PXFLODJHFRQWDLQVDWOHDVWWKUHHVLJQDOOLQJFRPSRXQGV WKDW LQGXFH KRUPRJRQLD IRUPDWLRQ DQG LQIOXHQFH F\DQREDFWHULDO JHQH H[SUHVVLRQ DQG WKH JURZWK RI FRPSDWLEOH Nostoc VWUDLQV 6HFWLRQ ,9% 'XULQJ VWDJH WZR RIWKH LQIHFWLRQ SURFHVV F\DQREDFWHULD WKDW KDYH DFFXPXODWHG RQ WKH JODQG DUH LQGXFHG WR GLIIHUHQWLDWHKRUPRJRQLDDQGGXULQJVWDJHWKUHHWKHVH
PLJUDWH WRZDUGV WKH LQWHULRU RIWKH JODQG YLD QDUURZ LQWHUFHOOXODU FKDQQHOV IRUPHG E\ WKH GLVVROXWLRQ RI FHOO ZDOOV GXULQJ WKH HDUO\ VWDJHV RIJODQG IRUPDWLRQ -RKDQVVRQ DQG %HUJPDQ %HUJPDQ HW DO E ,QWKH LQWHULRURIWKH JODQGWKH Gunnera FHOO ZDOOV GLVVROYH DQG WKH F\DQREDFWHULD HQWHU WKH FHOOV ZLWKRXW SHQHWUDWLQJ WKH KRVW SODVPDOHPPD VWDJHV IRXUDQGILYH ,WLV QRWFOHDU ZKHWKHUWKH GLVVROXWLRQ RI KRVW FHOO ZDOOV LV FDXVHG E\ WKH SODQW WKH F\DQREDFWHULD RU WKH RWKHU EDFWHULD SUHVHQW EXW RQO\ WKHF\DQREDFWHULDHQWHUWKHKRVWFHOOV%HUJPDQHWDO E ,Q WKH ILQDO VWDJH WKH Nostoc KRUPRJRQLD GLIIHUHQWLDWH DKLJK IUHTXHQF\ RIKHWHURF\VWV 6HFWLRQ 9,% DQG EHJLQ IL[LQJ QLWURJHQ ZKLOH WKH JODQG FKDQQHOV GLVDSSHDU UHQGHULQJ IXUWKHU LQIHFWLRQ LPSRVVLEOH $OWKRXJK LQ DOO RI WKH DERYH H[DPSOHV WKH F\DQREDFWHULD LQYDGH H[LVWLQJ SODQW VWUXFWXUHV WKH SUHVHQFH RI WKH F\DQRELRQWV GRHV LQGXFH VRPH FKDQJHV LQ WKH SODQW WLVVXHV )RU H[DPSOH LQ WKH F\FDG Mucrozamia communis PRVW XQLQIHFWHG SUHFRUDOORLG URRWV VHQHVFH ZKHUHDV LQIHFWHG SUHFRUDOORLGV DUH VWLPXODWHG WR GLIIHUHQWLDWH LQWR FRUDOORLGV $KHUQ DQG 6WDII 7KH ILODPHQWRXV SURWUXVLRQV)LJ) WKDWJURZLQWRWKHF\DQREDFWHULDO FRORQ\ LQAnthoceros DUH VWLPXODWHG E\ WKH SUHVHQFH RI WKH F\DQRELRQW 5RJHUV DQG 6WHZDUW DQG WKHUH LV HYLGHQFH WKDW F\DQREDFWHULD VWLPXODWH FHOO GLYLVLRQ RI VSHFLILF Gunnera FHOOV UHVXOWLQJ LQ WKH IRUPDWLRQ RI D V\PELRWLF RUJDQ ZLWKLQ WKH VWHP JODQG %HUJPDQ HW DO (YHQ LQ D QRQKRVW SODQW VXFK DV WKH FDUURW (Daucus carota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eosiphon pyriforme LV XQXVXDO EHFDXVH WKH F\DQRELRQWLVLQWUDFHOOXODUHQWHULQJWKHIXQJDOK\SKDH E\ HQGRF\WRVLV 0ROOHQKDXHU .OXJH HW DO
'*$GDPV
Fig. 8. 7KHDQJLRVSHUPGunnera chilensis. $ 6FDQQLQJHOHFWURQPLFURJUDSKVKRZLQJDVWHPJODQGORFDWHGDWWKHEDVHRIWZROHDISHWLROHV 7KHJODQGLVMXVWEUHDNLQJWKURXJKWKHSODQWHSLGHUPLVH DQGWKHLQGLYLGXDOSDSLOODHS FDQEHVHHQ VHSDUDWHGE\FKDQQHOV OHDGLQJLQWRWKH VWHPWLVVXH % /RQJLWXGLQDO VHFWLRQ RIDVWHP JODQG 7KH FKDQQHOV DUURZV VHSDUDWLQJWKH SDSLOODH S DUH OLQHG E\ D OD\HU RIHOHFWURQ GHQVHFHOOV 5HSURGXFHGZLWKSHUPLVVLRQIURP%HUJPDQHWDO1HZ3K\WRO &DPEULGJH8QLYHUVLW\3UHVV %DUV P
0ROOHQKDXHU HW DO 2QO\ LPPRWLOH ³SULPRUGLD´ WKDW GHYHORS IURP PRWLOH KRUPRJRQLD DUH LQFRUSRUDWHG E\ WKH KRVW 0ROOHQKDXHU HW DO 6FKLLOMOHUHWDO :KHQ WKH WLSV RI WKH IXQJDOK\SKDHHQFRXQWHUVXLWDEOHNostoc ILODPHQWVWKH IXQJDO ZDOO VRIWHQV DQGWKH F\DQREDFWHULDO YHJHWDWLYH FHOOVEXWQRWDQ\H[LVWLQJKHWHURF\VWVDUHHQJXOIHGE\ WKH IXQJDO SODVPDOHPPD E\ D SURFHVV RI SKDJRF\WRVLV )LJ & ' 0ROOHQKDXHU HW DO 0ROOHQKDXHU DQG 0ROOHQKDXHU 1HZ KHWHURF\VWV GHYHORS IURP WKH HQJXOIHG YHJHWDWLYH FHOOV 2QFH LQIHFWHG WKH K\SKDH VZHOO WR IRUP SHDU VKDSHG PXOWLQXFOHDWH EODGGHUV XS WR PP LQ GLDPHWHU DQG PP LQ OHQJWK FRQWDLQLQJ WKH
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ostoc ILODPHQWV RFFXS\ WKH LQQHU VXUIDFH RI WKHEODGGHUV LQ DFXSVKDSHG FRPSDUWPHQWNQRZQ DV
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Fig. 9. 7KHOLFKHQLichina confinis. $ 9LHZRIWKHWKDOOXV % /RQJLWXGLQDO VHFWLRQ RISDUWRIWKHWKDOOXV VKRZLQJWKHF\DQRELRQW&\ DQGP\FRELRQW0\ %DUV PP$ DQG P % 5HSURGXFHGZLWKSHUPLVVLRQ IURP-DQVRQ HWDO 1HZ3K\WRO &DPEULGJH8QLYHUVLW\3UHVV
WKH V\PELRVRPH ERUGHUHG E\ D KRVW PHPEUDQH 6FKOHU HW DO 7KH UHJLRQ EHWZHHQ WKH V\PELRVRPH PHPEUDQH DQG WKH Nostoc FHOO ZDOO NQRZQ DV WKH V\PELRVRPH VSDFH LV ILOOHG ZLWK DQ DPRUSKRXV PDWHULDO DV DUH RWKHU DUHDV RI WKH FRPSDUWPHQW QRW RFFXSLHG E\ WKH F\DQRELRQW 7KH SUHFLVH QDWXUH RI WKH DPRUSKRXV PDWHULDO LV QRW NQRZQ EXW LW DSSHDUV WR FRQWDLQ FRPSRQHQWV FKDUDFWHULVWLF RI WKH IXQJDO FHOO ZDOO LQFOXGLQJ SRO\VDFFKDULGHV DQG ILEULOODU FKLWLQ DQG LV SUREDEO\ SURGXFHG E\ WKH IXQJDO V\PELRVRPH SODVPD PHPEUDQH6FKOHUHWDO 7KH V\PELRVRPH VSDFH ZKLFK LV HIIHFWLYHO\ WKH LQWHUIDFH EHWZHHQ WKH Nostoc FHOO ZDOO DQG WKH IXQJDO SODVPD PHPEUDQH KDV VLPLODULWLHV ZLWK WKH V\PELRWLF LQWHUIDFH EHWZHHQ WKH IXQJDO ZDOO DQG WKH SODQW SODVPD PHPEUDQH LQ DUEXVFXODU P\FRUUKL]DV 6FKOHU HW DO ,QGHHG WKH IXQJXV Geosiphon pyriforme LV FORVHO\ UHODWHGWRWKH DUEXVFXODUP\FRUUKL]DIRUPLQJIXQJLRI WKHJHQXVGlomus 6FKOHUHWDO
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zolla OHDI FDYLW\ EUDQFKHG DQG VLPSOH KDLUV )LJ ' SRVVHVV WUDQVIHU FHOO FKDUDFWHULVWLFV DQG DUH WKRXJKW WR SOD\ D UROH LQ H[FKDQJH RI QXWULHQWV EHWZHHQ WKH F\DQRELRQW DQG WKH SODQW &DOYHUW HW DO %UDXQ+RZODQG DQG 1LHU]ZLFNL%DXHU %UDQFKHG KDLUV DUH WKRXJKW WR EH LQYROYHG LQ QLWURJHQ H[FKDQJH DQG VLPSOH KDLUV
LQFDUERQ H[FKDQJH &DOYHUWHWDO $VLPLODU WUDQVIHUFHOOPRUSKRORJ\LVVHHQLQWKHHORQJDWHGFHOOV WKDW WUDYHUVH WKH RSHQ DUHD RFFXSLHG E\ F\DQREDFWHULD EHWZHHQ WKH LQQHU DQG RXWHU FRUWH[ RI WKH F\FDG FRUDOORLG URRWV )LJ & /LQGEODG HW DO ,Q OLFKHQV WKHUH LV FORVH SK\VLFDO FRQWDFW EHWZHHQ WKH F\DQRELRQW DQG WKH IXQJDO K\SKDH ZLWKRXW WKH ODWWHU SHQHWUDWLQJ WKH F\DQREDFWHULDO FHOO ZDOO 5DL E ,Q DGGLWLRQ IXQJDO KDXVWRULD ZKLFK DUH VSHFLDOLVHG SHJOLNH SURMHFWLRQV RI WKH K\SKDH FDQ EH IRXQG FORVHO\ DVVRFLDWHG ZLWK F\DQREDFWHULDO ILODPHQWV DQG PD\ SOD\ D UROH LQ QXWULHQW H[FKDQJH 5DL -DQVRQ HW DO $KPDGMLDQ +RZHYHU WKHUH LV FRQIXVLRQ DERXWWKHIUHTXHQF\DQGIXQFWLRQRIKDXVWRULDDQGWKH H[WHQWWRZKLFKWKH\FDQSHQHWUDWHWKHF\DQRELRQWFHOO ZDOOV$KPDGMLDQ ,Q FRQWUDVW QR DGGLWLRQDO VWUXFWXUHV DUH QHHGHG LQ Gunnera DQG Geosiphon pyriforme EHFDXVH WKH LQWUDFHOOXODU ORFDWLRQ RI WKH F\DQRELRQWHQVXUHVHIILFLHQWH[FKDQJHRIPHWDEROLWHV
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zolla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ostoc punctiforme WKDW LV IROG PRUH LQIHFWLYH WKDQ WKH ZLOGW\SH LQ Anthoceros &RKHQ HW DO 7ZR JHQHV hrmA DQG hrmU, ZHUH LGHQWLILHGDVSDUWRIDQRSHURQ DQGH[SUHVVLRQRIERWK ZDV LQGXFHG E\ DQ DTXHRXV H[WUDFW RI Anthoceros WLVVXH EXW QRW E\ WKH KRUPRJRQLD LQGXFLQJ IDFWRU +,) RIWKHKRUQZRUW 7KH DTXHRXV H[WUDFW FRQWDLQV D KRUPRJRQLD UHSUHVVLQJ IDFWRU +5) WKDW LQKLELWV +,)LQGXFHG KRUPRJRQLD IRUPDWLRQ LQ ZLOGW\SH 1 punctiforme, DQG LW VHHPV WKDW WKH PXWDQW LV PRUH LQIHFWLYH EHFDXVH LW FRQWLQXHV WR UHVSRQG WR +,) SURGXFLQJ KRUPRJRQLD DQG H[WHQGLQJ LWV LQIHFWLYH VWDWH 7KHVH REVHUYDWLRQV LPSO\ WKDW WKH JHQH SURGXFWV RI WKH hrmUA RSHURQ EORFN KRUPRJRQLXP IRUPDWLRQ SHUKDSV E\ WKH SURGXFWLRQ RI DQ LQKLELWRU RU E\ FDWDEROLVP RIDQ DFWLYDWRU &RKHQ DQG 0HHNV
% 0RUSKRORJLFDO0RGLILFDWLRQV 7KH JURVV FHOOXODU PRUSKRORJ\ RI F\DQREDFWHULD LV IUHTXHQWO\ DOWHUHG LQ V\PELRVLV ,Q OLYHUZRUWV DQG KRUQZRUWV F\DQRELRQW YHJHWDWLYH FHOOV EHFRPH HQODUJHG DQG VSKHULFDO DQG WKH FHOO WR FHOOMXQFWLRQV ZHDNHQ VXFKWKDWHYHQJHQWOHSUHVVXUHZLOOFDXVH WKH FHOOV WR SDUW FRPSDQ\ )LJ ) 0HHNV ,Q F\DQROLFKHQV ILODPHQWRXV F\DQRELRQWV VXFK DV Scytonema DQG Calothrix FDQ EHFRPH XQLFHOOXODU 5DL DQG Nostoc FHOO VL]H LQFUHDVHV $KPDGMLDQ ,Q PDWXUH Azolla OHDI FDYLWLHV YHJHWDWLYH FHOO VL]H LQFUHDVHV VLJQLILFDQWO\ DQG DOWKRXJKFRQVLVWHQWZLWKLQHDFKILODPHQWFHOOVL]HDQG VKDSH FDQ EH KLJKO\ YDULDEOH EHWZHHQ ILODPHQWV %UDXQ+RZODQGDQG1LHU]ZLFNL%DXHU 3HWHUV 7KH Nostoc F\DQRELRQW RI Cycas revoluta LQFUHDVHV LQ FHOO VL]H ZKHQ V\PELRWLFDOO\ DVVRFLDWHG 6KDUPDHWDO DVGRHVWKHNostoc V\PELRQWRI Geosiphon pyriforme DIWHUSKDJRF\WRVLVE\WKHIXQJDO KRVW .OXJH HW DO 0ROOHQKDXHU HW DO
&KDSWHU 6\PELRWLF,QWHUDFWLRQV
Table 2. +HWHURF\VWIUHTXHQF\DQGJOXWDPLQHV\QWKHWDVHFKDUDFWHULVWLFVRIV\PELRWLFDOO\DVVRFLDWHGF\DQREDFWHULD
+RVW
+HWHURF\VW IUHTXHQF\D
*OXWDPLQH
V\QWKHWDVH
$PRXQW RI SURWHLQE
6SHFLILF DFWLYLW\E
)RUP RI FRPELQHG QLWURJHQ UHOHDVHG
&\FDGV
FLWUXOOLQHJOXWDPLQH
Gunnera
1+
+RUQZRUWVDQGOLYHUZRUWV
1+
Azolla
1+
/LFKHQV ELSDUWLWH WULSDUWLWH
1+ 1+
D
+HWHURF\VWIUHTXHQFLHVDUHH[SUHVVHGDVDSHUFHQWDJHRIWRWDOFHOOV 7\SLFDOYDOXHVIRUIUHHOLYLQJF\DQREDFWHULDDUH 9DOXHVDUHIRUWKHV\PELRQWDVDSHUFHQWDJHRIWKHVDPHF\DQREDFWHULXPLQWKHIUHHOLYLQJVWDWH 7DEOHFRPSLOHGIURP%HUJPDQHWDOE DQG5DLD E
E
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zolla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zolla F\DQRELRQW KDYH EHHQ WDNHQ WR LQGLFDWH WKDW KHWHURF\WV DUH VWLOO IXQFWLRQDO LQ EDVDO OHDIFDYLWLHV&DQLQLHWDO D
,Q F\FDG FRUDOORLG URRWV WKH IUHTXHQF\ RI KHWHURF\VWVLV ORZHVWDWWKHURRWWLS DQGKLJKHVWLQWKH EDVDO ROGHU SDUWV ZKHUH KLJK IUHTXHQFLHV RI WZR WKUHH RU HYHQ IRXU DGMDFHQW KHWHURF\VWV DUH IRXQG /LQGEODG HW DO D E /LQGEODG DQG %HUJPDQ 3HWHUV 6LPLODUO\ KHWHURF\VW IUHTXHQF\ LQ Gunnera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
V\PELRWLF FRORQLHV DQG WKHUH DUH UHDVRQV WR EHOLHYH WKDW WKH PHFKDQLVPV WKDW UHJXODWH KHWHURF\VW GHYHORSPHQWDQGIUHTXHQF\LQV\PELRVLVDUHGLIIHUHQW WR WKRVH RSHUDWLQJ LQ WKH IUHHOLYLQJ VWDWH )RU H[DPSOH LQ IUHHOLYLQJ F\DQREDFWHULDWKH VWLPXOXV WR GHYHORS KHWHURF\VWV LV QLWURJHQ VWDUYDWLRQ :RON HW DO \HW LQ V\PELRVLV ZLWK SODQWV WKH F\DQREDFWHULD VKRZ QR VLJQ RI QLWURJHQ VWDUYDWLRQ VHHWKHHQGRI6HFWLRQ9,&IRUDGLVFXVVLRQRIWKLV )XUWKHU HYLGHQFH FRPHV IURP VWXGLHV RI D Nostoc PXWDQWGHIHFWLYHLQQLWUDWHDVVLPLODWLRQ&DPSEHOODQG 0HHNV 1LWUDWH IDLOV WR UHSUHVV QLWURJHQ IL[DWLRQDQGKHWHURF\VWGHYHORSPHQWLQWKHIUHHOLYLQJ PXWDQW VWUDLQ EXW UHSUHVVHV ERWK ZKHQ WKH VWUDLQ LV V\PELRWLFDOO\ DVVRFLDWHG ZLWK WKH KRUQZRUW Anthoceros. $PPRQLXP LV DQ HIIHFWLYH UHSUHVVRU LQ ERWK ZLOGW\SH DQG PXWDQW ZKHWKHU IUHHOLYLQJ RU V\PELRWLFDOO\ DVVRFLDWHG 7KH UHSUHVVLYH HIIHFW RI QLWUDWHLQWKHV\PELRWLFDOO\DVVRFLDWHGPXWDQWGRHVQRW UHVXOW IURP WKH DFFXPXODWLRQ RI 1+ JHQHUDWHG E\ WKHUHGXFWLRQRI12 WR1+ LQ Anthoceros, EHFDXVH 1+ SRROV DUH WKH VDPH LQ WLVVXH IURP KRUQZRUW JURZQ LQ WKH SUHVHQFH DQG DEVHQFH RI 12 7KH LPSOLFDWLRQ RI WKHVH REVHUYDWLRQV LV WKDW QLWURJHQDVH H[SUHVVLRQ DQG KHWHURF\VW GHYHORSPHQW LQ V\PELRWLFDOO\ DVVRFLDWHG Nostoc DUH FRQWUROOHG E\ SODQWVLJQDOV $ JUHDWHU XQGHUVWDQGLQJ RIWKH IDFWRUV LQYROYHG LQ WKH UHJXODWLRQ RIKHWHURF\VW GHYHORSPHQW LQ V\PELRVLV PD\ FRPH IURP WKH DSSOLFDWLRQ RI PROHFXODU JHQHWLF WHFKQLTXHV WR F\DQRELRQWV VXFK DV Nostoc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amia DQG Cycas LQ ZKLFK KHWHURF\VW IUHTXHQF\ LV KLJKHVWLQWKHEDVDOUHJLRQ \HWQLWURJHQDVHDFWLYLW\ LV KLJKHVW DW WKH FRUDOORLG URRW WLS /LQGEODG DQG
'*$GDPV %HUJPDQ 1RWZLWKVWDQGLQJ WKHVH REVHUYDWLRQV WKHUH LV D JHQHUDO FRUUHODWLRQ EHWZHHQ WKH LQFUHDVH LQ KHWHURF\VW IUHTXHQF\ RI PDQ\ V\PELRWLFDOO\ DVVRFLDWHG F\DQREDFWHULD DQG HOHYDWHG UDWHVRIQLWURJHQIL[DWLRQ
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zolla *ULOO &DLROD HW DO 0XWXVNLQ DQG .ROHVQLNRY LQ ZKLFK DGGLWLRQDO SURWHFWLRQ IURP VXSHUR[LGH UDGLFDOV PD\ EH SURYLGHG E\ LURQ VXSHUR[LGH GLVPXWDVH VKRZQ E\ LPPXQRJROG ODEHOOLQJ WR EH KLJKHVW LQ F\DQRELRQWV LQ OHDYHV ZLWK WKHKLJKHVWQLWURJHQDVHDFWLYLW\&DQLQLHWDO a ,Q FRQWUDVW WKH V\PELRWLF FRORQLHV LQ WKH F\FDG Mucrozamia riedlei DUH WKRXJKW WR EH PLFURDHURELF DQG WKHIUHVKO\LVRODWHGF\DQRELRQWVH[KLELWLQFUHDVHG VHQVLWLYLW\ WR R[\JHQ /LQGEODG HW DO E SRVVLEO\ EHFDXVH RI WKH LQFRPSOHWH GHYHORSPHQW RI KHWHURF\VW ZDOO OD\HUV LQ WKH DEVHQFH RI R[\JHQ 5LSSNDDQG6WDQLHU ,Q PDQ\ F\DQREDFWHULDSODQW V\PELRVHV WKH F\DQREDFWHULD DUH FDSDEOH RI VXSSO\LQJ WKH HQWLUH QLWURJHQUHTXLUHPHQWVRIWKHKRVW ,QHYHQWKHODUJHVW Gunnera SODQWVZLWKOHDYHVWZRPHWHUVDFURVVWKLVLV DFKLHYHGE\DF\DQRELRQWPDVVWKDWPD\EHDVOLWWOHDV RIWKHWRWDOSODQWZHLJKW2VERUQH HWDO ,QGHHG LQ WKH FDVH RI Gunnera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
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV UDWKHU WKDQ WRWDO KHWHURF\VWV SRVVLEO\ EHFDXVH WKH PDQ\ PXOWLSOH DGMDFHQW KHWHURF\VWV SUHVHQW LQ IRU H[DPSOH Gunnera PD\ IXQFWLRQ LQHIILFLHQWO\ DV D UHVXOW RI SRRU WUDQVIHU RI SKRWRV\QWKDWH IURP YHJHWDWLYH FHOOV %HUJPDQ HW DO ,Q Gunneru DW OHDVW WKH ORZHUHG UDWH RI QLWURJHQDVH DFWLYLW\LQROGHUWLVVXHVLVQRWGXHWRUHGXFHGOHYHOVRI QLWURJHQDVH SURWHLQ EXW LV PRVW OLNHO\ GXH WR D UHGXFWLRQ LQWKH VXSSO\ RISKRWRV\QWKDWHV 6RGHUEDFN HWDO 'LQLWURJHQ IL[HG E\ WKH F\DQRELRQW LV UHOHDVHG WR WKH KRVW DV DPPRQLD LQ WKH FDVH RI Anthoceros, Blasia, Azolla, Gunneru 0HHNV HW DO D E %HUJPDQHWDO 6LOYHVWHUHWDO DQGPDQ\ OLFKHQV5DLHWDO DQGLQDQXQNQRZQRUJDQLF IRUP SRVVLEO\ FLWUXOOLQH RU JOXWDPLQH LQ F\FDGV 7DEOH 7KH SURSRUWLRQ RI FRPELQHG QLWURJHQ UHWDLQHG E\ WKH V\PELRQW FDQ EH DV OLWWOH DV LQ VRPHOLFKHQV5DL D$KPDGMLDQ LQ Gunneru 6LOYHVWHUHWDO LQWKHKRUQZRUW Anthoceros 0HHNV HW DO D DQG LV WKRXJKW WR EH LQAzollu 0HHNVHWDO E 3HWHUV DQG0HHNV ,QWKHFDVHRIAzolla WKLVOHDGV WR DPPRQLXPFRQFHQWUDWLRQV RI P0 LQ OHDIFDYLWLHV DWWKHDSH[WRP0LQWKHEDVDOROGHVW OHDIFDYLWLHV &DQLQL HW DO $PPRQLD UHOHDVH E\ WKH F\DQRELRQW LV D FRQVHTXHQFH RIGHFUHDVHG DFWLYLW\ RI JOXWDPLQH V\QWKHWDVH WKH ILUVW HQ]\PH LQ WKH JOXWDPLQH V\QWKHWDVHJOXWDPDWH V\QWKDVH *6 *2*$7 SDWKZD\ ZKLFK LV WKH SULPDU\ URXWH RI DPPRQLD DVVLPLODWLRQ LQ F\DQREDFWHULD DQG LQ EU\RSK\WHVDQGAzolla 0HHNV %HUJPDQHWDO EXW QRW LQ OLFKHQV ZKLFK HPSOR\ JOXWDPDWH GHK\GURJHQDVH*'+5DL E 7KHUH DUH VHYHUDO PHDQV E\ ZKLFK WKH GHFUHDVH LQ *6 DFWLYLW\LVDFKLHYHG7DEOH ,QAzolla LWUHVXOWV IURP D GHFUHDVH LQ WKH DPRXQW RI HQ]\PH LQ WKH F\DQRELRQW /HH HW DO 3HWHUV DQG 0HHNV ZKLFK FRUUHODWHV ZLWK D WHQIROG GHFUHDVH LQ WKHOHYHORI*6 (glnA) P51$1LHU]ZLFNL%DXHUDQG +DVHONRUQ 6LPLODUO\LQF\DQROLFKHQVERWK*6 DQG*2*$7DFWLYLW\LVUHGXFHGE\RYHUGXHWRD UHGXFWLRQ LQ WKH DPRXQW RI*6 SURWHLQ 5DL D ,Q FRQWUDVW LQ WKH KRUQZRUW Anthoceros DQ XQGHWHUPLQHG SRVWWUDQVODWLRQDO PRGLILFDWLRQ RI WKH HQ]\PHLVWKRXJKWWREHUHVSRQVLEOHIRUWKHGHFUHDVHG *6 DFWLYLW\ EHFDXVH WKH DPRXQW RI *6 SURWHLQ LQ IUHHOLYLQJ DQG V\PELRWLFDOO\ DVVRFLDWHG Nostoc LV OLWWOHGLIIHUHQW7DEOH-RVHSKDQG0HHNV /HH HW DO 5DL HWDO 0HHNV ,Q WKH Gunneru V\PELRQW*6VSHFLILFDFWLYLW\LVRIWKDW LQWKHIUHHOLYLQJVWDWHDOWKRXJKWWKHDPRXQWRISURWHLQ
LV XQFKDQJHG 7DEOH %HUJPDQ HW DO E 6LOYHVWHUHWDO ,Q FRQWUDVW *6 DFWLYLWLHV DQG SURWHLQ OHYHOV LQ F\FDGV DUH OLWWOH FKDQJHG LQ WKH V\PELRWLFDOO\ DVVRFLDWHG Nostoc /LQGEODG DQG %HUJPDQ DQG WKLV LV FRQVLVWHQW ZLWK WKH UHOHDVH RIRUJDQLF QLWURJHQ UDWKHU WKDQ 1+ WR WKH SODQW7DEOH3DWHHWDO ,Q WKH F\DQROLFKHQV +DOOERP HW DO EU\RSK\WHV5DLHWDO DQGAzolla %HUJPDQDQG 5DL WKH OHYHO RI*6 SURWHLQ LQ KHWHURF\VWV LV VLPLODU WR WKDW RI YHJHWDWLYH FHOOV +RZHYHU LQ WKH DSLFDO UHJLRQV RIWKH OLFKHQ Lichina confinis WKDOOXV *6 SURWHLQ FDQQRW EH GHWHFWHG LQ KHWHURF\VWV RI WKH Calothrix V\PELRQW DOWKRXJK LW UHPDLQV UHODWLYHO\ DEXQGDQW LQ YHJHWDWLYH FHOOV -DQVRQ HW DO 7KLV FRQWUDVWV ZLWK WKH VLWXDWLRQ LQ IUHHOLYLQJ F\DQREDFWHULD LQ ZKLFK KHWHURF\VWV KDYH DSSUR[LPDWHO\ WZLFH WKH *6 DFWLYLW\ DQG SURWHLQ OHYHOVRIYHJHWDWLYHFHOOV%HUJPDQHWDO 5DLHW DO 5HQVWURP.HOOQHUHWDO %HUJPDQHW DOE ,QLWLDO XSWDNH RI WKH DPPRQLD UHOHDVHG E\ WKH F\DQRELRQWV LQ EU\RSK\WHV DQG Azolla RFFXUV YLD WKH *6*2*$7 SDWKZD\ RI WKH KRVW 0HHNV HW DO E3HWHUVHWDO 0HHNV 5DL ZKHUHDV LQ F\DQROLFKHQV LW RFFXUV YLD IXQJDO JOXWDPDWH GHK\GURJHQDVH 5DL 7KH IRUP LQ ZKLFK QLWURJHQ LV WUDQVSRUWHG IURP WKH V\PELRWLF WLVVXH WR WKH UHVW RIWKH KRVW LV NQRZQ LQ RQO\ D IHZ FDVHV ,Q Azollu WKH PDMRU FRPSRXQGV PRYLQJ IURP WKH OHDI FDYLW\ WR WKH VWHP DSH[ DUH WKRXJKW WR EH JOXWDPDWH JOXWDPLQH DPPRQLD DQG SRVVLEO\ D JOXWDPDWH GHULYDWLYH 3HWHUV HW DO %UDXQ +RZODQG DQG 1LHU]ZLFNL%DXHU %HUJPDQ HW DO ZKHUHDVLQF\FDGVJOXWDPDWHDQGFLWUXOOLQH PRYH IURP WKH URRWV WR WKH VWHP 3DWH HW DO %HUJPDQ HW DO DQG LQ WULSDUWLWH F\DQROLFKHQV DODQLQH LV WKRXJKW WR EH IRUP RIQLWURJHQ WUDQVIHUUHG IURP WKH FHSKDORGLDWR WKH PDLQ WKDOOXV 5DL ,Q Gunneru monoica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
F\DQRELRQW 7KHUH DUH VHYHUDO UHDVRQV IRU WKLV FRQFOXVLRQ )LUVWO\ WKH SK\FRELOLSURWHLQV ZKLFK VHUYH DV ERWK SKRWRV\QWKHWLF DFFHVVRU\ SLJPHQWV DQG QLWURJHQUHVHUYHVLQF\DQREDFWHULD DUH VWLOOSUHVHQWLQ WKH F\DQRELRQWV RI EU\RSK\WHV 5DL HW DO 0HHNV Azolla .DSODQ HW DO %UDXQ +RZODQG DQG 1LHU]ZLFNL%DXHU F\FDGV /LQGEODG DQG %HUJPDQ DQG Gunnera 6RGHUEDFN DQG %HUJPDQ 6HFRQGO\ WKH XQLTXHF\DQREDFWHULDO QLWURJHQ UHVHUYH F\DQRSK\FLQ D FRSRO\PHU RI DUJLQLQH DQG DVSDUWLF DFLG 6LPRQ LV DOVR SUHVHQW LQ WKH F\DQRELRQWV RI EU\RSK\WHV5DLHWDO Gunnera 6RGHUEDFNHW DO -DJHUHWDO F\FDGV/LQGEODGHWDO D DQGAzolla %HUJPDQHWDO E 1LWURJHQ VWDUYDWLRQ ZRXOG EH H[SHFWHG WR UHVXOW LQ EUHDNGRZQ RI QLWURJHQ UHVHUYHV DQG KHQFH D UHGXFWLRQ LQ WKH FHOOXODU FRQWHQWV RI ERWK SK\FRELOLSURWHLQV DQG F\DQRSK\FLQ
' &2 $VVLPLODWLRQ DQG 7UDQVIHURI&DUERQ 7KH SULPDU\ URXWH RI&2 IL[DWLRQ LQ IUHHOLYLQJ DQG V\PELRWLFDOO\DVVRFLDWHG F\DQREDFWHULD LV WKH &DOYLQ ELVSKRVSKDWH F\FOH ZLWK ULEXORVH FDUER[\ODVHR[\JHQDVH 5XELVFR DV WKH SULPDU\ FDUER[\ODWLQJ HQ]\PH 7DELWD 7KH UDWHV RI OLJKWGHSHQGHQW 2 HYROXWLRQ DQG &2 IL[DWLRQ LQ WKH F\DQRELRQWV RI EU\RSK\WHV 0HHNV F\FDGV /LQGEODG DQG %HUJPDQ /LQGEODG HW DO E DQG Gunnera %RQQHWW 6RGHUEDFNDQG %HUJPDQ DUH HLWKHU JUHDWO\ UHGXFHG RU XQGHWHFWLEOH LPPHGLDWHO\ DIWHU WKHLU VHSDUDWLRQ IURP WKH KRVW SODQW 7DEOH ,Q WKHVH FDVHV WKH F\DQRELRQW JURZV SKRWR RU FKHPRKHWHURWURSKLFDOO\ UHFHLYLQJ IL[HG FDUERQ IURP LWV SKRWRV\QWKHWLF KRVW SUREDEO\ LQ WKH IRUP RI VXFURVH 6WHLQEHUJ DQG 0HHNV %HUJPDQ HW DO 6RGHUEDFN DQG %HUJPDQ DV LV WKH FDVH ZLWK Azolla 3HWHUV DQG 0HHNV %UDXQ+RZODQG DQG 1LHU]ZLFNL %DXHU 3HWHUV 7KHF\DQRELRQWRIAzolla FDQ IL[ &2 EXWWKLV SUREDEO\ DFFRXQWV IRU QR PRUH WKDQ RI WKH FDUERQ IL[DWLRQ RI WKH V\PELRVLV .DSODQDQG3HWHUV 3HWHUVDQG0HHNV DQG WKH OHYHO RI5XELVFR P51$ LV DW OHDVW ILYHIROG ORZHU WKDQ LQ IUHHOLYLQJ Anabaena 1LHU]ZLFNL %DXHU DQG +DVHONRUQ 0RVW &2 IL[DWLRQ LV WKRXJKW WR RFFXU LQ YHJHWDWLYH FHOOV DW WKH DSLFDO PHULVWHP RI Azolla, ZLWK OLWWOH LQ WKH ROGHU OHDYHV %UDXQ+RZODQG DQG 1LHU]ZLFNL%DXHU 6XUSULVLQJO\ LPPHGLDWHO\DIWHULVRODWLRQIURPAzolla, Anabaena azollae KDV D UDWH RI OLJKWGHSHQGHQW &2
'*$GDPV IL[DWLRQ VLPLODU WR WKDW RI WKH IUHHOLYLQJ RUJDQLVP 7DEOH ,PPHGLDWHO\DIWHULWVVHSDUDWLRQIURPV\PELRVLVWKH Nostoc V\PELRQW RI Anthoceros KDV D UDWH RI OLJKW GHSHQGHQW &2 IL[DWLRQ HLJKWIROG ORZHU WKDQ WKH VDPH F\DQREDFWHULXP JURZQ LQ WKH IUHHOLYLQJ VWDWH 7DEOH 6WHLQEHUJDQG0HHNV 0HHNV 7KH OHYHO RI5XELVFR SURWHLQ LV OLWWOH GLIIHUHQW LQ WKH WZR FDVHV 6WHLQEHUJ DQG 0HHNV 5DL HW DO 0HHNV LPSO\LQJ WKDW DFWLYLW\ LV UHJXODWHG E\ SRVW WUDQVODWLRQDO PRGLILFDWLRQ RI WKH HQ]\PH E\ DQ XQNQRZQ PHFKDQLVP 6WHLQEHUJ DQG 0HHNV 0HHNV ,Q FRQWUDVW WKHIUHVKO\ LVRODWHG F\DQRELRQW RI Gunnera chilensis VKRZV 5XELVFR DFWLYLW\ KLJKHU WKDQ WKH IUHHOLYLQJ Nostoc, HYHQ WKRXJK WKH V\PELRWLFDOO\ DVVRFLDWHG V\PELRQW KDV ORZHUHG &2IL[LQJ DFWLYLW\ 7DEOH 6RGHUEDFN DQG %HUJPDQ 7KLV UXOHV RXW D SRVW WUDQVODWLRQDO FRQWURO VXFK DV WKDW LQ Anthoceros. ,W VHHPV WKDW WKH SKRWRV\QWKHWLF PDFKLQHU\ RI WKH V\PELRQW LV ODUJHO\ LQWDFW ZLWK FKORURSK\OO D 6RGHUEDFN HW DO SK\FRELOLSURWHLQV DQG 5XELVFR 6RGHUEDFN DQG %HUJPDQ EHLQJ SUHVHQW DW OHYHOV HTXLYDOHQW WR WKH IUHHOLYLQJ V\PELRQW DQG WKDW D UHGXFWLRQ LQ SKRWRV\QWKHWLF HOHFWURQ WUDQVSRUW LV UHVSRQVLEOH IRU WKH UHGXFHG LQ vivo &2IL[LQJDFWLYLW\ RIWKH F\DQRELRQW $ VLPLODU VLWXDWLRQPD\DULVHLQF\FDGVVXFKDV Cycas revoluta, LQ ZKLFK GHVSLWH WKH ODFN RI in vivo &2 IL[DWLRQ FRQVLGHUDEOH 5XELVFR DFWLYLW\ FDQ EH PHDVXUHG LQ FUXGH H[WUDFWV RI WKH IUHVKO\ LVRODWHG F\DQRELRQW 7DEOH /LQGEODG HW DO /LQGEODG DQG %HUJPDQ $Q H[FHSWLRQ PD\ EH WKH F\DQRELRQWIURP Cycas circinalis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
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV
Table 3. &KDUDFWHULVWLFV RI OLJKWGHSHQGHQW &2 IL[DWLRQ DQG ULEXORVH ELVSKRVSKDWH FDUER[\ODVH 5XELVFR LQ V\PELRWLFDOO\ DVVRFLDWHG F\DQREDFWHULD
+RVW
/LJKWGHSHQGHQW &2 IL[DWLRQD
5XELVFR $PRXQW RI SURWHLQD
6SHFLILF DFWLYLW\D
Azolla
a F
QGE
QG
+RUQZRUW$punctatus
G
G
G
&\FDGV
H
QG
H
Gunnera
/LFKHQV ELSDUWLWH WULSDUWLWH
QG aL
I
J
aK aK
I QG aL
D E 9DOXHVDUHH[SUHVVHGDVDSHUFHQWDJHRIWKRVHIRUWKHIUHHOLYLQJF\DQREFDWHULD 7KH DPRXQW RISURWHLQ KDV QRW EHHQ GHWHUPLQHG EXW 5XELVFR P51$ LV RI WKDW LQ IUHHOLYLQJ Anabaena 1LHU]ZLFNL%DXHU DQG +DVHONRUQVHH6HFWLRQ9,' F G H 7DEOH FRPSLOHG IURP WKH IROORZLQJ UHIHUHQFHV 3HWHUV DQG 0HHNV 6WHLQEHUJ DQG 0HHNV /LQGEODG HW DO I 6RGHUEDFNDQG%HUJPDQ 6|GHUElFNDQG %HUJPDQ %HUJPDQDQG5DL L5DL HWDO QG QRWGHWHUPLQHG
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i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
'*$GDPV
D Richeliu intracellularis, WKH F\DQRELRQW RI Rhizosoleniu, IL[HV ERWK &2 DQG 1 IRU LWV GLDWRP KRVW 5DL D ,Q FRQWUDVW WKH XQLFHOOXODU F\DQRELRQWV RI WKH GLQRIODJHOODWHV Ornithocerus magnifcus DQG Ornithocerus steinii, VHHP WR SURYLGH IL[HG FDUERQ UDWKHU WKDQ QLWURJHQ IRU WKHLU QRQ SKRWRV\QWKHWLF KRVW DOWKRXJK WKH\ PD\ DOVR EH SKDJRWURSKLFDOO\ FRQVXPHG ZKHQ WKH\ VHQHVFH -DQVRQHWDOD
9,, 5HFRQVWLWXWLRQRIWKH6\PELRVHV 5HFRQVWUXFWLRQ RI V\PELRVHV E\ LQWURGXFLQJ D[HQLF FXOWXUHV RI WKH WZR V\PELRWLF SDUWQHUV KDV EHHQ VXFFHVVIXOO\DFKLHYHG ZLWK DOO RIWKHSODQW V\PELRVHV H[FHSW Azolla, WKLVEHLQJWKHRQO\FDVHLQZKLFKLWKDV QRW EHHQ SRVVLEOH WR JURZ WKH PDMRU F\DQRELRQW LQ IUHHFXOWXUH +RZHYHU WKHDUWLILFLDOUHFRQVWUXFWLRQ RI WKH Azolla V\PELRVLVKDVEHHQDFKLHYHGE\UHPRYDORI D PHJDVSRURFDUS DSLFDO FDS FRQWDLQLQJ WKH F\DQRELRQW DQG WUDQVIHUULQJ WKLV WR WKH GHFDSLWDWHG PHJDVSRURFDUS RI DQRWKHU Azolla SODQW /LQ HW DO /LQ DQG :DWDQDEH :DWDQDEH HW DO %UDXQ+RZODQG DQG 1LHU]ZLFN%DXHU ,Q FRQWUDVW WKH EU\RSK\WH V\PELRVHV KDYH SURYHG SDUWLFXODUO\ HDV\ WR UHFRQVWLWXWH 3ODWH H QRWMXVW ZLWK WKH RULJLQDO F\DQRELRQW EXW ZLWK F\DQRELRQWV IURP Gunnera, F\FDGV DQG HYHQ IUHHOLYLQJ VWUDLQV 0HHNV.LPXUDDQG1DNDQR $OO RI WKH V\PELRWLFDOO\FRPSHWHQW F\DQREDFWHULD DUH KRUPRJRQLDIRUPLQJ VWUDLQV RI WKH JHQXV Nostoc. 6LPLODU REVHUYDWLRQV KDYH EHHQ PDGH ZLWK Gunnera ZKLFK FDQ EH LQIHFWHG E\ ERWK V\PELRWLF DQG IUHH OLYLQJ Nostoc VWUDLQV %RQQHWW %HUJPDQHWDO E -RKDQVVRQ DQG %HUJPDQ EXW QRW E\ KRUPRJRQLDIRUPLQJ F\DQREDFWHULD RI RWKHU JHQHUD VXFK DV Calothrix, Fischerella DQG Chlorogloeopsis -RKDQVVRQDQG%HUJPDQ 5HFRQVWLWXWLRQ H[SHULPHQWV VXFK DV WKHVH DUH YHU\ KHOSIXO LQ VWXG\LQJ WKH LQIHFWLRQ SURFHVV DQG WKH HVWDEOLVKPHQWRIV\PELRWLFFRORQLHV DQG FDQSURYLGH LQIRUPDWLRQ RQ WKH VSHFLILFLW\ RI WKH DVVRFLDWLRQ 7KH\FRQILUPWKHEURDGFRPSHWHQFHRINostoc VSS DV V\PELRQWVEXWDUHOLNHO\WRJLYHDIDOVHLPSUHVVLRQRI WKH EUHDGWK RI VWUDLQV FDSDEOH RI LQIHFWLQJ SODQWV LQ WKH ILHOG EHFDXVH VXFK H[SHULPHQWV DUH DUWLILFLDO LQ WKHLU XVH RIDVLQJOH D[HQLF SODQWDQG D VLQJOH D[HQLF F\DQREDFWHULXP (YHQFRPSHWLWLRQH[SHULPHQWVXVLQJ VHYHUDO F\DQREDFWHULD DW RQFH 0HHNV GR QRW PLPLF WKH QDWXUDO HQYLURQPHQW LQ ZKLFK PDQ\ F\DQREDFWHULDO VWUDLQV ZLOOEHSUHVHQW QRWWRPHQWLRQ WKH XQNQRZQ LQIOXHQFHV RI WKH PDQ\ RWKHU EDFWHULD
DQG HXNDU\RWHV )RU H[DPSOH Calothrix DQG Chlorogloeopsis, ERWK FDSDEOH RI IRUPLQJ KRUPRJRQLD FDQ LQIHFWPhaeoceros LQWKH ODERUDWRU\ \HW WKH ODWWHU KDV QRWEHHQ IRXQG LQ ILHOG VDPSOHV RI DQ\ EU\RSK\WH DQG WKH IRUPHU RQO\ YHU\ UDUHO\ SRVVLEO\ UHIOHFWLQJ ERWK WKH UHODWLYH DEXQGDQFH RI WKHVHVWUDLQVLQWKHVRLO DQG WKHLULQDELOLW\WRFRPSHWH ZLWK WKH QDWXUDO SRSXODWLRQV RI Nostoc :HVW DQG $GDPV $OWKRXJKLWLV RIWHQSRVVLEOHWR JURZVHSDUDWHO\WKH SKRWRELRQW DQG P\FRELRQW SDUWQHUV RI OLFKHQV UHV\QWKHVLV RI WKH V\PELRVLV KDV SURYHG H[WUHPHO\ GLIILFXOW 7KLV LV LQ SDUW EHFDXVH RI WKH YHU\ VORZ JURZWK UDWHV RI WKH P\FRELRQWV LQ SDUWLFXODU +RZHYHUWKHUHDUHDQLQFUHDVLQJQXPEHURIH[DPSOHV RI SDUWLDO RU FRPSOHWH UHV\QWKHVHV RI ERWK ELSDUWLWH DQG WULSDUWLWH OLFKHQV $KPDGMLDQ 6WRFNHU:RUJRWWHUDQG7XUN +LOO 6WRFNHU:RUJRWWHU ,QWKHFDVHRIGeosiphon, Nostoc VSS RWKHU WKDQ N. punctiforme FDQ EH LQFRUSRUDWHG E\ WKH IXQJXV DV HQGRV\PELRQWV .OXJH HW DO ,QGHHG LW LV WKRXJKW WKDW LQ QDWXUH Geosiphon DQGWKHEU\RSK\WHVBlasia DQG Anthoceros FDQH[FKDQJHF\DQRELRQWV0ROOHQKDXHU .OXJH HWDO $WWHPSWV WR FRQVWUXFW DUWLILFLDO DVVRFLDWLRQV EHWZHHQ V\PELRWLF DQG IUHHOLYLQJ F\DQREDFWHULD DQG SODQWV KDYH LQYROYHG WKH LQWURGXFWLRQ RI F\DQREDFWHULD LQWR KLJKHU SODQW SURWRSODVWV RU WKH JURZWKRIF\DQREDFWHULDZLWKSODQWWLVVXHFHOOFXOWXUHV DQG SODQW UHJHQHUDWHV *XVHY DQG .RU]KHQHYVND\D DQG.RU]KHQHYVND\DHWDO
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
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV PDLQWHQDQFH RISODQWF\DQREDFWHULD DVVRFLDWLRQV PD\ RSHQ WKHSRVVLELOLW\ RIHQKDQFLQJH[LVWLQJ V\PELRVHV RUHYHQHVWDEOLVKLQJQRYHORQHVZLWKIRUH[DPSOHFURS SODQWV &\DQREDFWHULD VXFK DV Nostoc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
$FNQRZOHGJHPHQWV 0DQ\ WKDQNV WR %LUJLWWD %HUJPDQ WKH ODWH 'DYLG +DOO 3HWHU /LQGEODG ,DQ /XFDV 6DQGUD 1LHU]ZLFNL %DXHU 'LHWHU 0ROOHQKDXHU DQG $UWKXU 6FKOHU IRU SURYLGLQJ SKRWRJUDSKV DQG IRU FRPPXQLFDWLQJ GDWD SULHUWRSXEOLFDWLRQ
5HIHUHQFHV $GDPV'* 0XOWLFHOOXODULW\LQF\DQREDFWHULD ,Q 0RKDQ6 'RZ&DQG&ROH-$HGV 3URNDU\RWLF6WUXFWXUHDQG)XQFWLRQ$ 1HZ3HUVSHFWLYH 6\PS6RF*HQ0LFURELRO $GDPV '* &\DQREDFWHULD ,Q 6KDSLUR - DQG 'ZRUNLQ 0 HGV %DFWHULDDV 0XOWLFHOOXODU2UJDQLVPV SS 2[IRUG 8QLYHUVLW\3UHVV1HZ
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¶V VXQVFUHHQV"1DWXUDO3URGXFW5HSRUWV %DQGDUDQD\DNH:0 %HPLV-( DQG%RXUQH '- 8OWUDYLROHW DEVRUELQJ SLJPHQWV IURP WKH PDULQH VSRQJH '\VLGHD KHUEDFHD ,VRODWLRQ DQG VWUXFWXUH RI D QHZ P\FRVSRULQH &RPS %LRFKHP 3K\VLRO & 3KDUPDFRORJ\ WR[LFRORJ\ DQG HQGRFULQRORJ\ %DU ( DQG 7HO2U ( (IIHFW RI OLJKW DQG R[\JHQ RQ QLWURJHQDVH DFWLYLW\ DQG GLQLWURJHQDVH UHGXFWDVH )HSURWHLQ FRQWHQW LQ $]ROOD$QDEDHQD DVVRFLDWLRQ - 3ODQW 3K\VLRO %HUHQJXHU- 5RMR) GH3HGUR0$ 3IDQ]DJO % DQG/|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
VWUDLQ F\DQREDFWHULD LVRODWHG IURP Geosiphon pyriforme P\FRWD -3K\FRO %ODQN 5- ,V WKH FHQWUDO LQFOXVLRQ ERG\ RI WKH F\DQHOOH RI Cyanophora paradoxa DFDUER[\VRPH"(QGRF\WRELRVLV&HOO5HV %RJQHU( :DVWOKXEHU5 6FKOHJO ,DQG /RRV ( *O\FRJHQ DP\ODVH DQG DOSKDJOXFRVLGDVH DV SRVVLEOH FRPSRQHQWV LQ WKH JOXFRVHUHOHDVH V\VWHP RI WKH F\DQRELRQW RI Peltigera horizontalis SDUWLDOSXULILFDWLRQ DQG FKDUDFWHUL]DWLRQ 6\PELRVLV %RQQHWW+7 7KH Nostoc-Gunnera DVVRFLDWLRQ ,Q 5DL $1 HG &5& +DQGERRN RI6\PELRWLF &\DQREDFWHULD SS &5&3UHVV%RFD5DWRQ)ORULGD %UDVHOO+0 'DYLHV 6.DQG0DWWD\-3 1LWURJHQ IL[DWLRQ DVVRFLDWHG ZLWK EU\RSK\WHV FRORQL]LQJ EXUQW VLWHV LQ 6RXWKHUQ 7DVPDQLD$XVWUDOLD-%U\RO %UDXQ+RZODQG (% DQG 1LHU]ZLFNL%DXHU 6$ AzollaAnabaena V\PELRVLV ELRFKHPLVWU\ SK\VLRORJ\ XOWUDVWUXFWXUH DQG PROHFXODU ELRORJ\ ,Q 5DL $1 HG &5& +DQGERRN RI 6\PELRWLF &\DQREDFWHULD SS &5& 3UHVV %RFD 5DWRQ )ORULGD %UDXQ+RZODQG (% /LQGEODG 3 1LHU]ZLFNL%DXHU 6$ DQG %HUJPDQ % 'LQLWURJHQDVH UHGXFWDVH )HSURWHLQ RI QLWURJHQDVH LQ WKH F\DQREDFWHULDO V\PELRQWV RI WKUHH Azolla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zolla LPSOLHV DIXQFWLRQDO UROHLQPHWDEROLFH[FKDQJH3URWRSODVPD &DOYHUW+(3HUNLQV6.DQG3HWHUV*$ 6SRURFDUSVWUXFWXUH LQWKHKHWHURVSRURXVZDWHUIHUQAzolla mexicana 3UHVO 6FDQQLQJ (OHFWURQ0LFURVF &DPSEHOO(/+DJHQ.'&RKHQ0)6XPPHUV0/DQG0HHNV-& 7KH devR JHQH SURGXFW LV FKDUDFWHULVWLF RIUHFHLYHUV RI WZRFRPSRQHQWUHJXODWRU\V\VWHPVDQGLVHVVHQWLDOIRUKHWHURF\VW GHYHORSPHQW LQ WKH ILODPHQWRXV F\DQREDFWHULXP Nostoc VS VWUDLQ$7&&-%DFWHULRO &DPSEHOO(/DQG0HHNV-& &KDUDFWHULVWLFV RIKRUPRJRQLD IRUPDWLRQE\ V\PELRWLF Nostoc VSS LQUHVSRQVHWR WKHSUHVHQFH RI Anthoceros punctatus RU LWV H[WUDFHOOXODU SURGXFWV $SSO (QYLURQ0LFURELRO &DPSEHOO (/ DQG 0HHNV -& (YLGHQFH IRU SODQWPHGLDWHG UHJXODWLRQ RI QLWURJHQDVH H[SUHVVLRQ LQ WKH Anthoceros-Nostoc V\PELRWLFDVVRFLDWLRQ-*HQ0LFURELRO &DQLQL$%HUJPDQ%&LYLWDUHDOH3 5RWLOLR*DQG*ULOOL&DLROD0 F /RFDOL]DWLRQ RI LURQVXSHUR[LGH GLVPXWDVH LQ WKH F\DQRELRQWRIAzollafiliculoides /DP 3URWRSODVPD
'*$GDPV &DQLQL $ *ULOOL &DLROD 0 &LYLWDUHDOH 3 DQG *DOLD]]R )E 6XSHUR[LGH GLVPXWDVH LQ V\PELRWLF IUHHOLYLQJ DQG ZLOG Anabaena DQG Nostoc 1RVWRFDOHV &\DQRSK\WD 3K\FRORJLD &DQLQL $ *ULOOL &DLROD 0 DQG )HUUXVFL / D 4XDQWLWLDWLYH DQG TXDOLWDWLYH '1$ YDULDWLRQV LQ Anabaena azollae 6WUDVE OLYLQJLQAzollafilliculoides /DP &\WRPHWU\ &DQLQL $ *ULOOL &DLROD 0 DQG 0DVFLQL 0 $PPRQLXP FRQWHQWQLWURJHQDVHDFWLYLW\DQGKHWHURF\VWIUHTXHQF\ZLWKLQ WKH OHDI FDYLWLHV RI Azolla filiculoides /DP )(06 0LFURELRO /HWW &DUUDSLoR ) $UH EDFWHULD WKH WKLUG SDUWQHU RI WKH AzollaAnabaena V\PELRVLV"3ODQW6RLO &DUUDSLoR ) DQG 7DYDUHV 5 1HZ GDWD RQ WKH AzollaAnabaena V\PELRVLV , 0RUSKRORJLFDO DQG KLVWRFKHPLFDO DVSHFWV ,Q 6NLQQHU )$ %RGGH\ 50 DQG )HQGULN , HGV 1LWURJHQ )L[DWLRQ :LWK 1RQ/HJXPHV SS .OXZHU $FDGHPLF3XEOLVKHUV'RUGUHFKW &DVVPDQ.*'H'DWWD6.$PDUDQWH67/LERRQ636DPVRQ0, DQG 'L]RQ 0$ /RQJWHUPFRPSDULVRQ RIWKH DJURQRPLF HIILFLHQF\ DQG UHVLGXDO EHQHILWV RI RUJDQLF DQG LQRUJDQLF QLWURJHQ VRXUFHV IRU WURSLFDO ORZODQG ULFH ([SO $JULF &DXGDOHV 5 0RUHDX 5$ :HOOV -0 DQG $QWRLQH $' &HOOXODU OLSLG DQG IDWW\ DFLG FRPSRVLWLRQ RI F\DQRELRQWV IURP Azolla caroliniana. 6\PELRVLV &DXGDOHV 5:HOOV-0 DQG $QWRLQH$' &HOOXODUIDWW\DFLG FRPSRVLWLRQ RI V\PELRWLF F\DQREDFWHULD LVRODWHG IURP WKH DTXDWLFIHUQAzolla. -*HQ0LFURELRO &DXGDOHV 5 :HOOV -0 $QWRLQH $' DQG %XWWHUILHOG -( )DWW\ DFLG FRPSRVLWLRQ RI V\PELRWLF F\DQREDFWHULD IURP GLIIHUHQWKRVWSODQW(Azolla) VSHFLHV(YLGHQFHIRUFRHYROXWLRQRI KRVWDQGV\PELRQW ,QW-6\VWHP%DFWHULRO &RKHQ 0) DQG 0HHNV -& $ KRUPRJRQLXP UHJXODWLQJ ORFXV hrmUA, RIWKH F\DQREDFWHULXPNostoc punctiforrne VWUDLQ $7&& DQGLWVUHVSRQVHWR DQH[WUDFWRIDV\PELRWLFSODQW SDUWQHU Anthoceros punctatus. 0RO 3ODQW 0LFUREH,QWHUDFWLRQV &RKHQ 0) :DOOLV -* &DPSEHOO (/ DQG 0HHNV -& 7UDQVSRVRQ PXWDJHQHVLV RINostoc VS VWUDLQ $7&& D ILODPHQWRXV F\DQREDFWHULXP ZLWK PXOWLSOH FHOOXODU GLIIHUHQWLDWLRQDOWHUQDWLYHV 0LFURELRO &RUUHGRU-(:LONLQVRQ &5 9LFHQWH 93 0RUHOO -0 DQG2WHUR ( 1LWUDWH UHOHDVH E\ &DULEEHDQ UHHI VSRQJHV /LPQRO 2FHDQRJU &RZDQ ,5 /DQJH 2/ DQG *UHHQ 7*$ &DUERQGLR[LGH H[FKDQJH LQ OLFKHQV GHWHUPLQDWLRQ RI WUDQVSRUW DQG FDUER[\ODWLRQFKDUDFWHULVWLFV3ODQWD &R[ * &RPSDULVRQ RIProchloron IURP GLIIHUHQW KRVWV , 6WUXFWXUDO DQG XOWUDVWUXFWXUDO FKDUDFWHULVWLFV 1HZ 3K\WRO 'DOWRQ '$ DQG &KDWILHOG -0 $ QHZ QLWURJHQIL[LQJ F\DQRSK\WHKHSDWLF DVVRFLDWLRQ Nostoc DQG Porella. $P - %RW 'HOZLFKH &) DQG 3DOPHU -' 7KH RULJLQ RI SODVWLGV DQG WKHLU VSUHDG YLD VHFRQGDU\ V\PELRVLV 3ODQW 6\VWHP (YRO 6 'H
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV 'LRQLVLR6HVH 0/ 6KLPDGD $ 0DUX\DPD 7 DQG 0L\DFKL 6 &DUERQLF DQK\GUDVH DFWLYLW\ RI Prochloron VS LVRODWHG IURPDQDVFLGLDQKRVW$UFK0LFURELRO 'RGGV :. 3KRWRV\QWKHVLV RIWZR PRUSKRORJLHV RINostoc parmelioides &\DQREDFWHULD DVUHODWHGWRFXUUHQWYHORFLWLHV DQG GLIIXVLRQSDWWHUQV-3K\FRO 'RGGV:.DQG0DUUD- %HKDYLRUVRIWKHPLGJHCricotopus 'LSWHUD&KLURQRPLGDH UHODWHG WR PXWXDOLVP ZLWK Nostoc parmelioides &\DQREDFWHULD $TXDW ,QVHFWV 'RGGV:.*XGGHU'$DQG0ROOHQKDXHU' 7KHHFRORJ\RI Nostoc. -3K\FRO 'RXJODV 6( (XNDU\RWHHXNDU\RWH HQGRV\PELRVHV LQVLJKWV IURPVWXGLHVRIDFU\SWRPRQDGDOJD%LR6\VWHPV 'RXJODV 6( &KORURSODVW RULJLQV DQG HYROXWLRQ ,Q %U\DQW '$ HG 7KH 0ROHFXODU %LRORJ\ RI &\DQREDFWHULD SS .OXZHU$FDGHPLF3XEOLVKHUV'RUGUHFKW 'XQKDP '* DQG )RZOHU . 7D[RQRP\ DQG VSHFLHV UHFRJQLWLRQ LQ Azolla /DP ,Q Azolla 8WLOL]DWLRQ SS ,QWHUQDWLRQDO5LFH5HVHDUFK,QVWLWXWH/RV%DxRV3KLOLSSLQHV 'XRQJ 73 DQG 7LHGMH -0 1LWURJHQ IL[DWLRQ E\ QDWXUDOO\ RFFXUULQJ GXFNZHHGF\DQREDFWHULDO DVVRFLDWLRQV &DQ - 0LFURELRO (VNHZ '/ &DHWDQR$QROOpV * %DVVDP %- DQG *UHVVKRII 30 '1$ DPSOLILFDWLRQ ILQJHUSULQWLQJ RI WKH AzollaAnabaena V\PELRVLV 3ODQW0RO%LRO (QGHUOLQ&6 DQG 0HHNV-& 3XUHFXOWXUHDQGUHFRQVWLWXWLRQ RI WKH Anthoceros-Nostoc V\PELRWLF DVVRFLDWLRQ 3ODQWD )DXONQHU-' 8QVRQ0'DQG%HZOH\&$ 7KHFKHPLVWU\RI VRPH VSRQJHV DQG WKHLU V\PELRQWV 3XUH $SSO &KHP )D\ 3 2[\JHQ UHODWLRQV RI QLWURJHQ IL[DWLRQ LQ F\DQREDFWHULD0LFURELRO5HY )HLJH *% DQG -HQVHQ 0 %DVLF FDUERQ DQG QLWURJHQ PHWDEROLVPRIOLFKHQV ,Q5HLVVHU:HG $OJDHDQG6\PELRVHV 3ODQWV $QLPDOV )XQJL 9LUXVHV ,QWHUDFWLRQV ([SORUHG SS %LRSUHVV/WG%ULVWRO )RPL&DQG*ULOOL&DLROD0 3K\VLRORJLFDOFKDUDFWHUL]DWLRQ RIWKHEDFWHULD LVRODWHG IURPWKH DTXDWLF IHUQ Azolla. ,Q %RWKH + GH %UXLMQ )- DQG 1HZWRQ :( HGV 1LWURJHQ )L[DWLRQ +XQGUHG
)XFKV % 6XWWQHU3 6WHUQHU 6 :DVWOKXEHU 5 DQG /RRV ( 'LVSURSRUWLRQDWLQJ WUDQVJO\FRV\ODVH 'HQ]\PH LQ JUHHQ DOJDH DQG F\DQREDFWHULD SDUWLDO SXULILFDWLRQ DQG FKDUDFWHUL]DWLRQ Z 1DWXUIRUVFKXQJ&$-RXUQDORI%LRVFLHQFHV *DOORQ -5 7DQVOH\ 5HYLHZ 1R 5HFRQFLOLQJ WKH LQFRPSDWLEOH1 IL[DWLRQDQG21HZ3K\WRO *DQWDU0.HUE\1:DQG5RZHOO3D &RORQL]DWLRQRIZKHDW (Triticum vulgare / E\ 1IL[LQJ F\DQREDFWHULD ,, $Q XOWUDVWUXFWXUDOVWXG\1HZ3K\WRO *DQWDU0 .HUE\1:DQG5RZHOO3 &RORQL]DWLRQRIZKHDW (Triticum vulgare / E\ 1IL[LQJ F\DQREDFWHULD ,,, 7KH UROH RIDKRUPRJRQLDSURPRWLQJIDFWRU1HZ3K\WRO *DQWDU 0 .HUE\ 1: 5RZHOO 3 DQG 2EUHKW Z E &RORQL]DWLRQ RI ZKHDW (Triticum vulgare / E\ 1IL[LQJ F\DQREDFWHULD,$VXUYH\RIVRLOF\DQREDFWHULDOLVRODWHVIRUPLQJ DVVRFLDWLRQVZLWKURRWV 1HZ3K\WRO *DQWDU 0 .HUE\ 1: 5RZHOO 3 2EUHKW = DQG 6FULPJHRXU & E &RORQL]DWLRQ RI ZKHDW (Triticum vulgare / E\ 1 IL[LQJ F\DQREDFWHULD ,9 'DUN QLWURJHQDVH DFWLYLW\ DQG HIIHFWV RIF\DQREDFWHULD RQ QDWXUDO 1 DEXQGDQFH LQ WKH SODQWV 1HZ 3K\WRO *DQWDU 0 5RZHOO3 .HUE\1:DQG6XWKHUODQG ,:D 5ROH RI H[WUDFHOOXODU SRO\VDFFKDULGH LQ WKH FRORQL]DWLRQ RI ZKHDW (Triticum vulgare / URRWV E\ 1IL[LQJ F\DQREDFWHULD %LRO )HUWLO6RLOV *HEKDUGW -6 DQG 1LHU]ZLFNL%DXHU 6$ ,GHQWLILFDWLRQ RI D FRPPRQ F\DQREDFWHULDO V\PELRQW DVVRFLDWHG ZLWK Azolla VSS WKURXJK PROHFXODU DQG PRUSKRORJLFDO FKDUDFWHUL]DWLRQ RI IUHH OLYLQJ DQG V\PELRWLF F\DQREDFWHULD $SSO (QYLURQ 0LFURELRO *HKULJ+ 6FKOHU$DQG .OXJH 0 Geosiphon pyriforme, DIXQJXVIRUPLQJHQGRF\WRELRVLVZLWKNostoc F\DQREDFWHULD LV DQ DQFHVWUDO PHPEHU RI WKH *ORPDOHV HYLGHQFH E\ 668 ULERVRPDO51$DQDO\VLV-0RO(YRO *RUGRQ 1 $QJHO '/ 1HRUL $ .UHVV 1 DQG .LPRU % +HWHURWURSKLF GLQRIODJHOODWHV ZLWK V\PELRWLF F\DQREDFWHULD DQG QLWURJHQ OLPLWDWLRQ LQ WKH *XOI RI $TDED 0DULQH (FRORJ\ 3URJUHVV6HULHV *ULIILWKV '- DQG7KLQK/9 3KRWRV\QWKHVLV E\ in situ DQG LVRODWHG Prochloron 3URFKORURSK\WD DVVRFLDWHG ZLWK WKH GLGHPQLGDVFLGLDQV6\PELRVLV *ULIILWKV'-DQG7KLQK/9 &XUUHQWVWDWXV RIWKHWD[RQRP\ RIWKH JHQXV Prochloron. ,Q 6WDOH\-7 HG %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ YRO SS :LOOLDPV DQG :LONLQV%DOWLPRUH *ULOOL &DLROD 0 &DQLQL $ DQG 0RVFRQH ' 2[\JHQ FRQFHQWUDWLRQ QLWURJHQDVH DFWLYLW\ DQG KHWHURF\VW IUHTXHQF\ LQ WKH OHDI FDYLWLHV RI Azolla filiculoides /DP )(06 0LFURELRO /HWW *ULOOL&DLROD0 )RUQL&DQG&DVWDJQROD0 %DFWHULD LQWKH Azolla-Anabaena DVVRFLDWLRQ6\PELRVLV *ULOOL &DLROD 0 )RUQL & DQG &DVWDJQROD 0 Anabaena azollae DNLQHWHV LQ WKH VSRURFDUSV RIAzolla filiculoides /DP 6\PELRVLV *UREEHODDU1 6FRWW:( +DWWLQJK : DQG 0DUVKDOO - 7KH LGHQWLILFDWLRQ RI WKH FRUDOORLG URRW HQGRSK\WHV RI WKH VRXWKHUQ $IULFDQ F\FDGV DQG WKH DELOLW\ RIWKH LVRODWHV WR IL[ GLQLWURJHQ 6RXWK$IULFDQ-%RW *XVHY09DQG.RU]KHQHYVND\D7* $UWLILFLDODVVRFLDWLRQV ,Q5DL$1HG &5&+DQGERRNRI 6\PELRWLF&\DQREDFWHULDSS &5&3UHVV%RFD5DWRQ)ORULGD +lOOERP / %HUJPDQ % DQG 5DL $1 ,PPXQRJROG ORFDOL]DWLRQ RI JOXWDPLQH V\QWKHWDVH LQ WKH F\DQRELRQWV RI WKH
OLFKHQV Peltigera aphthosa DQG Peltigera canina. /LFKHQ 3K\VLRO%LRFKHP +DQGD\DQL'(GUDGD5$3URNVFK3:UD\9:LWWH/9DQ6RHVW 5:0 .XQ]PDQQ $ 6RHGDUVRQR )RXU QHZ KLRDFWLYH SRO\EURPLQDWHG GLSKHQ\O HWKHUV RI WKH VSRQJH Dysidea herbacea IURPZHVW6XPDWUD ,QGRQHVLD-1DWXUDO3URGXFWV +HOPFKHQ7$%KDWWDFKDU\D'DQG0HONRQLDQ0 $QDO\VHV RIULERVRPDO 51$ VHTXHQFHV IURP JODXFRF\VWRSK\WH F\DQHOOHV SURYLGH QHZ LQVLJKWV LQWR WKH HYROXWLRQDU\ UHODWLRQVKLSV RI SODVWLGV -0RO(YRO +LOO '- 7KH FRQWURO RI WKH FHOO F\FOH LQ PLFURELDO V\PELRQWV1HZ3K\WRO +LOO'- 7KHQDWXUHRIWKHV\PELRWLFUHODWLRQVKLSLQOLFKHQV (QGHDYRXU1HZ6HULHV +LQGH 5 3LURQHW ) DQG %RURZLW]ND 0$ ,VRODWLRQ RI Oscillatoria spongeliae, WKH ILODPHQWRXV F\DQREDFWHULDO V\PELRQWRIWKHPDULQHVSRQJHDysidea herbacea. 0DULQH%LRO +LURVH ( 0DUX\DPD 7 &KHQJ / DQG /HZLQ 5$ ,QWUDFHOOXODU V\PELRVLV RI D SKRWRV\QWKHWLF SURNDU\RWH Prochloron VS LQ D FRORQLDO DVFLGLDQ ,QYHUWHEUDWH %LRO +ROWRQ 5: 6WDP :7 DQG %RHOH%RV 6$ '1$'1$ UHDVVRFLDWLRQ VWXGLHV ZLWK '1$ IURP Prochloron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ostoc VS LQ V\PELRVLV ZLWK Gunnera manicata E\(6,DQG((/6 %RW$FWD -DQVRQ6&DUSHQWHU(- DQG%HUJPDQ% D ,PPXQRODEHOOLQJ RI SK\FRHU\WKULQ ULEXORVH ELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH DQG QLWURJHQDVH LQ WKH XQLFHOOXODU F\DQRELRQWV RI Ornithocercus VSS 'LQRSK\FHDH 3K\FRORJLD -DQVRQ 6 5DL $1 DQG %HUJPDQ % 7KH PDULQH OLFKHQ Lichina confinis 2 ) 0XOO F $J XOWUDVWUXFWXUH DQG ORFDOL]DWLRQ RIQLWURJHQDVH JOXWDPLQH V\QWKHWDVH SK\FRHU\WKULQ DQG ULEXORVH ELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH LQ WKH F\DQRELRQW1HZ3K\WRO -DQVRQ65DL$1DQG%HUJPDQ%E ,QWUDFHOOXODUF\DQRELRQW Richelia intracellularis 8OWUDVWUXFWXUH DQG LPPXQR ORFDOL]DWLRQ RI SK\FRHU\WKULQ QLWURJHQDVH 5XELVFR DQG JOXWDPLQHV\QWKHWDVH0DULQH%LRO -RKDQVVRQ & (VWDEOLVKPHQW RI WKH Gunnera-Nostoc V\PELRVLV3K'WKHVLV6WRFNKROP8QLYHUVLW\6ZHGHQ -RKDQVVRQ & DQG %HUJPDQ % (DUO\ HYHQWV GXULQJ WKH HVWDEOLVKPHQW RI WKH Gunnera/Nostoc V\PELRVLV 3ODQWD -RKDQVVRQ & DQG %HUJPDQ % 5HFRQVWUXFWLRQ RI WKH V\PELRVLV RI Gunnera manicata /LQGHQ F\DQREDFWHULDO VSHFLILFLW\1HZ3K\WRO
'*$GDPV -RVHSK &0 DQG 0HHNV -& 5HJXODWLRQ RI H[SUHVVLRQ RI JOXWDPLQH V\QWKHWDVH LQ D V\PELRWLF Nostoc VWUDLQ DVVRFLDWHG ZLWK Anthocerospunctatus. -%DFWHULRO .DSODQ'&DOYHUW+(DQG3HWHUV*$ 7KHAzolla-Anabaena azollae UHODWLRQVKLS ;,, 1LWURJHQDVH DFWLYLW\ DQG SK\FRELOLSURWHLQV RIWKHHQGRSK\WHDV DIXQFWLRQ RIOHDIDJH DQG FHOOW\SH3ODQW3K\VLRO .DSODQ 'DQG3HWHUV *$ ,QWHUDFWLRQ RIFDUERQ PHWDEROLVP LQ WKH Ada-Anabaena V\PELRVLV 6\PELRVLV .DUGLVK16LOYHUVWHLQ /)OHPLQJHU*DQG*DOXQ0 /HFWLQ IURP WKH OLFKHQ Nephroma leavigatum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zolla V\PELRVLV E\ '1$DPSOLILFDWLRQ:RUOG-0LFURELRO%LRWHFKQRO .LPXUD- DQG 1DNDQR7 5HFRQVWLWXWLRQ RID Blasia-Nostoc V\PELRWLF DVVRFLDWLRQ XQGHU D[HQLF FRQGLWLRQV 1RYD +HGZLJLD .OHLQ ( %DU ( )RPL & 0DONLQ 6 DQG 7HO2U ( 7KH DSSOLFDWLRQ RI FU\R6(0 WHFKQLTXHV WR WKH VWXG\ RI WKH V\PELRWLF DVVRFLDWLRQ LQ WKH Azolla OHDI FDYLW\ - 0LFURVFRS\ .OXJH 0 0ROOHQKDXHU ' DQG 0ROOHQKDXHU 5 3KRWRV\QWKHWLF FDUERQ DVVLPLODWLRQ LQ Geosiphon pyriforme .XW]LQJ ) Y :HWWVWHLQ DQ HQGRV\PELRWLF DVVRFLDWLRQ RI IXQJXVDQGF\DQREDFWHULXP3ODQWD .OXJH 0 0ROOHQKDXHU ' DQG 0ROOHQKDXHU 5 Geosiphon pyrifone .W]LQJ YRQ :HWWVWHLQ D SURPLVLQJ V\VWHP IRU VWXG\LQJHQGRF\DQRVHV3URJ%RW .OXJH 0 0ROOHQKDXHU ' 0ROOHQKDXHU 5 DQG .DSH 5 Geosiphon pyrifone, DQ HQGRV\PELRWLF FRQVRUWLXP RI D IXQJXV DQG D F\DQREDFWHULXP (Nostoc), IL[HV QLWURJHQ %RW $FWD .QLJKW &' DQG $GDPV '* $ PHWKRG IRU VWXG\LQJ FKHPRWD[LV LQ QLWURJHQIL[LQJ F\DQREDFWHULXPSODQW V\PELRVHV 3K\VLRO0ROHF3ODQW3DWKRO .RLNH ,
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ostoc E\UHFRPELQDQW'1$ 6\PELRVLV /HRQDUGL ' &DQLQL $ DQG )RPL & ,PPXQRORJLFDO FRPSDULVRQEHWZHHQ$UIKUREDFIHULVRODWHV DQGEDFWHULDOLYLQJLQ $]ROODILOLFXORLGHV/DP 6\PELRVLV /HZLQ 5$ 2UGHU3URFKORUDOHV ,Q 6WDOH\-7HG %HUJH\ V 0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ YRO SS :LOOLDPVDQG:LONLQV%DOWLPRUH /HZLQ 5$ 7KH FHOO VDS RI 3URFKORURQ 3URFKORURSK\WD 3K\FRORJLD /HZLQ 5$ DQG &KHQJ / HGV 3URFKORURQ D 0LFURELDO (QLJPD&KDSPDQDQG+DOO1HZ
&\DQREDFWHULD SS .OXZHU $FDGHPLF 3XEOLVKHUV 1HWKHUODQGV /RWWL) *LRYDQQHWWL/ 0DUJKHUL0&9HQWXUD6 DQG0DWHUDVVL 5 'LYHUVLW\ RI '1$ PHWK\ODWLRQ SDWWHUQ DQG WRWDO '1$ UHVWULFWLRQ SDWWHP LQ V\PELRWLF 1RVWRF :RUOG - 0LFURELRO DQG %LRWHFKQRO /XFDV ,$1 6\PELRQWV RI WKH WURSLFDO 'LQRSK\VLDOHV 'LQRSK\FHDH 2SKHOLD 0DGDQ $3 DQG 1LHU]ZLFNL%DXHU 6$ ,Q VLWX GHWHFWLRQ RI WUDQVFULSWV IRU ULEXORVH ELVSKRVSKDWH FDUER[\ODVH LQ F\DQREDFWHULDOKHWHURF\VWV-%DFWHULRO 0i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azollae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
0ROOHQKDXHU ' DQG 0ROOHQKDXHU5 1RVWRF LQ V\PELRVLV 7D[RQRPLFLPSOLFDWLRQV$OJRORJLFDO6WXGLHV 0ROOHQKDXHU ' 0ROOHQKDXHU 5 DQG .OXJH 0 6WXGLHV RQ LQLWLDWLRQ DQG GHYHORSPHQW RI WKH SDUWQHU DVVRFLDWLRQ LQ Geosiphon pyriforme .XW] Y :HWWVWHLQ D XQLTXH HQGRF\WRELRWLF V\VWHP RI D IXQJXV *ORPDOHV DQG WKH F\DQREDFWHULXP Nostoc punctiforme .W] +DULRW 3URWRSODVPD 0XU/5DQG%XUJHU:LHUVPD7 7KHRUGHU3URFKORUDOHV ,Q %DORZV$7ULLSHU+*'ZRUNLQ0+DUGHU:DQG6FKOHLIHU.+ HGV 7KH 3URNDU\RWHV 9RO ,, SS 6SULQJHU9HUODJ +HLGHOEHUJ 0XWXVNLQ $$ DQG .ROHVQLNRY 3$ 6\PELRWLF AzollaAnabaena DVVRFLDWLRQ VRPH SKRWRV\QWKHWLF FKDUDFWHULVWLFV QLWURJHQDVH DFWLYLW\ DQG R[LGDWLYH V\VWHPV 3KRWRV\QWKHWLFD 1LHU]ZLFNL%DXHU 6$ Azolla-Anabaena V\PELRVLV 8VH LQ DJULFXOWXUH ,Q 5DL $1 HG &5& +DQGERRN RI 6\PELRWLF &\DQREDFWHULDSS &5&3UHVV%RFD5DWRQ)ORULGD 1LHU]ZLFNL%DXHU 6$ DQG $XOILQJHU + 8OWUDVWUXFWXUDO FKDUDFWHUL]DWLRQ RI HXEDFWHULD UHVLGLQJ ZLWKLQ OHDI FDYLWLHV RI V\PELRWLF DQG F\DQRELRQWIUHH Azolla mexicana. &XUUHQW 0LFURELRO 1LHU]ZLFNL%DXHU 6$ DQG $XOILQJHU + 2FFXUUHQFH DQG XOWUDVWUXFWXUDO FKDUDFWHUL]DWLRQ RI EDFWHULD LQ DVVRFLDWLRQ ZLWK DQG LVRODWHG IURP Azolla caroliniana. $SSO (QYLURQ 0LFURELRO 1LHU]ZLFNL%DXHU 6$ DQG +DVHONRUQ 5 'LIIHUHQFHV LQ P51$ OHYHOV LQ Anabaena OLYLQJ IUHHO\ RU LQ V\PELRWLF DVVRFLDWLRQZLWKAzolla. (0%2- 2EUHKW = .HUE\ 1: *DQWDU0 DQG 5RZHOO 3 (IIHFWV RI URRWDVVRFLDWHG 1IL[LQJ F\DQREDFWHULD RQ WKH JURZWK DQG QLWURJHQ FRQWHQW RIZKHDW (Triticum vulgare / VHHGOLQJV %LRO )HUWLO6RLOV 2GLQWVRY 96 1LWURJHQ IL[DWLRQ LQ Prochloron 3URFKORURSK\WD DVFLGLDQ DVVRFLDWLRQV LV Prochloron UHVSRQVLEOH(QGRF\WRELRVLVDQG&HOO5HVHDUFK 2OVRQ 55 /LJKWHQKDQFHG JURZWK RI WKH DVFLGLDQ Didemenum molle/Prochloron VS V\PELRVLV 0DULQH %LRO %HUOLQ 2VERUQH %$ &RPSDULVRQV RI SKRWRV\QWKHVLV DQG SURGXFWLYLW\ RI Gunnera tinctoria 0ROLQD 0LUEHO ZLWK DQG ZLWKRXW WKH SK\FRELRQW 1RVWRF punctiform / 3ODQW &HOO DQG (QYLURQ 2VERUQH%$&XOOHQ$-RQHV3:DQG&DPSEHOO*- 8VHRI QLWURJHQ E\ WKH Nostoc-Gunnera tinctoria 0ROLQD 0LUEHO V\PELRVLV1HZ3K\WRO 2VERUQH %$ 'RULV ) &XOOHQ $ 0F'RQDOG 5 &DPSEHOO * DQG 6WHHU0: Gunnera tinctoria: DQXQXVXDOQLWURJHQIL[LQJ LQYDGHU %LRVFLHQFH 2WW 6 3KRWRV\PELRGHPHV DQG WKHLU GHYHORSPHQW LQ Peltigera venosa. /LFKHQRORJLVW 3DHUO+: 1 IL[DWLRQ QLWURJHQDVHDFWLYLW\ DWWULEXWHG WR D VSHFLILF Prochloron 3URFKORURSK\WD DVFLGLDQ DVVRFLDWLRQ RQ 3DODDX0LFURQHVLD 0DULQH%LRO%HUOLQ 3DOHQLN % DQG 6ZLIW + &\DQREDFWHULDO HYROXWLRQ DQG SURFKORURSK\WH GLYHUVLW\ DV VHHQ LQ '1$GHSHQGHQW 51$ SRO\PHUDVHJHQHVHTXHQFHV-3K\FRO 3DOPTYLVW . 3KRWRV\QWKHWLF &2XVH HIILFLHQF\ LQ OLFKHQV DQG WKHLU LVRODWHG SKRWRELRQWV WKH SRVVLEOH UROH RI D &2 FRQFHQWUDWLQJPHFKDQLVP3ODQWD 3DOPTYLVW . 8SWDNH DQG IL[DWLRQ RI &2 LQ OLFKHQ SKRWRELRQWV6\PELRVLV
'*$GDPV 3DOPTYLVW . 0i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eltigera aphthosa SKRWRV\PELRGHPH1HZ3K\WRO 3HQQHOO 5, &HOO VXUIDFH DUDELQRJDODFWDQ SURWHLQV DUDELQRJDODFWDQV DQG SODQW GHYHORSPHQW ,Q &ROORZ -$ DQG *UHHQ -5 HGV 3HUVSHFWLYHV LQ 3ODQW &HOO 5HFRJQLWLRQ SS &DPEULGJH8QLYHUVLW\3UHVV&DPEULGJH 3HUNLQV 6. DQG 3HWHUV *$ 7KH Azolla-Anabaena V\PELRVLV HQGRSK\WH FRQWLQXLW\ LQ WKH Azolla OLIHF\FOH LV IDFLOLWDWHG E\ HSLGHUPDO WULFKRPHV , 3DUWLWLRQLQJ RI WKH HQGRSK\WLF Anabaena LQWR GHYHORSLQJ VSRURFDUSV 1HZ 3K\WRO 3HUUDMX %799 5DL $1 .XPDU $3DQG 6LQJK +1 Cycas circinalis-Anabaena cycadeae V\PELRVLV SKRWRV\QWKHVLV DQG WKHHQ]\PHV RIQLWURJHQ DQG K\GURJHQ PHWDEROLVP LQ V\PELRWLF DQGFXOWXUHGAnabaena cycadeue. 6\PELRVLV 3HWHUV*$ Azolla DQGRWKHUSODQWF\DQREDFWHULDV\PELRVHV $VSHFWVRIIRUPDQGIXQFWLRQ3ODQW6RLO 3HWHUV *$DQG0HHNV-& 7KHAzolla-Anabaena V\PELRVLV EDVLFELRORJ\ $QQX5HY3ODQW3K\VLRO 3HWHUV *$ .DSODQ ' 0HHNV -& %X]E\ .0 0DUVK %+ DQG &RUELQ-/ $VSHFWVRIQLWURJHQDQGFDUERQ LQWHUFKDQJH LQ WKH Azolla-Anabaena V\PELRVLV ,Q /XGGHQ 3: DQG %XPV -( HGV 1LWURJHQ )L[DWLRQ DQG &2 0HWDEROLVP SS (OVHYLHU1HZ
&KDSWHU 6\PELRWLF,QWHUDFWLRQV WD[RQRPLF GLVWULEXWLRQ DQG H[LVWHQFH RI UHJLRQV RI KRPRORJ\ ZLWK V\PELRWLF JHQHV RI Rhizobium. &DQ - 0LFURELRO 3OD]LQVNL - )UDQFKH & /LX && /LQ 7 6KDZ : *XQQLQJ %(6 DQG 5ROIH %* 7D[RQRPLF VWDWXV RIAnabaena azollae: $QRYHUYLHZ3ODQW6RLO 3OD]LQVNL - 7D\ORU5 6KDZ : &URIW / 5ROIH %* DQG *XQQLQJ %(6 E ,VRODWLRQ RI Agrobacterium VS VWUDLQ IURP WKH Azolla OHDIFDYLW\)(060LFURELRO/HWW 3OD]LQVNL - =KHQJ 4 7D\ORU5 &URIW/ 5ROIH %*DQG *XQQLQJ %(6 D '1$ SUREHV VKRZ JHQHWLF YDULDWLRQ LQ F\DQREDFWHULDO V\PELRQWV RI WKH Azolla IHUQ DQG D FORVHU UHODWLRQVKLS WR IUHHOLYLQJ Nostoc VWUDLQV WKDQ WR IUHHOLYLQJ Anabaena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nthoceros-Nostoc V\PELRVLV LQPXQRHOHFWURQPLFURVFRSLF ORFDOL]DWLRQ RIQLWURJHQDVH JOXWDPLQH V\QWKHWDVH SK\FRHU\WKULQ DQG ULEXORVHELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH LQ WKH F\DQRELRQWDQG WKHFXOWXUHG IUHHOLYLQJ LVRODWH Nostoc - *HQ0LFURELRO 5DL $1 %RUWKDNXU 0 6RGHUEDFN ( DQG %HUJPDQ % ,PPXQRJROG ORFDOL]DWLRQ RIK\GURJHQDVH LQ WKH F\DQREDFWHULDO SODQW V\PELRVHV Peltigera canina, Anthoceros punctatus DQG Gunnera megallanica. 6\PELRVLV 5DL $1 5RZHOO 3 DQG 6WHZDUW:'3 1LWURJHQDVH DFWLYLW\ DQG GDUN &2 IL[DWLRQ LQ WKH OLFKHQ Peltigera aphthosa :LOOG 3ODQWD 5DVPXVVHQ 8 -RKDQVVRQ & DQG %HUJPDQ % (DUO\ FRPPXQLFDWLRQ LQ WKH Gunnera-Nostoc V\PELRVLV 3ODQW LQGXFHG FHOO GLIIHUHQWLDWLRQ DQG SURWHLQ V\QWKHVLV LQ WKH F\DQREDFWHULXP 0ROHFXODU 3ODQW0LFUREH ,QWHUDFWLRQV 5DVPXVVHQ8-RKDQVVRQ&5HQJOLQ$3HWHUVVRQ&DQG%HUJPDQ % $ PROHFXODU FKDUDFWHUL]DWLRQ RI WKH Gunnera-Nostoc V\PELRVLV FRPSDULVRQ ZLWK Rhizobium- DQG AgrobacteriumSODQWLQWHUDFWLRQV1HZ3K\WRO 5HQVWURP.HOOQHU( 5DL $1 DQG %HUJPDQ % &RUUHODWLRQ EHWZHHQQLWURJHQDVHDQGJOXWDPLQHV\QWKHWDVHH[SUHVVLRQ LQ WKH F\DQREDFWHULXP Anabaena cylindrica. 3K\VLRO3ODQW80: 5LSSND 5 DQG 6WDQLHU 5< 7KH HIIHFWV RIDQDHURELRVLV RQ QLWURJHQDVH V\QWKHVLV DQG KHWHURF\VW GHYHORSPHQW E\ 1RVWRFDFHDQF\DQREDFWHULD-*HQ0LFURELRO 5RELQV5-+DOO'26KL'-7XUQHU5-DQG5KRGHV0-& 0XFLODJHDFWVWRDGKHUHF\DQREDFWHULDDQGFXOWXUHGSODQWFHOOVWR ELRORJLFDODQGLQHUWVXUIDFHV)(060LFURELRO/HWW 5RJHU 3$ %LRORJLFDO 1 IL[DWLRQ DQG LWV PDQDJHPHQW LQ ZHWODQGULFHFXOWLYDWLRQ)HUWLOLVHU5HVHDUFK 5RJHUV *$ DQG 6WHZDUW :'3 7KH F\DQRSK\WHKHSDWLF V\PELRVLV , 0RUSKRORJ\ DQG SK\VLRORJ\ 1HZ 3K\WRO
5RWKHU-$$]L]$.DULP1+DQG:KLWWRQ%$ (FRORJ\RI GHHSZDWHU ULFHILHOGV LQ %DQJODGHVK 1LWURJHQ IL[DWLRQ E\ EOXHJUHHQDOJDOFRPPXQLWLHV+\GURELRORJLD 5R]HQ $ $UDG + 6FKRQIHOG 0 DQG 7HO2U ( )UXFWRVH VXSSRUWV JO\FRJHQ DFFXPXODWLRQ KHWHURF\VW GLIIHUHQWLDWLRQ 1 IL[DWLRQ DQG JURZWK RI WKH LVRODWHG F\DQRELRQW Anabaena azollae. $UFK0LFURELRO 5R]HQ $ 6FKRQIHOG 0 DQG 7HO2U ( )UXFWRVHHQKDQFHG GHYHORSPHQW DQG JURZWK RIWKH 1IL[LQJ F\DQRELRQWAnabaena azollae. =1DWXUIRUVFKF 5XHW]OHU . 0DQJURYH VSRQJH GLVHDVH LQGXFHG E\ F\DQREDFWHULDO V\PELRQWV )DLOXUH RI D SULPLWLYH LPPXQH V\VWHP" 'LV$TXDW2UJ5: 6FKUHLEHU 8 *DGHPDQQ 5 5DOSK 3- DQG /DUNXP $:' $VVHVVPHQW RI SKRWRV\QWKHWLF SHUIRUPDQFH RI Prochloron LQ Lissoclinum patella LQ KRVSLWH E\ FKORURSK\OO IOXRUHVFHQFH PHDVXUHPHQWV3ODQW&HOO3K\VLRO 6FKURHWHU % ,Q VLWX SKRWRV\QWKHWLF GLIIHUHQWLDWLRQ RI WKH JUHHQ DOJDO DQG WKH F\DQREDFWHULDO SKRWRELRQW LQ WKH FUXVWRVH OLFKHQ Placopsis contortuplicata. 2HFRORJLD 6FKURHWHU % *UHHQ 7*$ .DSSHQ / DQG 6HSSHOW 5' &DUERQ GLR[LGH H[FKDQJH DW VXE]HUR WHPSHUDWXUHV )LHOG PHDVXUHPHQWV RQ Umbilicaria aprina LQ $QWDUFWLFD &U\SWRJDP %RW 6FKOHU$%RQIDQWH3 6FKQHSI( 0ROOHQKDXHU' DQG .OXJH 0 &KDUDFWHUL]DWLRQ RI WKH Geosiphon pyriforme V\PELRVRPH E\ DIILQLW\ WHFKQLTXHV FRQIRFDO ODVHU VFDQQLQJ PLFURVFRS\&/60 DQGHOHFWURQPLFURVFRS\3URWRSODVPD 6FKLL%OHU$0H\HU7*HKULJ+DQG.OXJH0 9DQDWLRQVRI OHFWLQ ELQGLQJ VLWHV LQ H[WUDFHOOXODU JO\FRFRQMXJDWHV GXULQJ WKH OLIH F\FOH RI Nostoc punctiforme, D SRWHQWLDOO\ HQGRV\PELRWLF F\DQREDFWHULXP (XU -3K\FRO 6FKLL%OHU $ 0ROOHQKDXHU ' 6FKQHSI ( DQG .OXJH 0 Geosiphon pyriforme, DQ HQGRV\PELRWLF DVVRFLDWLRQ RI IXQJXV DQG F\DQREDFWHULD WKH VSRUH VWUXFWXUH UHVHPEOHV WKDW RI DUEXVFXODUP\FRUUKL]DO$0 IXQJL%RW$FWD 6FKOHU$ 6FKQHSI( 0ROOHQKDXHU' DQG .OXJH 0 7KH IXQJDO EODGGHUV RI WKH HQGRF\DQRVLV Geosiphon pyriforme, D *ORPXVUHODWHGIXQJXVFHOOZDOOSHUPHDELOLW\LQGLFDWHVDOLPLWHG SRUHUDGLXVRIRQO\QP3URWRSODVPD 6KDUPD$ 0LVKUD'3 DQG .XPDU $ &KDUDFWHUL]DWLRQ RID V\PELRWLFKHWHURF\VWRXV1IL[LQJF\DQREDFWHULXPIURP Cycas FRUDOORLGURRWV:RUOG-0LFURELRO%LRWHFKQRO 6KHDWK 5* DQG &ROH .0 %LRJHRJUDSK\ RI VWUHDP PDFURDOJDHLQ1RUWK$PHULFD- 3K\FRO 6KL'-%URXHUV0+DOO'2DQG5RELQV5- 7KHHIIHFWVRI LPPRELOL]DWLRQ RQ WKH ELRFKHPLFDO SK\VLRORJLFDO DQG PRUSKRORJLFDO IHDWXUHV RI Anabaena azollae. 3ODQWD 6KLPDGD$ .DQDL 6 DQG 0DUX\DPD7 3DUWLDO VHTXHQFH RI ULEXORVHELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH DQG WKH SK\ORJHQ\RIProchloron DQG Prochlorococcus 3URFKORUDOHV - 0RO (YRO 6LPRQ5' ,QFOXVLRQERGLHVLQWKHF\DQREDFWHULD ,Q)D\3 DQG9DQ%DDOHQ&HGV 7KH&\DQREDFWHULDSS 6LOYHVWHU:%3DUVRQV5DQG:DWW3: 'LUHFWPHDVXUHPHQW RIUHOHDVH DQG DVVLPLODWLRQ RIDPPRQLD LQ WKH Gunnera-Nostoc V\PELRVLV1HZ3K\WRO 6RGHUEDFN ( 'HYHORSPHQWDO SDWWHUQV LQ WKH NostocGunnera V\PELRVLV3K'WKHVLV6WRFNKROP8QLYHUVLW\ 6ZHGHQ $NDGHPLWU\FN$%
6RGHUEDFN ( DQG %HUJPDQ % 7KH Nostoc-Gunnera magellanica V\PELRVLV SK\FRELOLSURWHLQV FDUER[\VRPHV DQG 5XELVFRLQWKHF\DQRELRQW3K\VLRO3ODQW 6RGHUEDFN ( DQG %HUJPDQ % 7KH Nostoc-Gunnera V\PELRVLV FDUERQ IL[DWLRQ DQG WUDQVORFDWLRQ 3K\VLRO 3ODQW 6RGHUEDFN( /LQGEODG 3 DQG %HUJPDQ% 'HYHORSPHQWDO SDWWHUQV UHODWHG WR QLWURJHQ IL[DWLRQ LQ WKH Nostoc-Gunnera magellanica /DPV\PELRVLV3ODQWD 6SLOOHU+ DQG *XQDVHNDUDQ 0 $PPRQLDH[FUHWLQJ PXWDQW RIWKH F\DQREDFWHULXP Anabaena variabilis VXSSRUWV JURZWK RI ZKHDW $SSO0LFURELRO %LRWHFKQRO 6SLOOHU + 6WDOOLQJV -U : :RRGV 7 DQG *XQDVHNDUDQ 0 5HTXLUHPHQW IRU GLUHFW DVVRFLDWLRQ RI DPPRQLDH[FUHWLQJ Anabaena variabilis PXWDQW 6$ ZLWK URRWV IRU PD[LPDO JURZWK DQG \LHOG RI ZKHDW $SSO 0LFURELRO %LRWHFKQRO 6WDFNHEUDQGW(6HHZDOGW()RZOHU9-DQG6FKOHLIHU.+ 7KH UHODWHGQHVV RI Prochloron VS LVRODWHG IURP GLIIHUHQW GLGHPQLGDVFLGLDQKRVWV$UFK0LFURELRO 6WDII,$ DQG $KHUQ &3 6\PELRVLV LQ F\FDGV ZLWK VSHFLDO UHIHUHQFH WR Macrozamia communis. ,Q 6WHYHQVRQ ': DQG 1RUVWRJ-.HGV 7KH%LRORJ\6WUXFWXUHDQG6\VWHPDWLFVRIWKH &\FDGDOHV 3URFHHGLQJVRI&<&$' WKHVHFRQGLQWHUQDWLRQDO FRQIHUHQFH RQ F\FDG ELRORJ\ SS 3DOP DQG &\FDG 6RFLHWLHVRI$XVWUDOLD/WG4XHHQVODQG$XVWUDOLD 6WDP :7 %RHOH%RV 6$ DQG 6WXOS %. *HQRW\SLF UHODWLRQVKLSV EHWZHHQ Prochloron VDPSOHV IURP GLIIHUHQW ORFDOLWLHVDQGKRVWVDV GHWHUPLQHGE\'1$'1$UHDVVRFLDWLRQV $UFK0LFURELRO 6WHLQEHUJ 1$ DQG 0HHNV -& 3KRWRV\QWKHWLF &2 IL[DWLRQ DQG ULEXORVH ELVSKRVSKDWH FDUER[\ODVHR[\JHQDVH DFWLYLW\ RI Nostoc VS VWUDLQ 8&' LQ V\PELRWLF DVVRFLDWLRQ ZLWK Anthoceros punctatus. -%DFWHULRO 6WHLQEHUJ 1$ DQG 0HHNV -& 3K\VLRORJLFDO VRXUFHV RI UHGXFWDQWIRUQLWURJHQIL[DWLRQDFWLYLW\LQNostoc VS VWUDLQ8&' LQ V\PELRWLF DVVRFLDWLRQ ZLWK Anthoceros punctatus. %DFWHULRO 6WHUJLDQRX .. DQG )RZOHU . &KURPRVRPH QXPEHUV DQG WD[RQRPLF LPSOLFDWLRQV LQ WKH IHUQ JHQXV Azolla $]ROODFHDH 3ODQW6\VW(YRO 6WHZDUW :'3 DQG 5RJHUV *$ 7KH F\DQRSK\WHKHSDWLF V\PELRVLV ,, 1LWURJHQ IL[DWLRQDQG WKHLQWHUFKDQJHRIQLWURJHQ DQGFDUERQ 1HZ3K\WRO 6WLUHZDOW 9/ 0LFKDORZVNL &% /RIIHOKDUGW : %RKQHUW +- DQG %U\DQW '$ 1XFOHRWLGH VHTXHQFH RIWKH F\DQHOOHJHQRPH IURP Cyanophora paradoxa. 3ODQW 0RO %LRO 5HSRUWHU 6WRFN 3$ DQG 6LOYHVWHU:% 3KORHP WUDQVSRUW RIUHFHQWO\ IL[HG QLWURJHQ LQ WKH Gunnera-Nostoc V\PELRVLV 1HZ 3K\WRO 6WRFNHU:RUJRWWHU ( ([SHULPHQWDO FXOWLYDWLRQ RI OLFKHQV DQGOLFKHQV\PELRQWV&DQ-%RWVXSSOHPHQW 66 6WRFNHU:RUJRWWHU( DQG 7XUN5 7KDOOXV IRUPDWLRQ of WKH F\DQREDFWHULDO OLFKHQ Peltigera didactyla IURP VRUHGLD XQGHU ODERUDWRU\FRQGLWLRQV%RW$FWD 6WRFNHU:RUJRWWHU ( DQG 7XUN 5 $UWLILFLDO UHV\QWKHVLV RI WKDOOL RI WKH F\DQREDFWHULDO OLFKHQ Peltigera praetextata XQGHU ODERUDWRU\FRQGLWLRQV/LFKHQRORJLVW 6WRFNHU:RUJRWWHU ( DQG 7XUN 5 $UWLILFLDO UHV\QWKHVLV RI WKH SKRWRV\PELRGHPH Pelfigera leucophlebia XQGHU ODERUDWRU\ FRQGLWLRQV&U\SWRJDP%RW 6WUDVEXUJHU ( 8EHU Azolla, SS +HUPDQ 'DELV 9HUODJ-HQD
' * $GDPV 6WUDVEXUJHU ( 'DV %RWDQLVFKH 3UDFWLFXP *XVWDY )LVFKHU -HQDDQG/HLS]LJ 6ZLIW + 7KH F\WRORJ\ RI Prochloron. ,Q /HZLQ 5$ DQG &KHQJ / HGV Prochloron, D 0LFURELDO (QLJPD SS &KDSPDQDQG+DOO1HZ
&KDSWHU 6\PELRWLF ,QWHUDFWLRQV VRPDWLF HPEU\RV RIFDUURW WUHDWHG ZLWK DKLJK PROHFXODUZHLJKW H[WUDFWIURPDPDULQHF\DQREDFWHULXP 3ODQW&HOO5HS :DNH + 8PHWVX + 2]HNL < 6KLPRPXUD . DQG 0DWVXQDJD 7 , ([WUDFWV RI PDULQH F\DQREDFWHULD VWLPXODWHG VRPDWLF HPEU\RJHQHVLVRIDaucus carota / 3ODQW&HOO5HS :DOODFH :+ DQG *DWHV -( ,GHQWLILFDWLRQ RI HXKDFWHULD LVRODWHG IURP OHDI FDYLWLWLHV RI IRXU VSHFLHV RI WKH 1IL[LQJ Azolla IHUQ DV Arthrobacter &RQQ DQG 'LPPLFN $SSO (QYLURQ 0LFURELDO :DUG $. 'DKP &1 DQG &XPPLQJV .: Nostoc &\DQRSK\WD SURGXFWLYLW\ LQ 2UHJRQ VWUHDP HFRV\VWHPV LQYHUWHEUDWH LQIOXHQFHV DQG GLIIHUHQFHV EHWZHHQ PRUSKRORJLFDO W\SHV -3K\FRO :DVPDQQ&& /|HIIHOKDUGW:DQG %RKQHUW +- &\DQHOOHV RUJDQL]DWLRQDQGPROHFXODUELRORJ\,Q)D\3DQG9DQ%DDOHQ& HGV 7KH&\DQREDFWHULDSS(OVHYLHU$PVWHUGDP :DWDQDEH,/LQ&5DPLUH]&/DSLV076DQWLDJR9HQWXUD7DQG /LX && 3K\VLRORJ\ DQG DJURQRP\ RIAzolla-Anabaena V\PELRVLV ,Q 6NLQQHU )$ %RGGH\ 50 DQG )HQGULN , HGV 1LWURJHQ )L[DWLRQ ZLWK 1RQ/HJXPHV SS .OXZHU $FDGHPLF3XEOLVKHUV'RUGUHFKW :DWDQDEH,DQG/LX && ,PSURYLQJQLWURJHQIL[LQJV\VWHPV DQG LQWHJUDWLQJ WKHP LQWR VXVWDLQDEOH ULFH IDQQLQJ 3ODQW 6RLO :HUQHU ' 6\PELRVLV RI 3ODQWV DQG 0LFUREHV SS &KDSPDQDQG+DOO/RQGRQ :HVW 1 DQG $GDPV '* 3KHQRW\SLF DQG JHQRW\SLF FRPSDULVRQ RI V\PELRWLF DQG IUHHOLYLQJ F\DQREDFWHULD IURP D VLQJOHILHOGVLWH $SSO(QYLURQ0LFURELRO :KDWOH\ )5 DQG $OEHUWH 56 %LRFKHPLFDO IHDWXUHV RI Prochloron. ,Q /HZLQ 5$ DQG &KHQJ / HGV Prochloron, D 0LFURELDO(QLJPDSS&KDSPDQDQG+DOO1HZ
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hitton and M . Potts (eds), The Ecology of Cyanobacteria, Kluwer Academic Publishers . Printed in the Netherlands
&$6XWWOH
,;
& 6LJQLILFDQFH &RQFOXVLRQ
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ynechococcus VSS DUH WKH PRVW DEXQGDQW F\DQRSKDJHV 6W\ORYLULGDH DQG 3RGRYLULGDH DUH PRVW FRPPRQO\ LVRODWHG IURP IUHVK ZDWHUV 0RUSKRORJLFDO HYLGHQFH DOVR VXJJHVWV WKDW IUHVKZDWHU DQG PDULQH P\RYLUXVHV DUH PRUHFORVHO\UHODWHG WRHDFKRWKHUWKDQWKH\DUHWR RWKHUEDFWHULRSKDJHV &\DQRSKDJHV WKDWLQIHFWSK\FRHU\WKULQ ULFK Synechococcus VSSFDQEHH[WUHPHO\DEXQGDQWLQFRDVWDOPDULQHHQYLURQPHQWVZKHUHWKH\FDQRFFXUDWWLWHUV LQH[FHVVRI P/ DQG J RIVHGLPHQW,QVXUIDFHZDWHUV DEXQGDQFHYDULHVRYHURUGHUV RIPDJQLWXGHRQD VHDVRQDOEDVLV$EXQGDQFHIROORZVWKDWRISynechococcus, ZLWKHYLGHQFHIRUDWKUHVKROGLQSynechococcus RIFD WR P/ EH\RQG ZKLFK F\DQRSKDJH DEXQGDQFH LQFUHDVHV JUHDWO\ ,Q QHDUVKRUH ZDWHUV WKH KLJK FRQFHQWUDWLRQV RI F\DQRSKDJHV DQG Synechococcus UHVXOW LQ KLJK HQFRXQWHU IUHTXHQFLHV DQG VHOHFWLRQ IRU Synechococcus FRPPXQLWLHV WKDW DUH ODUJHO\ UHVLVWDQW WR LQIHFWLRQ (QFRXQWHUV DUH PXFK OHVV IUHTXHQW RIIVKRUH DQGWKLVOHDGVWRWKHDSSHDUDQFHRIDFRPPXQLW\WKDWDSSHDUVWRKDYHORZUHVLVWDQFHWRLQIHFWLRQ ,QIUHVKZDWHUV YLUXVHV ZKLFK LQIHFW ILODPHQWRXV F\DQREDFWHULD DSSHDU WR EH PRVW DEXQGDQW DQG DOVR VKRZ VWURQJ VHDVRQDO G\QDPLFV KRZHYHU HYHQ LQ WKH PRVW HXWURSKLF HQYLURQPHQWV WLWHUV DUH RUGHUV RI PDJQLWXGH OHVV WKDQ LQ SURGXFWLYHFRDVWDOZDWHUV/LWWOHHIIRUWZDVPDGHWRVFUHHQIUHVKZDWHUVIRUF\DQRSKDJHVWKDWLQIHFWSK\FRHU\WKULQ ULFK Synechococcus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ynechococcus RQDGDLO\ EDVLV ,Q DGGLWLRQ WR O\WLF LQIHFWLRQ O\VRJHQLFDVVRFLDWLRQVZHUHFOHDUO\GHPRQVWUDWHGLQILODPHQWRXVDQGXQLFHOOXODUF\DQREDFWHULD EXWWKH HFRORJLFDO LPSOLFDWLRQV RI O\VRJHQ\ UHPDLQ XQH[SORUHG (QYLURQPHQWDO IDFWRUV DQG WKH SK\VLRORJLFDO VWDWH RI F\DQREDFWHULDFOHDUO\DIIHFWF\DQRSKDJHF\DQREDFWHULDOLQWHUDFWLRQVEXWUHPDLQSRRUO\XQGHUVWRRG
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
EORRPV :KHQ LW EHFDPH DSSDUHQW WKDW F\DQRSKDJHV ZHUHXQOLNHO\WREHXVHIXODVELRORJLFDOFRQWURODJHQWV HFRORJLFDOLQYHVWLJDWLRQVODUJHO\FHDVHG ,QWKHODVWIHZ\HDUVWKHUHZDVDUHVXUJHQFHRIZRUN IRFXVHG RQ WKH HFRORJ\ RI F\DQRSKDJHV SDUWLFXODUO\ ZLWK UHVSHFW WR WKRVH ZKLFK LQIHFW PDULQH Synechococcus VSS &DUU DQG 0DQQ 7KHVH HIIRUWV VWHPPHG IURP REVHUYDWLRQV WKDW YLUXVOLNH SDUWLFOHV ZHUH W\SLFDOO\ SUHVHQW LQ QDWXUDO ZDWHUV DW DEXQGDQFHV LQ H[FHVV RI P/ %HUJK HW DO D VLJQLILFDQW SURSRUWLRQ RI F\DQREDFWHULD LQ PDULQH SK\WRSODQNWRQ FRPPXQLWLHV DSSHDUHG WR FRQWDLQ YLVLEOH YLUXV SDUWLFOHV 3URFWRU DQG )XKUPDQ DQG YLUXVHV WKDW LQIHFW F\DQREDFWHULD ZHUH UHDGLO\ LVRODWHG IURP PDULQH ZDWHUV 6XWWOH HW DO 6XEVHTXHQWO\ LW ZDV GLVFRYHUHG WKDW LQIHFWLRXVF\DQRSKDJHV FDQEHH[WUHPHO\ DEXQGDQWLQ
&KDSWHU &\DQRSKDJHV VHDZDWHU ZLWK FRQFHQWUDWLRQV IUHTXHQWO\ LQ H[FHVV RI WR P/ 6XWWOHDQG&KDQ 6XWWOHHWDO :DWHUEXU\DQG9DORLV *LYHQWKHJOREDO VLJQLILFDQFH RI F\DQREDFWHULD DV SULPDU\ SURGXFHUV DQG QLWURJHQ IL[HUV DV ZHOO DV WKH SRWHQWLDO RI F\DQRSKDJHV WR VHUYH DV PRUWDOLW\ DJHQWV YHFWRUV RI JHQHWLF LQIRUPDWLRQ DQG UHJXODWRUV RI FRPPXQLW\ VWUXFWXUH LW LV WLPHO\ WR FRQVLGHU ZKDW WUDQVSLUHG LQ WKH GHFDGH VLQFH WKH ODVWUHYLHZ RIF\DQRSKDJHV ZDV SXEOLVKHG 8QOLNH SUHYLRXV UHYLHZV WKDW W\SLFDOO\ SURYLGHG H[WHQVLYH LQIRUPDWLRQ RQ F\DQRSKDJH ELRORJ\ WKH SULPDU\ HPSKDVLV RIWKLV FRQWULEXWLRQ LV WR IRFXV RQ DVSHFWV WKDW DUH RI HFRORJLFDO DQG HQYLURQPHQWDOLPSRUWDQFH
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nternational Committee on Taxonomy of Viruses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nacystis DQG Synechococcus 6DIIHUPDQ HW DO 0RUSKRORJLFDO FKDUDFWHULVWLFV LQFOXGH DQ LVRPHWULF KHDG GLDPHWHU QP DFRQWUDFWLOH WDLO RI [ QPWKDWFRQWUDFWVWR QP 5HSUHVHQWDWLYHV IURPWKLV JURXS DOVR LQIHFWILODPHQWRXV F\DQREDFWHULD DVVLJQHGWRWKHJHQXVAnabaena HJ*URPRY 7KHUH LV FRQVLGHUDEOH YDULDWLRQ LQ VL]H DQG PRUSKRORJ\DPRQJLVRODWHVZLWKLQWKLVJURXS ERWKLQ WKH VL]H RIWKH FDSVLG DQG LQ WKH VWUXFWXUH RIWKH WDLO *URPRY 6XWWOH DQG &KDQ :DWHUEXU\ DQG 9DORLV ZKLFK PD\ VXJJHVW WKDW WKH SURSRVHG JHQXV FRQVLVWV RID QXPEHU RIVSHFLHV )RU H[DPSOH :DWHUEXU\ DQG 9DORLV LVRODWHG IRXU GLVWLQFW PRUSKRW\SHV IURP D VLQJOH ZDWHU VDPSOH 7KLV YDULDWLRQ ZDV DOVR REVHUYHG DW WKH PROHFXODU OHYHO WKH QXPEHU RI FDSVLG SURWHLQV UHSRUWHG YDULHG IURP IRUWZRPDULQHLVRODWHV:LOVRQHWDO WRIRUWKHW\SHVSHFLHVWKHPRO*&FRQWHQWZDV GHWHUPLQHG WR YDU\ IURP FD WR DQG WKH JHQRPH PROHFXODUZHLJKWIURP [ WR [ GDOWRQV $GROSK DQG +DVHONRUQ 6DIIHUPDQ HW DO 6KHUPDQ DQG &RQQHOO\ 7KH ODWWHU FRUUHVSRQG WR JHQRPLF FRPSOH[LWLHV RIFD WR NE EDVHG RQ EDVH SDLUV SHU GDOWRQ IRU ODPEGD %DVHG RQ UHVWULFWLRQ PDSSLQJ %DQFURIW DQG 6PLWK WKH JHQRPH VL]H RI WZR FORVHO\ UHODWHG F\DQRSKDJHV $1 DQG $1 +X HW DO ZDV NE ZKLOH $1 DQG $/ .R]\DNRY ZHUHFDNE :LOVRQHWDO UHSRUWHGJHQRPH VL]HV IRUVHYHUDOPDULQH LVRODWHVWR UDQJH IURPFD WR NE 1RWH WKDW DW OHDVW RQH RIWKH F\DQRSKDJHV LGHQWLILHGDVDF\DQRVW\ORYLUXVLQ:LOVRQHWDO ZDV VXEVHTXHQWO\ UHFODVVLILHG DV D F\DQRP\RYLUXV :LOVRQ HW DO 7KH ,&79 FODVVLILHG SKDJHV ZLWKLQ WKLV JURXS DV XQDVVLJQHG VSHFLHV LQ WKH 0\RYLULGDH0XUSK\HWDO
&$6XWWOH
Fig. 1. 5HSUHVHQWDWLYHV RIYLUXVHV WKDW LQIHFW F\DQREDFWHULD 6FDOH EDU HTXDOV QP D &\DQRP\RYLUXV Myoviridae LVRODWH 63:0 &\DQRP\RYLUXVHVSRVVHVVDFRQWUDFWLOHWDLOWKDWLVVHSDUDWHGIURPWKHFDSVLGE\DQHFN 1RWHWKHSUHVHQFHRIQHFNKDLUVDUURZV DQXQXVXDO IHDWXUH WKDW LV IRXQG RQ VRPH F\DQRP\RYLUXV LVRODWHV IURP IUHVK DQG PDULQH ZDWHUV E &\DQRSRGRYLUXV Podoviridue LVRODWH 6%%3, &\DQRSRGRYLUXVHV DUH GLVWLQJXLVKHG E\ VKRUW QRQFRQWUDFWLOH WDLOV F &\DQRVW\ORYLUXV Siphoviridae LVRODWH 6%%6 &\DQRVW\ORYLUXVHV KDYHDORQJQRQFRQWUDFWLOHWDLOWKDWFDQDSSHDUIOH[LEOHLQPLFURJUDSKV
&\DQRSRGRYLUXV
&\DQRVW\ORYLUXV
&\DQRSRGRYLUXVHV )LJ E EHORQJ ZLWKLQ WKH 3RGRYLULGDHEXWQRJHQXVZDV DVVLJQHGE\WKH,&79 7KH\ DUH SUHVHQW LQPDULQH DQG IUHVK ZDWHUV DQG DUH FRPPRQO\ LVRODWHG IURP ZDVWH VWDELOL]DWLRQ SRQGV HJ +X HW DO 7KH LVRPHWULF FDSVLGV DUH W\SLFDOO\VLPLODULQGLDPHWHUWRF\DQRVW\ORYLUXVHVEXW WKH WDLOV DUH PXFK VKRUWHU 7KH W\SH VSHFLHV LV &\DQRSKDJH/33 WKDWLQIHFWV DOJDH DVVLJQHG WR WKH JHQHUD Lyngbya, Plectonema and Phormidium 6FKQHLGHU HW DO 6KHUPDQ DQG +DVHONRUQ 7KH LVRPHWULFKHDG LV QPLQ GLDPHWHUDQG WKHWDLOWRQPORQJWKHUHDUHDERXWVWUXFWXUDO SURWHLQV DQG WKH JHQRPH LV [ GDOWRQV FD NE 7KHUHSRUWVRIPRO *&FRQWHQWVUDQJHIURP WKHORZVLQ/33/XIWLJDQG+DVHONRUQ WR WKH PLG V LQ 60 0DF.HQ]LH DQG +DVHONRUQ D %DQFURIW DQG 6PLWK SHUIRUPHG UHVWULFWLRQ PDSSLQJ RI&\DQRSRGRYLUXV $/ ZKLFK LQIHFWV Anabaena 3&& .R]\DNRY DQG FRQFOXGHG WKDWWKH JHQRPH ZDV DSSUR[LPDWHO\ NE LQVL]H
9LUXVHV ZLWKLQ WKH SURSRVHG JHQXV Cyanostylovirus EHORQJ WR WKH IDPLO\ 6LSKRYLULGDH )LJ F 7KH W\SH VSHFLHV &\DQRSKDJH 6 LQIHFWV WKH JHQXV Synechococcus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nabaena DQG Nostoc; WKH $6 JURXS LQIHFWV F\DQREDFWHULD FODVVLILHG DV Anacystis nidulans DQG Synechococcus cedrorum; DQG WKH 60 JURXSLQIHFWVF\DQREDFWHULDLGHQWLILHGDV6 elongatus DQG Microcystis aeruginosa. 0DWWHUV DUH FRPSOLFDWHG IXUWKHU EHFDXVH WKH YDOLGLW\ RIPDQ\ RIWKHVH VSHFLHV KDV EHHQ FDOOHG LQWR TXHVWLRQ 6HFWLRQ ,,,% DQG &KDSWHU 0RUH UHFHQWO\ WKHUH ZDV DQ HIIRUW WR LQFOXGH LQIRUPDWLRQ DERXW WKH WD[RQRPLF DIILOLDWLRQ RI F\DQRSKDJHV LQ WKHLU VWUDLQ GHVLJQDWLRQV 6XWWOH DQG &KDQ :LOVRQ HW DO DQG LW LV UHFRPPHQGHG WKDW WKLV SURFHGXUH EH IROORZHG LQ WKH IXWXUH 7KH VXJJHVWHG QRPHQFODWXUH IRU QHZ F\DQRSKDJHLVRODWHVLV&\DQRSKDJH;[<<=DDZKHUH Xx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
%(YROXWLRQ $OWKRXJKWKHRULJLQRIF\DQRSKDJHVLV XQNQRZQ LWLV SRVVLEOH WR PDNH LQIHUHQFHV UHJDUGLQJ WKHLU HYROXWLRQDU\ KLVWRU\ 7KH YDVW DUUD\ RI JHQHWLF DUFKLWHFWXUHWKDWLVIRXQGLQYLUXVHVLQGLFDWHVWKDWWKH\ DUHSRO\SK\OHWLFDQGGRQRWVKDUHDFRPPRQDQFHVWRU 7KH PRVWFRPPRQ H[SODQDWLRQIRUWKHLU RULJLQLV WKDW WKH\ UHSUHVHQW HVFDSHG JHQHWLF PDWHULDO IURP RUJDQLVPV &\DQRSKDJHV DUH XQOLNHO\ WR KDYH DULVHQ
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
,,, 'LYHUVLW\ $ *HQHWLF'LYHUVLW\ 7KH PRUSKRORJ\ DQG PROHFXODU FKDUDFWHULVWLFV RIWKH SURSRVHG JHQHUD RIF\DQRSKDJHV VXJJHVW WKDW JHQHWLF GLYHUVLW\ PD\ EH KLJK DOWKRXJK WKHUH DUH IHZ GDWD ZKLFK VSHFLILFDOO\ H[DPLQH GLYHUVLW\ ZLWKLQ
F\DQRSKDJHV 0XUDGRY HW DO H[DPLQHG ILYH LVRODWHV RI YLUXVHV LQ WKH 137 JURXS ZKLFK LQIHFW F\DQREDFWHULDDVVLJQHG WR WKHJHQHUDPlectonema DQG Nostoc. %DVHG RQ WKHLU GHVFULSWLRQ LW DSSHDUV WKDW WKHVH LVRODWHV ZHUH Podoviridae. 7KH\ IRXQG D GLIIHUHQW SDWWHUQ LQ WKH QXPEHU RI UHVWULFWLRQ IUDJPHQWV REWDLQHG ZLWK (FR59 DQG 6DX$ UHVWULFWLRQ HQGRQXFOHDVHV LQGLFDWLQJ WKDW HDFK RI WKH LVRODWHVZDVJHQHWLFDOO\GLIIHUHQW:LOVRQHWDO DOVR H[DPLQHG UHVWULFWLRQ IUDJPHQW OHQJWK SRO\PRUSKLVPV 5)/3 LQ VHYHUDO F\DQRSKDJHV WKDW LQIHFW PDULQH Synechococcus VS VWUDLQ '& V\QRQ\P :+ $OWKRXJK RULJLQDOO\ FKDUDFWHUL]HG DV F\DQRP\RYLUXVHV 6%0 630 6:+0 DQG F\DQRVW\ORYLUXVHV 6%6 636 DW OHDVW RQH RI WKH VW\ORYLUXVHV ZDV UHDVVLJQHG WR WKH Myoviridae :LOVRQHWDO 5HJDUGOHVV RIWKHLU WD[RQRP\WKH YLUXVHV IHOO LQWRWKUHH JURXSV EDVHG RQ 5)/3 UHVWULFWLRQ IUDJPHQW OHQJWK SRO\PRUSKLVP DQDO\VHV ,Q DVLPLODUVWXG\ XQSXEOLVKHG &KDQ DQG 6XWWOHH[DPLQHGWKUHHF\DQRP\RYLUXVHV63:06 3:063:0 $OWKRXJKDOORIWKHVHYLUXVHVZHUH DOVR RULJLQDOO\ LVRODWHG XVLQJ Synechococcus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
% +RVW5DQJH /33*URXS 7KHUH LV FRQVLGHUDEOH FRQIXVLRQ LQ WKH OLWHUDWXUH UHJDUGLQJ WKH KRVW VSHFLILFLW\ RI YLUXVHV WKDW LQIHFW F\DQREDFWHULD $ JRRG GHDO RI WKH FRQIXVLRQ
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yngbya, Plectonema DQG Phormidium 6DIIHUPDQ DQG 0RUULV 3DGDQ DQG 6KLOR VXJJHVWHG WKDW WKH YHU\ VLPLODU PRO * & FRQWHQWV DQG EXR\DQW GHQVLWLHV J FP RIWKH '1$ LVRODWHG IURP VXVFHSWLEOH LVRODWHV RI WKHVH JHQHUD (GHOPDQ HW DO LQGLFDWHG WKDW WKHVH VWUDLQV ZHUH FORVHO\ UHODWHG ,W ZDV VXJJHVWHG EDVHG RQ WKH JHQHWLF DQG PRUSKRORJLFDO VLPLODULWLHV DPRQJ WKH /33 JURXS RI F\DQREDFWHULD WKDW WKH\ EH LQFOXGHG LQ D VLQJOH VSHFLHV Schizothrix calcicola 'URXHW 6LPLODUO\ 6WDP UHFRPPHQGHG WKH PHUJHU RI WKH JHQHUD ZKLOH 5LSSND DQG +HUGPDQ SODFHGF\DQREDFWHULDVXVFHSWLEOHWR LQIHFWLRQE\/33 3&& LQWR D FORVHO\UHODWHGJHQXV(Leptolyngbya, &OXVWHU ZLWKLQ WKH 2UGHU 2VFLOODWRULDOHV +RZHYHU UHFHQW ZRUN 7XUQHU LQGLFDWHGWKDWVRPHPHPEHUVDVVLJQHG WRWKLVJHQXVEHORQJHGLQRWKHUWD[RQRPLFJURXSVDQG WKDW Leptolyngbya ZDVSUREDEO\QRWDYDOLGJHQXVDQG VKRXOGEHIXUWKHUVXEGLYLGHG 7KHPRVWH[WHQVLYHVWXG\RQWKHKRVWUDQJHRI/33 ZDV FRQGXFWHG E\-RKQVRQ DQG 3RWWV 7KH\ H[DPLQHGWKHVHQVLWLYLWLHVRIVWUDLQVDQGYDULDQWVRI F\DQREDFWHULD WR O\VLV E\ WKH /33 DUFKDHW\SH DV ZHOODVVHYHUDO/33 VHURW\SHV/33VHURW\SHVDQG QHZ /33 LVRODWHV 7KH\ DOVR FRQFOXGHG WKDW WKH KRVW UDQJHRI/33F\DQRSKDJHV ZDVUHVWULFWHG WRPHPEHUV RI WKH Lyngbya, Plectonema DQG Phormidium
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³/33´ JHQHUD $OWKRXJK GLIIHUHQW JURXSV RI JHQHWLFDOO\ UHODWHG F\DQREDFWHULD DUH EHJLQQLQJ WR EH TXLWH ZHOO GHILQHG :LOPRWWH 7XUQHU WKHUH DUH QXPHURXV H[DPSOHV ZKHUH F\DQREDFWHULD ZHUH LQFRUUHFWO\ DVVLJQHG WR D JLYHQ JHQXV &RQVHTXHQWO\ WKH /33 JURXS RI F\DQREDFWHULD VKRXOG QRW EH FRQVLGHUHG D WD[RQRPLFDOO\ YDOLG JURXS DQG LW LV OLNHO\ WKDW WKH F\DQREDFWHULDZKLFKDUHLQIHFWHGE\WKH/33JURXSRI F\DQRSKDJHV ZLOO XOWLPDWHO\ EH SODFHG LQ D VLQJOH JHQXV
$$1 1DQG13*URXSV &\DQRSKDJHV DOVR LQIHFW DQG FDXVH O\VLV RI ILODPHQWRXV F\DQREDFWHULD DVVLJQHG WR Nostoc VSS DQG Anabaena VSS DQGLQFOXGHERWKSRGRYLUXVHV $ O $/ 1$1VHULHVH[FHSW$1 DQG$1 DQGP\RYLUXVHV1$$1 DQG$1$/ VHULHV H[FHSW $/ $GROSK DQG +DVHONRUQ .R]\DNRY HW DO .R]\DNRY +X HW DO *URPRY $OWKRXJKWKHYLUXVHVUHSUHVHQW WZR IDPLOLHV WKH\ SRVVHVV ZLWK IHZ H[FHSWLRQV UHPDUNDEO\VLPLODUKRVWUDQJHVDQGLQIHFWERWKNostoc VSS DQGAnabaena VSS +X HW DO ,Q IXUWKHU VWXGLHV ZLWK F\DQRSKDJH 1 5LSSND DQG +HUGPDQ FRQILUPHG O\VLV RI F\DQREDFWHULD RULJLQDOO\ DVVLJQHGWRAnabaena variabilis 3&& $7&& Anabaena VS 3&& $7&& DQG Nostoc muscorum DQG Anabaena VS 3&& $7&& 3&& $7&& DQG %DVHGRQDYDULHW\RIFULWHULDWKH\FRQFOXGHG WKDW DOO RI WKHVH F\DQREDFWHULDO LVRODWHV EHORQJHG ZLWKLQ D YHU\ FORVHO\ UHODWHG WD[RQRPLF JURXS RI Nostoc &KDSWHU &RQVHTXHQWO\ WKH RULJLQDO FRQFOXVLRQ WKDW WKHVH YLUXVHV SRVVHVVHG D EURDG KRVW UDQJHZDVDQ DUWLIDFWRIF\DQREDFWHULDOQRPHQFODWXUH ,W LV DOVR LPSRUWDQW WR QRWH WKDWKRVWUDQJH PXWDWLRQV DOVR RFFXU VSRQWDQHRXVO\ LQ WKLV JURXS RI YLUXVHV 6DUPD DQG .DXU KHQFH FDXWLRQ PXVW EH
H[HUFLVHG ZKHQ XVLQJ VHQVLWLYLW\ WR F\DQRSKDJH LQIHFWLRQDVDWD[RQRPLFFULWHULRQ /LWWOH FDQ EH VDLG DERXW WKH 13 JURXS RI F\DQRSKDJHV 137 VWUDLQV WKDW ZHUH UHSRUWHG WR LQIHFW D YDULHW\ RIF\DQREDFWHULDO LVRODWHV DVVLJQHG WR WKH JHQHUD Nostoc DQG Plectonema 0XUDGRY HW DO %DVHG RQ WKH DXWKRUV¶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
$6DQG60*URXSV 6LPLODU GLIILFXOWLHV H[LVW ZLWK KRVW UDQJH VWXGLHV FRQGXFWHG RQ F\DQRSKDJH LVRODWHV WKDW LQIHFW XQLFHOOXODU F\DQREDFWHULD 7KH P\RYLUXV $6 ZDV RULJLQDOO\ UHSRUWHG WR LQIHFW Anacystis nidulans 87(; 3&& DQG Synechococcus cedrorum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ynechococcus elongatus 87(; 3&& DQG Microcystis aeruginosa 15& 6DIIHUPDQ HW DO E ZKLOH WKH P\RYLUXV $6 ZDV GHVFULEHG DV LQIHFWLQJ M. aeruginosa 15& DV ZHOO DV Synechococcus cedrorum 87(; 6DIIHUPDQHW DO 6XEVHTXHQWO\ LW ZDV VKRZQ WKDW 60 LQIHFWHG D QXPEHU RI FORVH UHODWLYHV WR 3&&
&$6XWWOH
3&&3&&3&&3&& ZKLFK EHORQJ ZLWKLQ D FORVHO\ UHODWHG FOXVWHU RI Synechococcus VSHFLHV LQFOXGLQJ S. cedrorum 3&& :DWHUEXU\ DQG 5LSSND 5LSSND DQG +HUGPDQ
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
$ )UHVKZDWHU 7KH H[LVWHQFH RI F\DQRSKDJHV ZDV GHPRQVWUDWHG E\ WKH SXULILFDWLRQ RI D YLUXV ZKLFK FDXVHG WKH O\VLV RI VHYHUDO LVRODWHV RI ILODPHQWRXV F\DQREDFWHULD EHORQJLQJ WR D JURXS RIFORVHO\ UHODWHG F\DQREDFWHULD WKDW ZHUH FODVVLILHG LQWR WKUHH JHQHUD Lyngbya, Plectonema DQG Phormidium 6DIIHUPDQ DQG 0RUULV )XUWKHU LQYHVWLJDWLRQ GHPRQVWUDWHG WKDW /33 F\DQRSKDJHV ZHUH IRXQG LQ ZDVWH VWDELOL]DWLRQSRQGV LQ WKH 8QLWHG 6WDWHV 6DIIHUPDQ DQG 0RUULV 6DIIHUPDQ HW DO D 6DIIHUPDQ ,QGLD 6LQJK DQG 6LQJK DQG 6FRWODQG 'DIW HW DO DV ZHOO DV LQ ILVKSRQGV LQ ,VUDHO 3DGDQ DQG 6KLOR DQG ULFH ILHOGV LQ ,QGLD 6LQJK 'HVSLWHWKHLUZLGHVSUHDGRFFXUUHQFHWKHWLWHUVRI/33 F\DQRSKDJHV ZHUH DW PRVW D IHZ KXQGUHG LQIHFWLRXV
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nabaenaNostoc JURXS DUHDOVR ZLGHVSUHDG DQG ZHUH IRXQG LQ ODNHV UHVHUYRLUV DQG VHZDJH VHWWOLQJ SRQGV $GROSK DQG +DVHONRUQ 0HQG]KXO HW DO .R]\DNRY +XHWDO 0RLVDHWDO 3KOLSVHWDO 7KH\ DUHOHVVIUHTXHQWO\LVRODWHG WKDQWKH/33F\DQRSKDJHV KRZHYHU DQG DOVRDSSHDU WR EH PXFK OHVV DEXQGDQW W\SLFDOO\ WKH\ DUH RQO\ GHWHFWDEOH IROORZLQJ DPSOLILFDWLRQ LQ HQULFKPHQW FXOWXUH $ WKLUG JURXS RI F\DQRSKDJHV ZDV UHSRUWHG WR LQIHFW F\DQREDFWHULD DVVLJQHG WR WKH JHQHUD Nostoc DQG Plectonema 0XUDGRY HW DO 7KHVH YLUXVHV ZHUH UHSRUWHG WR EH GLVWULEXWHG ZLGHO\ LQ WKH IRUPHU 6RYLHW 8QLRQ ZKHUH WKH\ ZHUH LVRODWHG IURP ILVK IDUPV DQG ZDVWH VWDELOL]DWLRQ SRQGV +LJK WLWHUV RI WKHVH YLUXVHV ZHUH IRXQG GXULQJ VXPPHU ZLWK DEXQGDQFHV UDQJLQJ IURP WR LQIHFWLRXV XQLWVP/ 0XUDGRYHWDO 7KH UDQJH RI RFFXUUHQFH RI YLUXVHV WKDW LQIHFW WKH Anacystis-Synechococcus JURXS RI F\DQREDFWHULD LV DVZLGHVSUHDGDVWKDWRIRWKHUIUHVKZDWHUF\DQRSKDJHV 6DIIHUPDQHWDO 6KHUPDQDQG&RQQHOO\ *URPRY .LPDQG&KRL DOWKRXJKWLWHUV ZHUH QRW UHSRUWHG URXWLQHO\ ,QWHUHVWLQJO\ DOWKRXJK SK\FRHU\WKULQULFK Synechococcus VSS FDQ EH GRPLQDQW LQ PDQ\ IUHVKZDWHU V\VWHPV HJ &DURQ HW DO 6WRFNQHUDQG 6KRUWUHHG WKHUHZHUH
IHZ DWWHPSWV WR LVRODWH YLUXVHV ZKLFK LQIHFW WKHVH F\DQREDFWHULD 2Q VHYHUDO RFFDVLRQV QDWXUDO YLUXV FRPPXQLWLHV IURP/DNH &RQVWDQFH LQ *HUPDQ\ ZHUH VFUHHQHG DQG YLUXVHV ZKLFK LQIHFWHG VHYHUDO SK\FRHU\WKULQULFK Synechococcus VWUDLQV ZHUH LVRODWHG &KDQ DQG 6XWWOH XQSXEOLVKHG GDWD 0RUH HIIRUWVVKRXOGEHPDGHWRVFUHHQIUHVKZDWHUVIRUWKHVH F\DQRSKDJHV EHFDXVH WKH\ DUH DEXQGDQW LQ PDULQH V\VWHPV ZKHUHWKH\UHSUHVHQWWKHRQO\ JURXSWKDWFDQ LQIHFW SK\FRHU\WKULQULFK LVRODWHV RI Synechococcus VHHEHORZ
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ynechococcus VSS LQ PDULQH ZDWHUV FRQWDLQHG YLVLEOH YLUXVOLNH SDUWLFOHV 3URFWRU DQG )XKUPDQ DQGWKDWYLUXVHVLQIHFWLQJPDMRUPDULQHSULPDU\ SURGXFHUV LQFOXGLQJ Synechococcus VSS FRXOG EH UHDGLO\ LVRODWHG IURP VHDZDWHU 6XWWOH HW DO 0RVW RI WKH F\DQRSKDJHV WKDW ZHUH LVRODWHG ZHUH 0\RYLULGDH DOWKRXJK 6W\ORYLULGDH DQG 3RGRYLULGDH ZHUH DOVR LVRODWHG 6XWWOH DQG &KDQ 6XWWOH HW DO :DWHUEXU\DQG9DORLV ,W ZDV VRRQ DSSDUHQW WKDW YHU\ KLJK FRQFHQWUDWLRQV RI F\DQRSKDJHV WKDW LQIHFW SK\FRHU\WKULQULFK VWUDLQV RI PDULQH Synechococcus VSS ZHUH SUHVHQW LQ D YDULHW\RIHQYLURQPHQWV ,QFRDVWDO ZDWHUV RIWKH*XOI RI 0H[LFR YLUXVHV WKDW LQIHFW Synechococcus 6WUDLQV '&DQG6<1 RFFXUUHGURXWLQHO\DWFRQFHQWUDWLRQV RI P/ DQG JUHDWHU 6XWWOH DQG &KDQ 6LPLODUO\:DWHUEXU\DQG9DORLV IRXQG LQIHFWLRXV F\DQRSKDJH FRQFHQWUDWLRQV ! P/ LQ :RRGV +ROH +DUERU ZKLOH *DU]D DQG 6XWWOH XQSXEOLVKHG GDWD UHFRUGHG DEXQGDQFHV LQ IMRUGV DORQJ WKH FRDVW RI%ULWLVK &ROXPELDUDQJLQJ EHWZHHQ 7KH KLJKHVW DQG P/ LQ $XJXVW FRQFHQWUDWLRQ RI LQIHFWLRXV F\DQRSKDJHV UHSRUWHG IRU
D QDWXUDO HQYLURQPHQW ZDV GXULQJ D EORRP RI Synechococcus VSS DW D VWDWLRQ RYHUO\LQJ WKH FRQWLQHQWDO VKHOI LQ WKH ZHVWHUQ *XOI RI 0H[LFR 6XWWOH HW DO 7KH DEXQGDQFH RIF\DQRSKDJHV WKDW LQIHFWHG Synechococcus 6WUDLQ '& ZDV FD P/ ZKLFKUHSUHVHQWHG DERXW RIWKHHQWLUHYLUDO FRPPXQLW\ DVGHWHUPLQHGIURPGLUHFWFRXQWV RIYLUDO SDUWLFOHV ,Q QHDUVKRUH ZDWHUV RIWKH *XOIRI0H[LFR 3RUW $UDQVDV 7H[DV DQG :RRGV +ROH LQIHFWLRXV F\DQRSKDJH FRQFHQWUDWLRQV UHDFKHG WKHLU KLJKHVW DEXQGDQFH GXULQJ WKH VXPPHU DQG IHOO ZKHQ ZDWHU WHPSHUDWXUHV ZHUH KLJKHVW DQG Synechococcus VSS ZDV PRVW DEXQGDQW :DWHUEXU\ DQG 9DORLV 6XWWOHDQG&KDQ 7KH DEXQGDQFH RI F\DQRSKDJHV WKDW LQIHFW PDQ\ QRQSK\FRHU\WKULQ FRQWDLQLQJ Synechococcus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ynechococcus VWUDLQV :+ DQG:+ ZHUHIUHTXHQWO\ VLPLODU EXW DWRWKHUWLPHVZHUHPRUHWKDQRQHRUGHURIPDJQLWXGH GLIIHUHQW :DWHUEXU\ DQG 9DORLV 6LPLODU REVHUYDWLRQVZHUHPDGHIRUFRDVWDOZDWHUVRIWKH*XOI RI 0H[LFR ZKHUH WKH FRQFHQWUDWLRQ RI YLUXVHV WKDW LQIHFW VRPH SK\FRHU\WKULQULFK VWUDLQV RI Synechococcus %* DQG 6<1 ZHUH OHVV WKDQ RI WKRVH LQIHFWLQJ Synechococcus 6WUDLQ '& 6XWWOHDQG&KDQ ,W LV FOHDU WKDW WKH DEXQGDQFH RI LQIHFWLRXV F\DQRSKDJHV LV WLHG WR WKDW RI Synechococcus VSS DQGQRWGLUHFWO\WRWHPSHUDWXUH$UHJUHVVLRQDQDO\VLV RQ VHDVRQDO GDWDFROOHFWHG GXULQJ DWKUHH\HDUSHULRG RQ WKH DEXQGDQFH RI F\DQRSKDJHV LQIHFWLQJ VHYHUDO VWUDLQV RI Synechococcus VSS YHUVXV WHPSHUDWXUH RQO\H[SODLQHG WR RIWKHYDULDQFH 6XWWOHDQG &KDQ 7KLV FRQFOXVLRQ LV FRQVLVWHQW ZLWK GDWD FROOHFWHG RQ D VHDZDUG WUDQVHFW IURP WKH FRDVW RI 7H[DV ,Q VXUIDFH ZDWHUV ERWK WKH DEXQGDQFH RI F\DQRSKDJHV WKDW LQIHFW Synechococcus 6WUDLQ '& DQG WKH FRQFHQWUDWLRQ RI Synechococcus FHOOV GURSSHGPRUHWKDQWHQIROGZKHUHDVWKHWHPSHUDWXUH LQFUHDVHG E\ DERXW & 6XWWOH DQG &KDQ $W
&$6XWWOH WKH VHDZDUG HQG RI WKH WUDQVHFW LQ ZDWHUV ZKLFK RYHUODLGWKHFRQWLQHQWDO VKHOI F\DQRSKDJH DEXQGDQFH GURSSHGIURPDERXW P/ DWWKHVXUIDFHWRFD P/ DW PGHSWK ZKLOH ZDWHUWHPSHUDWXUHGURSSHG WR DERXW & 5RGGD FRPSOHWHG D VHDVRQDO VWXG\LQWKH VDPHHQYLURQPHQWDQG FDPHWR D VLPLODU FRQFOXVLRQ 3ULQFLSOH &RPSRQHQW $QDO\VLV GHPRQVWUDWHG WKDW WKH DEXQGDQFH RI LQIHFWLRXV F\DQRSKDJHV ZDV UHODWHG ILUVW DQG PRVW VWURQJO\ WR WKH DEXQGDQFH RI Synechococcus DQG VHFRQGO\ WR WHPSHUDWXUH $W VWDWLRQV LQ WKH FHQWUDO *XOI RI 0H[LFR LQIHFWLRXV F\DQRSKDJHV ZHUH XQGHWHFWDEOH XQSXEOLVKHGGDWD )RU PDULQH HQYLURQPHQWV WKHUH LV HYLGHQFH IRU D WKUHVKROG LQ WKH DEXQGDQFH RI Synechococcus VSS EH\RQG ZKLFK LQIHFWLRXV F\DQRSKDJH FRQFHQWUDWLRQ LQFUHDVHV JUHDWO\ $ VKDUS LQFUHDVH LQ LQIHFWLRXV F\DQRSKDJH FRQFHQWUDWLRQV IURP FD P/ WR P/ ZDV REVHUYHG WR EH DVVRFLDWHG ZLWK Synechococcus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ynechococcus VS VWUDLQ '& ZHUH PHDVXUHG LQ WKH VHGLPHQW DQG RYHUOD\LQJ ZDWHU DWD VLWH NP RIIVKRUH LQ WKH *XOIRI0H[LFR LQ0D\$XJXVWDQG'HFHPEHURI 5RGGDHWDO LQUHYLHZ &\DQRSKDJH WLWHUV UDQJHG IURP [ P/ DW WKH VHD VXUIDFH WR [ P/ DW P MXVW DERYH WKH VHGLPHQW VXUIDFH ,Q WKH VHGLPHQW WKH WLWHU RI LQIHFWLRXV F\DQRSKDJHV GHFUHDVHG H[SRQHQWLDOO\ ZLWK GHSWK IURP [ P/ DW WKH ZDWHUVHGLPHQW LQWHUIDFH WR [ P/ DW FP EHORZWKHVHGLPHQWVXUIDFH)LJ 7KHVHVHGLPHQWV ZHUH DQDHURELFEHORZ FP 7KH VRXUFHRILQIHFWLRXV F\DQRSKDJHVDWFPGHSWKLQVHGLPHQWVO\LQJEHORZ WKH SKRWLF ]RQH LV DQ LQWULJXLQJ TXHVWLRQ 7KH FRQFHQWUDWLRQ RI F\DQRSKDJHV LQ WKH ZDWHU LPPHGLDWHO\DERYHWKHVHGLPHQWVXUIDFHZDVDERXWDQ RUGHU RI PDJQLWXGH OHVV WKDW DW WKH VHGLPHQWZDWHU LQWHUIDFH 7KLV VXJJHVWV WKDW WKH YLUXVHV SUREDEO\ DWWDFKWRSDUWLFOHV HLWKHUGLUHFWO\ RUYLDLQIHFWHG FHOOV DQG VLQN WR WKH VHGLPHQW VXUIDFH ZKHUH WKH\ DUH VXEVHTXHQWO\ EXULHG %DVHG RQ DEVROXWH GDWLQJ HVWLPDWHV VHGLPHQWDWLRQ UDWHV LQ WKLV UHJLRQ RI WKH
&KDSWHU &\DQRSKDJHV
9 )DWH RI &\DQRSKDJHV LQ WKH 1DWXUDO (QYLURQPHQW
Fig. 2. $EXQGDQFH RI F\DQRSKDJHV LQIHFWLQJ PDULQH Synechococcus 6WUDLQ '& LQ VHGLPHQWV NP RIIVKRUH LQ WKH *XOI RI 0H[LFR %DVHG RQ DEVROXWH GDWLQJ HVWLPDWHV DQG VHGLPHQWDWLRQUDWHVWKHYLUXVHVDWFPGHSWKDUHLQH[FHVVRI \HDUVROG
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hormidium persicinum 2KNL DQG )XMLWD 8QOLNH WKHSRGRYLUXVHVWKDWKDYH VKRUW QRQFRQWUDFWLOH WDLOV DQG LQIHFW RWKHU ILODPHQWRXV F\DQREDFWHULDWKHYLUXVZKLFKLQIHFWHGP. persicinum SRVVHVVHG D ORQJ IOH[LEOH WDLO DQG ZDV PRUSKRORJLFDOO\ VLPLODU WR VLSKRYLUXVHV $W SUHVHQW WKHUH LV QR LQIRUPDWLRQ RQ WKH GLVWULEXWLRQ RU DEXQGDQFHRIWKHVHF\DQRSKDJHVLQQDWXUH 7R GDWH YLUXVHV ZKLFK LQIHFW RWKHU PDULQH ILODPHQWRXVF\DQREDFWHULDZHUHQRWLVRODWHG ,WZRXOG EHSDUWLFXODUO\LQWHUHVWLQJLIYLUXVHV FDQEHIRXQGWKDW LQIHFW WKH RVFLOODWRULDQ Trichodesmium; D PDMRU SULPDU\SURGXFHUDQGQLWURJHQIL[HULQWURSLFDOZDWHUV &DUSHQWHU &DUSHQWHU DQG 5RPDQV &KDSWHU *LYHQ WKH SUHYDOHQFH RI RWKHU YLUXVHV ZKLFKLQIHFWILODPHQWRXVF\DQREDFWHULDLWVHHPVOLNHO\ WKDWWKH\H[LVW
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
WKHGHFD\UDWHVRIF\DQRSKDJHLVRODWHVXQGHUWKHVDPH FRQGLWLRQV *DU]D DQG 6XWWOH LQ UHYLHZ 'XULQJ WKH VSULQJDQG ZLQWHU GHFD\UDWHV RIF\DQRSKDJH LVRODWHV DQG QDWXUDO F\DQRSKDJH FRPPXQLWLHV ZHUH JHQHUDOO\ VLPLODU EXW GXULQJ VXPPHU GHFD\ UDWHV RI LVRODWHV ZHUH DV PXFK DV WZRIROG KLJKHUWKDQ LQ WKH QDWXUDO FRPPXQLWLHV $V WKH VHQVLWLYLW\ RI WKH LVRODWHV WR GDPDJLQJ UDGLDWLRQ GLG QRW FKDQJH WKHVH UHVXOWV LQGLFDWH WKDW WKH QDWXUDO YLUDO FRPPXQLW\ EHFDPH PRUH UHVLVWDQW WR GDPDJH FDXVHG E\ VXQOLJKW 7KH PHFKDQLVPZKHUHE\UHVLVWDQFHWR GDPDJLQJUDGLDWLRQ FDQ EH LQFUHDVHG LV XQNQRZQ EXW SRWHQWLDOO\ FRXOG LQYROYHFKDQJHVWRWKH'1$LWVHOI 7KH SUHVHQFH RI KLJK FRQFHQWUDWLRQV RI LQIHFWLRXV F\DQRSKDJHVLQVXUIDFHZDWHUV LVDWWULEXWDEOH LQSDUW WR LQFUHDVHG UHVLVWDQFH RI QDWXUDO FRPPXQLWLHV WR VXQOLJKW GXULQJ SHULRGV RI KLJKHVW 89% IOX[ DQG UHSDLU RI GDPDJHG '1$ E\ KRVWFHOO DQG YLUDO HQFRGHG PHFKDQLVPV 0LOOHU DQG .RNMRKQ )ULHGEHUJHWDO &KDSWHU GXULQJGDUNQHVVRU ZKHQWKH YLUXVHV DQG KRVWFHOOV DUH PL[HG EHORZ WKH VXUIDFH0L[LQJEHORZWKHVXUIDFHZLOOQRWRQO\UHVXOW LQDGHFUHDVHLQH[SRVXUHWR89% EXWZLOODOVRUHVXOW LQ D VKLIW LQ VSHFWUDO TXDOLW\ WR ORQJHU ZDYHOHQJWKV DOORZLQJWKHSRWHQWLDOUHSDLURIWKHGDPDJHG'1$E\ WKH OLJKWGHSHQGHQW HQ]\PH SKRWRO\DVH 7KH ORVV RI YLUDO LQIHFWLYLW\ IROORZLQJ H[SRVXUH WR VXQOLJKW LV ODUJHO\ WKH UHVXOW RI GDPDJH WR '1$ IURP WKH IRUPDWLRQRIS\ULPLGLQH GLPHUV H[SRVXUHWR VXQOLJKW JHQHUDOO\ ZLOO QRW LQWHUIHUH ZLWK WKH DWWDFKPHQW RI F\DQRSKDJHV WR KRVW FHOOV DQG WKH VXEVHTXHQW LQMHFWLRQ RI YLUDO QXFOHLF DFLGV 7KH GDPDJHG YLUDO '1$ FDQ WKHQ EH UHSDLUHG E\ KRVWFHOO PHFKDQLVPV VXFK DV SKRWRUHDFWLYDWLRQ LH WKH UHVWRUDWLRQ RI LQIHFWLYLW\ WR :LUUDGLDWHG YLUXVHV XSRQ H[SRVXUH WR YLVLEOH OLJKW LQ WKH SUHVHQFH RI KRVW EDFWHULD 'XOEHFFR %RWK ILODPHQWRXV DQG XQLFHOOXODU F\DQREDFWHULD DUH NQRZQ WR KDYH YHU\ HIILFLHQW OLJKWGHSHQGHQW PHFKDQLVPV WKDW FDQ UHSDLU 89GDPDJHG F\DQRSKDJH '1$ :X HW DO 6LQJK$VDWR$POD+ZDQJ/HHHW DO /HYLQHDQG7KLHO 0RUHRYHU LWZDV GHPRQVWUDWHG UHFHQWO\ WKDW SKRWRUHDFWLYDWLRQ FDQ EH YHU\ LPSRUWDQW LQ PDLQWDLQLQJ WKH LQIHFWLYLW\ RI QDWXUDO SKDJH FRPPXQLWLHV LQ WKH VHD :HLQEDXHU HW DO :LOKHOPHWDOLQSUHVVLQUHYLHZ 7KH VLJQLILFDQFH RI VXQOLJKW UHODWLYH WR RWKHU IDFWRUV UHVSRQVLEOH IRU WKH GHFD\ RI F\DQRSKDJH LQIHFWLYLW\ LQ WKH XSSHU PL[HG OD\HU RI WKH ZDWHU FROXPQ GHSHQGV RQ D QXPEHU RI WKLQJV LQFOXGLQJ LQVRODWLRQ DW WKH VXUIDFH WKH RSWLFDO FODULW\ RI WKH ZDWHUDQGWKHGHSWKRIWKHPL[HGOD\HU ,QFRQWUDVWWR
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
9, (IIHFWRI&\DQRSKDJHVRQ &\DQREDFWHULDO 3RSXODWLRQV DQG &RPPXQLWLHV &OHDUO\ WKH IDFW WKDW LQIHFWLRXV F\DQRSKDJHV H[LVW LQ DTXDWLF V\VWHPV DQG WKH IDFW WKDW WKH\ KDYH D UHODWLYHO\ VKRUW UHVLGHQFH WLPH VHH DERYH LPSOLHV WKDW WKH\ DUH UHVSRQVLEOH IRU VRPH SRUWLRQ RI F\DQREDFWHULDO PRUWDOLW\ *LYHQ WKDW WKLV PRUWDOLW\ LV VHOHFWLYH LW FDQ DIIHFW FRPPXQLW\ VWUXFWXUH RI SODQNWRQLFFRPPXQLWLHV7KHUHZHUHWZRSULPDU\IRFL IRU VWXGLHV RQ WKH LPSDFW RI F\DQRSKDJHV RQ WKH
&KDSWHU &\DQRSKDJHV PRUWDOLW\ RI F\DQREDFWHULD 7KH ILUVW DQG HDUOLHVW ZRUN GHDOW ZLWK IUHVKZDWHUV DQG ZDV SULPDULO\ FRQFHUQHG ZLWK WKH SRWHQWLDO XVH RI F\DQRSKDJHV DV ELRORJLFDO FRQWURO DJHQWV WR HOLPLQDWH RU UHGXFH QXLVDQFH DQG WR[LF EORRPV RI F\DQREDFWHULD 7KH LPSHWXV IRU WKLV ZRUN ZDV WKH PDMRU HFRORJLFDO DQG HFRQRPLF HIIHFWV RIVXFK EORRPV 7KH VHFRQG IRFXV ZDVWRWU\DQGGHWHUPLQHWKHHIIHFWRIF\DQRSKDJHVRQ WKH PRUWDOLW\ RI Synechococcus VSS LQ PDULQH HQYLURQPHQWV7KLVUHVHDUFKZDVGULYHQODUJHO\E\WKH LPSRUWDQFH RISynechococcus VSS DV PDMRU SULPDU\ SURGXFHUV LQ WKH PDULQH HQYLURQPHQW DQG WKH KLJK DEXQGDQFHRIFRRFFXUULQJF\DQRSKDJHV
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ynechococcus VSS 8QIRUWXQDWHO\ DOORIWKHPHWKRGV UHTXLUH VSHFLILF DVVXPSWLRQV ZKLFK ZHDNHQ WKH FRQFOXVLRQV 2QWKHRWKHUKDQG LWLVHQFRXUDJLQJWKDW HQWLUHO\ LQGHSHQGHQW PHWKRGV OHDG WR VLPLODU HVWLPDWHV RI F\DQREDFWHULDO PRUWDOLW\ DV D FRQVHTXHQFHRIYLUDOLQIHFWLRQ3ODWH F
)UHTXHQF\RIOQIHFWHG&HOOV 7KH ILUVW DWWHPSW WR GHGXFH WKH HIIHFWV RIYLUXVHV RQ WKH PRUWDOLW\ RI Synechococcus VSS LV DOVR SHUKDSV WKH PRVW HOHJDQW 7KH DSSURDFK ZDV EDVHG RQ WKH SULQFLSOH WKDW LI WKH SURSRUWLRQ RI WKH ODWHQW SHULRG WKH WLPH EHWZHHQ ZKHQ WKH FHOOV DUH LQIHFWHG DQG ZKHQ O\VLV EHJLQV WR RFFXU GXULQJ ZKLFK YLUDO SDUWLFOHV DUH YLVLEOH ZLWKLQFHOOV LV NQRZQ DQG LIWKH SURSRUWLRQRIFHOOVFRQWDLQLQJYLVLEOHYLUDOSDUWLFOHV LV DOVR NQRZQ WKHQ WKH SURSRUWLRQ RI WKH SRSXODWLRQ ZKLFK LV LQIHFWHG FDQ EH FDOFXODWHG 3URFWRU DQG )XKUPDQ H[DPLQHG VDPSOHV IURP ILYH ORFDWLRQV LQ WKH ZHVWHUQ $WODQWLF DQG &DULEEHDQ DQG GHWHUPLQHG WKDW RQ DYHUDJH WR RI F\DQREDFWHULDOFHOOVFRQWDLQHGYLVLEOHSKDJHSDUWLFOHV
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lectonema boryanum 87(; LQIHFWLRXV YLUXVHV DSSHDUHG DSSUR[LPDWHO\ KDOIZD\ WKURXJK WKH ODWHQWSHULRG 3DGDQHWDO +RZHYHUWKHUHDUH PDQ\GLIIHUHQFHVLQWKHUHSOLFDWLRQ RIF\DQRSKDJHV LQ ILODPHQWRXV DQG XQLFHOOXODU F\DQREDFWHULD DQG LW LV OLNHO\ LQDSSURSULDWH WR LQIHU GHWDLOV RIWKH O\WLF F\FOH LQ PDULQH Synechococcus VSS IURP REVHUYDWLRQV PDGH RQ IUHVKZDWHU ILODPHQWRXV F\DQREDFWHULD 'DWD IURP D IUHVKZDWHU Synechococcus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
RI WKH O\WLF F\FOH EDVHG RQ WKH GDWD RI 0DF.HQ]LHDQG+DVHONRUQE 7KLVZRXOGPHDQ WKDW LI WR RISynechococcus FHOOV FRQWDLQHG YLVLEOH YLUXV SDUWLFOHV WKHQ WR RI WKH WRWDO SRSXODWLRQ ZDV LQIHFWHG $V WKH O\WLF F\FOH RI F\DQRSKDJHVLQXQLFHOOXODUF\DQREDFWHULDUDQJHVIURP FD WRKRXUV0DF.HQ]LHDQG+DVHONRUQ E 6DIIHUPDQ HWDO 6XWWOH DQG &KDQ .LP DQG&KRL RQHFDQQRWVWDWHZLWK FHUWDLQW\ZKDW WKLVUHSUHVHQWVLQWHUPV RIGDLO\PRUWDOLW\ DOWKRXJKLW VHHPV OLNHO\ WKDW VHYHUDO SHUFHQW RI WKH Synechococcus SRSXODWLRQLVO\VHGRQDGDLO\EDVLV,Q DVWHDG\VWDWHV\VWHPLQZKLFKRIWKHFHOOVOLYHWR UHSURGXFHDQG GLH WKHQVRPHWKLQJWKDWUHPRYHV RI WKH FHOOV ZLOO DFFRXQW IRU RI WKH PRUWDOLW\ 3URFWRUDQG )XKUPDQ 7KHUHIRUH LI WR RIWKH SRSXODWLRQ LV LQIHFWHG DQG WKH O\WLF F\FOHLVOHVVWKDQWKHGRXEOLQJWLPHRIWKHFHOOVWKHQ WRRIWKHWRWDOPRUWDOLW\PD\EHWKHUHVXOWRIYLUDO LQIHFWLRQLIDOOLQIHFWHGFHOOVDUHNLOOHGE\YLUXVHV
'HFD\5DWHV $QRWKHUDSSURDFK WKDWZDV XVHG WR LQIHUWKH HIIHFWRI YLUXVHV RQ F\DQREDFWHULDO PRUWDOLW\ ZDV WR DVVXPH WKDWWKHUDWHV RIUHPRYDO DQGSURGXFWLRQRILQIHFWLRXV F\DQRSKDJHV ZHUH EDODQFHG ,IWKH QXPEHU RIYLUXVHV SURGXFHGSHUO\WLFHYHQWLVNQRZQWKHQWKHQXPEHURI FHOOV LQ ZKLFK O\VLV RFFXUUHG FDQ EH HVWLPDWHG IURP WKHQXPEHURIYLUXVHVWKDWZHUH ORVWIURPWKHV\VWHP 7KLV DSSURDFK ZDV ILUVW XVHG WR HVWLPDWH PRUWDOLW\ UDWHVLQPDULQHEDFWHULDFDXVHGE\SKDJHV+HOGDODQG %UDWEDN 6XWWOH DQG &KHQ 7KH VDPH DSSURDFKFDQEHXVHGIRUF\DQREDFWHULD ,QIHFWLRXV F\DQRSKDJHV DUH FRQWLQXRXVO\ ORVW IURP DTXDWLF HQYLURQPHQWV EHFDXVH RID YDULHW\ RIIDFWRUV LQFOXGLQJVRODUUDGLDWLRQ DQG DWWDFKPHQWWRSDUWLFOHV 7KH ILUVW DWWHPSWWR HVWLPDWH PRUWDOLW\ UDWHV LPSRVHG RQ QDWXUDO FRPPXQLWLHV RI Synechococcus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
&$6XWWOH UHVXOWHGLQ DQHVWLPDWHRI FHOOV P/ G LQZKLFK O\VLV ZRXOG KDYH WR RFFXU LIYLUDO SURGXFWLRQ ZDV WR EDODQFHWKHUHPRYDO UHVXOWLQJ IURP H[SRVXUH WR VRODU UDGLDWLRQ 7KH FRQFHQWUDWLRQ RISynechococcus DWWKLV ORFDWLRQ ZDV FHOOV P/ WKHUHIRUH LQ DVWHDG\ VWDWH V\VWHP LW ZDV HVWLPDWHG WKDW DERXW RIWKH FHOOVZRXOGKDYHWREHO\VHGRQDGDLO\EDVLVLQRUGHU WR EDODQFHWKHUHPRYDOUHVXOWLQJIURP VRODUUDGLDWLRQ $V PRUH GDWD EHFDPH DYDLODEOH VHH EHORZ LW ZDV DSSDUHQWWKDWWKH DYHUDJHGHFD\UDWH DQGEXUVWVL]H LQ QDWXUDO FRPPXQLWLHV ZHUH LQ HUURU D PDJQLWXGH RI DERXW WKUHHIROG DV RQH HUURU ZDV DQ RYHUHVWLPDWH DQG WKH RWKHU DQ XQGHUHVWLPDWH WKH HIIHFWV RI WKHVH SURFHVVHV RQ WKH HVWLPDWHG PRUWDOLW\ RI Synechococcus VSS VKRXOGEHPLQLPDO 0RUH UHFHQWO\ LQ VLWX GHFD\ UDWHV RI QDWXUDO FRPPXQLWLHV RI F\DQRSKDJHV ZHUH XVHG WR HVWLPDWH WKH HIIHFW RI YLUXVHV RQ WKH PRUWDOLW\ RI Synechococcus VSS *DU]D *DU]D DQG 6XWWOH LQ UHYLHZ ,Q WZR VWDWLRQV RIIWKH FRDVW RI7H[DV WKH GHFD\UDWHVRILQIHFWLRXV F\DQRSKDJHV ZHUH HVWLPDWHG WREH DQG G ZKHQ DYHUDJHGRYHUKRXUV DQGLQWHJUDWHG RYHUWKHVXUIDFHPL[HG OD\HUDQG P UHVSHFWLYHO\ 7KH FRQFHQWUDWLRQV RI F\DQRSKDJHV WKDW LQIHFWHG Synechococcus VWUDLQ '& LQWKHVHVDPSOHVZHUH [ DQG [ P/ UHVSHFWLYHO\ &RQVHTXHQWO\ EDODQFLQJ WKH UHPRYDO UDWHV RI LQIHFWLRXV F\DQRSKDJHV ZRXOG UHTXLUH DYHUDJH SURGXFWLRQ UDWHV RI [ DQG [ F\DQRSKDJHV P/ G UHVSHFWLYHO\ %DVHG RQ 7(0 REVHUYDWLRQV RIWKHVH VDPSOHV WKH DYHUDJH EXUVW VL]H ZDV HVWLPDWHG WR EH YLUXVHV SHU Synechococcus FHOOO\VHG6XWWOHHWDO 7KHUHIRUHWKHQXPEHUV RI Synechococcus FHOOV LQ ZKLFK O\VLV PXVW KDYH RFFXUUHG LQ RUGHUWR EDODQFHWKH PHDVXUHG ORVV UDWHV RI LQIHFWLRXV F\DQRSKDJHV ZHUH DQG ZKLFK UHSUHVHQWV DSSUR[LPDWHO\ DQG RI WKH Synechococcus SRSXODWLRQ UHVSHFWLYHO\ 7KLV LV D PLQLPXP HVWLPDWH DV LW RQO\ FRQVLGHUHG YLUXVHV WKDW FDXVHG O\VLV RI Synechococcus VWUDLQ '& ,Q DGGLWLRQLWDVVXPHGWKDWDOOSynechococcus FHOOVZHUH UHVSRQVLEOH IRUSURGXFWLRQ RILQIHFWLRXV F\DQRSKDJHV ,Q UHDOLW\ WKHUH PD\EH VXEVHWV RIWKH Synechococcus SRSXODWLRQZKLFKDUHUHVSRQVLEOHIRUPRVWRIWKHYLUDO SURGXFWLRQ ,WLV HQFRXUDJLQJWKDWUHVXOWV IURPWZR LQGHSHQGHQW DSSURDFKHV ± F\DQRSKDJHGHFD\UDWHH[SHULPHQWV DQG REVHUYDWLRQV RI WKH SURSRUWLRQ RI LQIHFWHG Synechococcus FHOOV LQ QDWXUDO FRPPXQLWLHV ± ZHUH FRQVLVWHQW 7KHVH HVWLPDWHV VXJJHVWWKDWWKH PRUWDOLW\ FDXVHG E\ F\DQRSKDJHV DOWKRXJK QRW ODUJH LV VLJQLILFDQW DQG UHVXOWV LQ WKH LQIHFWLRQ DQG O\VLV RI
&KDSWHU &\DQRSKDJHV VHYHUDO SHUFHQW RIWKH FHOOV LQ QDWXUDO Synechococcus FRPPXQLWLHVRQDGDLO\EDVLV
&RQWDFW5DWHV $WKLUGDSSURDFKWKDWFDQEHXVHGWRPDNHLQIHUHQFHV DERXWWKHPD[LPXPHIIHFWRIYLUXVHV RQWKHPRUWDOLW\ RI F\DQREDFWHULD LV WR H[DPLQH WKH FRQWDFW UDWHV EHWZHHQ YLUXVHV DQGSRWHQWLDO KRVWFHOOV 0XUUD\ DQG -DFNVRQ ,I WKH DEXQGDQFHV RI LQIHFWLRXV YLUXVHV DQG KRVW FHOOV DUH NQRZQ WKH FRQWDFW UDWH EHWZHHQ WKH WZR FDQ EH FDOFXODWHG 6XWWOH &KDQ DVSh G'Y 93ZKHUHSh LVWKH6KHUZRRG QXPEHU GLPHQVLRQOHVV IRU Synechococcus G LV WKH FHOO GLDPHWHU [ FP 'Y LV WKH GLIIXVLYLW\ RIWKH YLUXVHV [ FP G 9 LV WKHFRQFHQWUDWLRQRILQIHFWLRXVYLUXVHV FP DQG3LV WKH FRQFHQWUDWLRQ RI Synechococcus FP $Q HVVHQWLDOO\ HTXLYDOHQW FDOFXODWLRQ LV WR LQIHU LQIHFWLRQ UDWHV IURP WKHRUHWLFDO UDWHV RISKDJH DGVRUSWLRQ DQG PHDVXUHG FRQFHQWUDWLRQV RI Synechococcus DQG F\DQRSKDJHV :DWHUEXU\ DQG 9DORLV ,Q ERWK DSSURDFKHV WKH SRWHQWLDO LQIHFWLRQ UDWH LV GLUHFWO\ SURSRUWLRQDO WR WKH SURGXFW RIWKH YLUDO DQG KRVW FHOO DEXQGDQFHV $V WKH LQIHFWLRQ UDWH FDQQRW H[FHHG WKH UDWH DW ZKLFK YLUXVHV HQFRXQWHU RU DGVRUE WR KRVW FHOOV WKHVH UDWHV SURYLGH PD[LPXP HVWLPDWHV RI WKH YLUDO LQIHFWLRQUDWH RUWKHSRWHQWLDO IRUYLUDO LQIHFWLRQ 6XWWOH %DVHG RQ DEXQGDQFH GDWD IRU F\DQRSKDJHV DQG Synechococcus FHOOV LQ :RRGV +ROH +DUERU DQG ZDWHUV IDUWKHURIIVKRUH:DWHUEXU\ DQG9DORLV DUJXHG WKDW F\DQRSKDJHV FRXOG RQO\ EH UHVSRQVLEOH IRU WKH PRUWDOLW\ RI DW PRVW D IHZ SHUFHQW RI Synechococcus FHOOV RQ D GDLO\ EDVLV +RZHYHU LQ WKHVHZDWHUVWKHKLJKHVWHQFRXQWHUUDWHRFFXUUHGZKHQ WKH DEXQGDQFHV RI F\DQRSKDJHV DQG Synechococcus ZHUH [ DQG [ P/ UHVSHFWLYHO\ ,Q FRQWUDVW LQ WKH FRDVWDO ZDWHUV RI7H[DV F\DQRSKDJH DEXQGDQFHV DQG KHQFH HQFRXQWHU UDWHV FDQ EH PXFK KLJKHU DQG UHVXOW LQ PRVW Synechococcus FHOOV EHLQJ FRQWDFWHG E\ LQIHFWLRXV F\DQRSKDJH RQ D GDLO\ EDVLV 6XWWOH DQG &KDQ 7KHVH FDOFXODWLRQV DVVXPH WKDW WKH FRQFHQWUDWLRQ RI F\DQRSKDJHV WKDW LQIHFW VSHFLILFKRVWV VXFK DV Synechococcus 6WUDLQV '& RU :+ DUH LQGLFDWLYH RI WKH WRWDO QXPEHU RI LQIHFWLRXV F\DQRSKDJHV DQG WKDW DOO LQIHFWLRXV F\DQRSKDJHV KDYH WKH SRWHQWLDO WR LQIHFW DOO Synechococcus FHOOV ,W LV SUREDEO\ UHDVRQDEOH WR DVVXPHWKDWWKHDEXQGDQFHRIF\DQRSKDJHVWKDWLQIHFW WKHVHVWUDLQVRISynechococcus LVUHSUHVHQWDWLYHRIWKH WRWDOQXPEHURILQIHFWLRXVF\DQRSKDJHVKRZHYHU WKH
SURSRUWLRQ RIWKH Synechococcus SRSXODWLRQ WKDW FDQ EH LQIHFWHG E\ WKH PRVW DEXQGDQWF\DQRSKDJHV OLNHO\ YDULHVVHHEHORZ
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
1. +RVWFHOO5HVLVWDQFH 1RW RQO\ FDQ FRQWDFW UDWHV EH XVHG WR HVWDEOLVK WKH PD[LPXP QXPEHU RI FHOOV WKDW FDQ EH LQIHFWHG E\ F\DQRSKDJHV RQ D GDLO\ EDVLV EXW LI LQGHSHQGHQW HVWLPDWHV RI PRUWDOLW\ DV WKH UHVXOW RI LQIHFWLRQ DUH DYDLODEOHWKHQFRQWDFWUDWHV FDQDOVR EHXVHG WR LQIHU WKH SURSRUWLRQ RID Synechococcus SRSXODWLRQ WKDW LV UHVLVWDQW WR LQIHFWLRQ )RU H[DPSOH LQ WKH VXUIDFH PL[HG OD\HU RI WKH FRDVWDO ZDWHUV RI 7H[DV WKH SHUFHQWDJH RI Synechococcus FHOOV LQ ZKLFK O\VLV RFFXUUHGGXHWRYLUDOLQIHFWLRQZDVHVWLPDWHGWRUDQJH IURP SHU GD\ LQ QHDUVKRUH ZDWHUV WR SHU GD\ NP RIIVKRUH 6XWWOH DQG &KDQ 7KLV FDOFXODWLRQ DVVXPHG WKDW VRODU UDGLDWLRQ ZDV UHVSRQVLEOH IRU PRVW RIWKH GHVWUXFWLRQ RI LQIHFWLYLW\ DQG WKDW WKH EXUVW VL]H ZDV DERXW 6XWWOH DQG &KDQ ,Q IDFW WKH GHVWUXFWLRQ UDWHV DQG EXUVW VL]HVZHUHSUREDEO\RYHUHVWLPDWHGE\DERXWWKUHHIROG VHH DERYH &DOFXODWHG FRQWDFW UDWHV LQ WKH VXUIDFH PL[HG OD\HU DWWKH RIIVKRUH VWDWLRQ LQGLFDWHG WKDW RI Synechococcus ZHUH FRQWDFWHG E\ LQIHFWLRXV F\DQRSKDJHV RQ D GDLO\ EDVLV 7KLV HVWLPDWH ZDV VLPLODU WR WKH HVWLPDWHG PRUWDOLW\ UDWH RI SHU GD\ DQG LQGLFDWHG WKDW PRVW FRQWDFWV UHVXOWHG LQ LQIHFWLRQ KHQFH PRVW Synechococcus ZHUH QRW UHVLVWDQW WR PRVW F\DQRSKDJHV ,Q FRQWUDVW LQ QHDUVKRUH ZDWHUV WKH HVWLPDWHG PRUWDOLW\ UDWH RI Synechococcus DV DUHVXOWRILQIHFWLRQZDV RQO\ SHU GD\ \HW RISynechococcus ZHUH FRQWDFWHG E\F\DQRSKDJHVHDFKGD\&RQVHTXHQWO\IHZFRQWDFWV UHVXOWHGLQLQIHFWLRQDQGWKLVVXJJHVWVWKDWHLWKHUPRVW Synechococcus ZHUH UHVLVWDQW RU WKDW WKH HIILFLHQF\ RILQIHFWLRQZDVYHU\ORZ $GLIIHUHQWDSSURDFKXVHGE\:DWHUEXU\DQG9DORLV SURYLGHG IXUWKHU HYLGHQFH WKDW ZKHQ Synechococcus DQG F\DQRSKDJHV DUH DEXQGDQW PRVW Synechococcus DUH UHVLVWDQW WR WKH PRVW DEXQGDQW F\DQRSKDJHV &ORQDO LVRODWHV RI Synechococcus DQG F\DQRSKDJHVZHUHREWDLQHGE\VHULDOGLOXWLRQRIZDWHU FROOHFWHG IURP :RRGV +ROH +DUERU 7KH F\DQREDFWHULDLVRODWHG IURPWKH KLJKHVWGLOXWLRQV LH WKRVH WKDW ZHUH PRVW DEXQGDQW ZHUH UHVLVWDQW WR WKH PRVW DEXQGDQW F\DQRSKDJHV ZKHUHDV F\DQREDFWHULD WKDWZHUHLVRODWHGIURPORZHUGLOXWLRQVZHUHVHQVLWLYH WRPDQ\RIWKHF\DQRSKDJHV 7KH DXWKRUV DUJXHGWKDW WKH REVHUYDWLRQ WKDW PRVW Synechococcus ZHUH UHVLVWDQWWRLQIHFWLRQE\WKHLUFRRFFXUULQJSKDJHVZDV FRQVLVWHQW ZLWK UHVXOWV IURP FRQWLQXRXV FXOWXUHV RI KHWHURWURSKLF EDFWHULD /HQVNL DQG /HYLQ /HQVNL ,QWKHODWWHUWKHUHLVDWKHFXOPLQDWLRQ
&$ 6XWWOH LQVWDEOHSKDJHDQGKRVWSRSXODWLRQVZLWKSKDJHDERXW DQRUGHURIPDJQLWXGHOHVVDEXQGDQWWKDQEDFWHULD 7KH VFHQDULR LQ WKH *XOI RI 0H[LFR LV GLIIHUHQW EHFDXVH WKH DEXQGDQFH RIF\DQRSKDJHV LV IUHTXHQWO\ IROG JUHDWHU WKDQ WKDW RI Synechococcus. $V D FRQVHTXHQFH FRQWDFW UDWHV DUH VXIILFLHQWO\ KLJK WKDW VPDOO SRSXODWLRQV RI VHQVLWLYH FHOOV FRXOG QRW SHUVLVW XQOHVV VXFFHVVIXO FRQWDFWV ZHUH YHU\ ORZ /RZ FRQWDFW VXFFHVV ZDV DOVR SURSRVHG DV DQ H[SODQDWLRQ IRU WKH SHUVLVWHQFH RI QDWXUDO SRSXODWLRQV RI WKH PDULQH SK\WRIODJHOODWH Micromonas pusilla LQ WKH SUHVHQFH RIKLJK WLWHUV RI LQIHFWLRXV YLUXVHV &RWWUHOO DQG6XWWOH )XUWKHU HYLGHQFH WKDW ORZ FRQWDFW VXFFHVV PLJKW H[SODLQ WKH SHUVLVWHQFH RI Synechococcus LQ WKH SUHVHQFH RI KLJK F\DQRSKDJH WLWHUV ZDV REWDLQHG XVLQJIOXRUHVFHQWODEHOHGYLUXVHVDVSUREHVWRLGHQWLI\ FHOOVVXVFHSWLEOHWRLQIHFWLRQ+HQQHVHWDO ,Q FXOWXUHV RI D FRDVWDO FORQH RI Synechococcus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ynechococcus YLUDO UHFHSWRUV ZHUH REVHUYHG WR EH YHU\ UDUH RU ORFDOL]HG LQ D VPDOO UHJLRQ RI WKH FHOO VXUIDFH 2EVHUYDWLRQVWKDWRQO\ DSRUWLRQ RIFHOOV LQD SRSXODWLRQ ZHUH VXVFHSWLEOH WR LQIHFWLRQ DW DQ\ RQH WLPHDUHFRQVLVWHQWZLWKWKHGDWDDQG PRGHORI/HQVNL 7KHPRGHOVXJJHVWVWKDWRQHZD\WRPDLQWDLQ VWDELOLW\ LQSRSXODWLRQV RIKRVWFHOOV DQG YLUXVHV LV WR KDYH D ODUJH IUDFWLRQ RI WKH KRVW SRSXODWLRQ EH SURWHFWHGIURPSKDJHLQIHFWLRQ DWDQ\ JLYHQWLPH EXW QRWWR KDYHWKLV SURWHFWLRQ EH KHULWDEOHDV LV WKHFDVH IRUUHVLVWDQFH
&KDSWHU &\DQRSKDJHV )RU RWKHU LVRODWHV RI Synechococcus, YLUXVHV ZHUH REVHUYHG WR DWWDFK WR DOO FHOOV LQ D FORQDO SRSXODWLRQ +HQQHV HWDO 3HUKDSV FRQWLQXRXV H[SUHVVLRQ RI YLUDO UHFHSWRUV LV PRUH SUHYDOHQW LQ RIIVKRUH LVRODWHV ZKHUH FRQWDFW IUHTXHQFLHV DUH ORZHU ZKHUH WKHUH LV OHVV VHOHFWLRQ WR DYRLG LQIHFWLRQ 6XWWOH DQG &KDQ DQG ZKHUH ORZ QXWULHQW FRQFHQWUDWLRQV UHVXOWLQVHOHFWLRQIRUKLJKOHYHOV RIQXWULHQWWUDQVSRUW SURWHLQV $OWHUQDWLYHO\ VRPH F\DQREDFWHULD PD\ EHFRPH OHVV UHVLVWDQW WR LQIHFWLRQ DIWHU EHLQJ PDLQWDLQHG LQ FXOWXUH IRU D QXPEHU RI \HDUV :DWHUEXU\DQG9DORLV ,Q PDQ\ ZD\V WKH UHODWLRQVKLS EHWZHHQ F\DQREDFWHULDDQGF\DQRSKDJHVLQIUHVKZDWHUDSSHDUV VLPLODU WR WKDW LQ PDULQH V\VWHPV DOWKRXJK WKH ELRPDVV RI QLWURJHQIL[LQJ ILODPHQWRXV DQG FRORQLDO F\DQREDFWHULD DUH IUHTXHQWO\ PXFK PRUH DEXQGDQW LQ IUHVKZDWHU/LNHWKHLUPDULQHFRXQWHUSDUWV IUHVKZDWHU F\DQREDFWHULD ZKLFK DUH VHQVLWLYH WR LQIHFWLRQ DUH QHYHU GRPLQDQW ZKHQ SKDJHV ZKLFK LQIHFW WKHP DUH DEXQGDQW 6DIIHUPDQ DQG 0RUULV 3DGDQ DQG 6KLOR 7KLV REVHUYDWLRQ LPSOLHV VHOHFWLRQ IRU F\DQREDFWHULDO FRPPXQLWLHV WKDW DUH UHVLVWDQW WR LQIHFWLRQ E\ WKH GRPLQDQW SKDJH ,Q FRQWLQXRXV FXOWXUH H[SHULPHQWV WKH DEXQGDQFHV RI ILODPHQWRXV F\DQREDFWHULD DQG LQIHFWLRXV F\DQRSKDJHV XQGHUZHQW D VHULHV RI LQFUHDVLQJO\GDPSHQHG RXWRISKDVH RVFLOODWLRQV HYHQWXDOO\ UHVXOWLQJ LQ D VWDEOH FRH[LVWHQFH EHWZHHQ WKH SKDJH DQG WKH KRVW &RZOLVKDZ DQG 0UVD &DQQRQ HW DO %DUQHWHWDO 'XULQJWKHH[SHULPHQWV%DUQHWHW DO IRXQG WKDW WKHUH ZDV VHOHFWLRQ IRU ERWK PXWDQW F\DQRSKDJH DQG F\DQREDFWHULD DQG WKLV UHVXOWHG LQ D UDQJH RI DGVRUSWLRQ DQG LQIHFWLRQ HIILFLHQFLHV 7KH UHVXOWV ZHUH VLPLODU WR WKRVH REWDLQHGIURPWKH VWXG\RIFKHPRVWDWVRIEscherichia coli DQG 3KDJH 7 /HYLQ HW DO /HQVNL DQG /HYLQ DQG VXJJHVWHG WKDW D FRPELQDWLRQ RI GLIIHULQJJURZWKUDWHV DQGSKDJHVXVFHSWLELOLWLHVZHUH UHVSRQVLEOH IRU WKH FRH[LVWHQFH RI F\DQRSKDJHV DQG F\DQREDFWHULD ,Q DGGLWLRQ WR WKH KRVW UDQJH RI VSHFLILF F\DQRSKDJHVLWLV QHFHVVDU\WR DOVRFRQVLGHUKRVWFHOO UHVLVWDQFH :LGH YDULDWLRQV LQ WKH H[WHQW WR ZKLFK VSHFLILF VWUDLQV RI F\DQREDFWHULD DUH VXVFHSWLEOH WR LQIHFWLRQ E\ F\DQRSKDJHV ZDV GHPRQVWUDWHG PRVW FOHDUO\ IRU YLUXVHV WKDW LQIHFWHG PDULQH Synechococcus VSS 7LWHUV RIYLUXVHVWKDWLQIHFWVRPH VWUDLQV RISynechococcus FDQ UHDFK WR P/ ZKHUHDV WLWHUV RI YLUXVHV ZKLFK LQIHFW RWKHU VWUDLQV YDU\ IURP XQGHWHFWDEOH WR D IHZ P/ 6XWWOH DQG &KDQ :DWHUEXU\ DQG 9DORLV
([FOXVLYHO\ WKH LVRODWHV ZKLFK DUH PRVW VHQVLWLYH WR LQIHFWLRQ E\ D ZLGH UDQJH RIF\DQRSKDJHV EHORQJ WR WKH SK\FRHU\WKULQULFK JURXS RI Synechococcus VSS WKDW ZHUH WHUPHG 0DULQHFOXVWHU $ :DWHUEXU\ DQG 5LSSND ,Q FRQWUDVW SK\FRF\DQLQGRPLQDQW LVRODWHV ZHUH UHVLVWDQW WR LQIHFWLRQ E\ PRVW F\DQRSKDJHVLQQDWXUDOZDWHUV 3K\FRHU\WKULQULFK Synechococcus DUH E\ IDU WKH PRVW DEXQGDQW XQLFHOOXODU F\DQREDFWHULD LQ ROLJRWURSKLF RFHDQLF ZDWHUV :DWHUEXU\ HW DO DVZHOO DVLQ PDQ\ROLJRWURSKLF ODUJHODNHV &DURQ HW DO 6WRFNQHU DQG 6KRUWUHHG 3K\FRF\DQLQGRPLQDQW Synechococcus DUH UHVWULFWHG WR SURGXFWLYH ZDWHUV 3K\FRHU\WKULQULFK FHOOV DUH SKRWRV\QWKHWLFDOO\ PXFK PRUH HIILFLHQW WKDQ WKRVH ZLWK SK\FRF\DQLQ DV WKHLU GRPLQDQW SLJPHQW XQGHU WKH VKRUWHU ZDYHOHQJWKV FKDUDFWHULVWLF RIROLJRWURSKLF ZDWHU /HZLV HW DO $OVR WKH DEXQGDQFHV RI Synechococcus DQG LQIHFWLRXV F\DQRSKDJHV GHFUHDVH DV RQH PRYHV RIIVKRUH 6XWWOH DQG &KDQ WKHUHIRUH WKH FRQWDFW IUHTXHQF\ EHWZHHQ Synechococcus DQG F\DQRSKDJHV DOVR GHFUHDVHV 7KH ORZHU FRQWDFW IUHTXHQF\ PHDQV WKDW WKH SRWHQWLDO PRUWDOLW\ IURP YLUDO LQIHFWLRQ LV GHFUHDVHG UHODWLYHWR RWKHU IDFWRUV VXFK DV JUD]LQJ DQG UHVXOWV LQ OHVV VHOHFWLYH SUHVVXUH IRU F\DQREDFWHULD WR EH UHVLVWDQW WR LQIHFWLRQ
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lectonema boryanum 87(; LQ OLWHU SRQGV E\ WKH DGGLWLRQ RI &\DQRSKDJH /33 'HVMDUGLQV DQG 2OVRQ 7KH H[SHULPHQWV VKRZHG WKDW /33 FRXOG FRQWURO P. boryanum, EXW WKDWZDWHUTXDOLW\GLGQRWQHFHVVDULO\LPSURYHDVRWKHU
SK\WRSODQNWRQ LQFOXGLQJ F\DQREDFWHULD XOWLPDWHO\ GRPLQDWHG WKH SRQGV ,Q RWKHU SRQGV WKH SUHVHQFH RI F\DQRSKDJHV DSSHDUHG WR SUHFOXGH WKH HVWDEOLVKPHQW RIDEORRP ([SHULPHQWV ZLWK QDWXUDOO\ RFFXUULQJ F\DQREDFWHULDO EORRPV DOVR GHPRQVWUDWHG WKDW WKH DEXQGDQFH RI F\DQREDFWHULD FDQ EH UHGXFHG E\ WKH DGGLWLRQ RI F\DQRSKDJHV DQG EDFWHULDO SDWKRJHQV 0DUWLQ DQG %HQVRQ ,Q OLWHU HQFORVXUH H[SHULPHQWV YLUXVHV DQG RWKHU SDWKRJHQV RI F\DQREDFWHULD ZHUH DGGHG WR QDWXUDOO\ RFFXUULQJ EORRPV DQG WKLV UHVXOWHG LQ D VXEVWDQWLDO GHFUHDVH LQ F\DQREDFWHULDO ELRPDVV (YHQ WKH DGGLWLRQ RI F\DQRSKDJHV DORQH ZDV UHSRUWHG WR FDXVH D FRQVLGHUDEOHGHFUHDVHLQF\DQREDFWHULD 7KHSUHVHQFH RIYLUXVOLNH SDUWLFOHV ZLWKLQAphanizomenon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
9,, (QYLURQPHQWDODQG 3K\VLRORJLFDO (IIHFWVRQ&\DQRSKDJHV 7KHUH DUH D QXPEHU RI HQYLURQPHQWDO DQG SK\VLRORJLFDO FRQVLGHUDWLRQV WKDW DIIHFW WKH VWDELOLW\ DWWDFKPHQWDQGUHSOLFDWLRQRIF\DQRSKDJHVDQGZKLFK DUH FRQVHTXHQWO\ LPSRUWDQW IRU WKHLU SHUVLVWHQFH LQ QDWXUH9LUXVHVYDU\JUHDWO\LQWHUPVRIWKHLUVWDELOLW\ ZKLFK PDNHV LW GLIILFXOW WR PDNH JHQHUDOL]DWLRQV 1RQHWKHOHVVPDQ\F\DQRSKDJHVZHUHVKRZQWRUHWDLQ WKHLU LQIHFWLYLW\ RYHU D ZLGH UDQJH RIHQYLURQPHQWDO FRQGLWLRQV DOWKRXJK PRVW RI WKHVH VWXGLHV ZHUH
&$6XWWOH FRQGXFWHG ZLWK F\DQRSKDJH LVRODWHV UDWKHU WKDQ QDWXUDO F\DQRSKDJH FRPPXQLWLHV 0RUHRYHU PRVW RI WKHVH VWXGLHV ZHUH FRQGXFWHG RQ WLPH VFDOHV RI PLQXWHV WR KRXUV UDWKHU WKDQ GD\V WR ZHHNV PDNLQJ H[WUDSRODWLRQWRWKHQDWXUDOHQYLURQPHQWGLIILFXOW
$ (QYLURQPHQWDO6WDELOLW\RI,QIHFWLRXV &\DQRSKDJHV 7HPSHUDWXUH LV D NH\ HQYLURQPHQWDO YDULDEOH WKDW HIIHFWV LQIHFWLYLW\ 6WXGLHV RQ F\DQRSKDJH LVRODWHV UHYHDOHG WKDW YLUXVHV FDQ JHQHUDOO\ EH VWRUHG IRU D PRQWK RU PRUH DW & ZLWK OLWWOH ORVV RI WLWHU 6DIIHUPDQ DQG 0RUULV E 6DIIHUPDQ HW DO E DOWKRXJK IUHH]LQJ FDQ KDYH D ODUJH HIIHFW 'HVMDUGLQV DQG 2OVRQ ,Q VHDZDWHU VDPSOHV ZLWK SDUWLFXODWH PDWHULDO !P UHPRYHG F\DQRSKDJH WLWHUV ZHUH VWDEOH IRU VHYHUDO PRQWKV ZKHQ VWRUHG DW & 5RGGD DQG 6XWWOH XQSXEOLVKHG GDWD DQG WLWHUV LQH[FHVV RI P/ ZHUH PHDVXUHG LQ P ILOWHUHG VDPSOHV VWRUHG D \HDU RU PRUH 6XWWOH DQG &KDQ $W KLJKHU WHPSHUDWXUHV WLWHUV RI VRPH F\DQRSKDJH LVRODWHV ZHUH VWDEOH DQ KRXURUPRUH DW& 6DIIHUPDQ DQG0RUULV E 6DIIHUPDQ HW DO E ZKLOH RWKHU WKHUPRVHQVLWLYH VWUDLQV ZHUH UHSRUWHGWREHXQDEOHWRIRUPSODTXHVDW & 3DGDQ HW DO $W WHPSHUDWXUHV DERYH & LQIHFWLYLW\ ZDV W\SLFDOO\UDSLGO\ORVW 6DIIHUPDQ DQG 0RUULV E 3DGDQ HW DO 6DUPD DQG 6LQJK 7KH RQO\ GDWD RQ QDWXUDO F\DQRSKDJH FRPPXQLWLHV LQGLFDWHG WKDW GHFD\ UDWHV RILQIHFWLYLW\ LQXQWUHDWHGVHDZDWHUYDULHGIURPFD G EHWZHHQ DQG & WR DQG G DW DQG & UHVSHFWLYHO\ *DU]D *DU]D DQG 6XWWOH LQ UHYLHZ 7KH LQIHFWLYLW\ RIQDWXUDO F\DQRSKDJH FRPPXQLWLHV LV DOVR YHU\ VHQVLWLYH WR VRODUUDGLDWLRQ DQG WKLV FDQ EHWKHPDMRUORVVSURFHVVLQQDWXUDOZDWHUV6XWWOHDQG &KDQ *DU]D *DU]D DQG 6XWWOH LQ UHYLHZ )RU QDWXUDO F\DQRSKDJH FRPPXQLWLHV DWWZR VWDWLRQV RQ WKH FRQWLQHQWDO VKHOI RI WKH *XOI RI 0H[LFR VXQOLJKWZDVUHVSRQVLEOHIRUUHPRYDOUDWHV RI F\DQRSKDJH LQIHFWLYLW\ RI DQG G ZKHQ LQWHJUDWHG RYHUWKH VXUIDFHPL[HG OD\HU DQG DYHUDJHG RYHU K 6HFWLRQ ,9 &OHDUO\ WKH HIIHFW RI VRODU UDGLDWLRQ RQ WKH VXUYLYDO RI F\DQRSKDJHV LQ QDWXUDO ZDWHUV GHSHQGV VWURQJO\ RQ RWKHU HQYLURQPHQWDO IDFWRUV VXFK DV ZDWHU WUDQVSDUHQF\ VRODU DQJOH DQG LQVRODWLRQ 6LPLODU WR RWKHU SKDJHV WKHUH LV QR FRUUHODWLRQ EHWZHHQF\DQRSKDJHPRUSKRORJ\DQGWKHHIIHFWRIS+ RQ VWDELOLW\ DOWKRXJK PDQ\ IUHVKZDWHU F\DQRSKDJHV
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
% $GVRUSWLRQ $OWKRXJK VRPH F\DQREDFWHULD KDYH VWUDWHJLHV WR UHGXFH WKH IUHTXHQF\ ZLWK ZKLFK FRQWDFW ZLWK F\DQRSKDJHV UHVXOW LQ LQIHFWLRQ VHH DERYH LQ JHQHUDO PRVW FRQWDFWV EHWZHHQ EDFWHULRSKDJHV DQG KRVW FHOOV DUH WKRXJKW WR UHVXOW LQ DGVRUSWLRQ $QGHUVRQ 6FKZDUW] 7KLV LV DOVR WUXH IRU PDQ\ F\DQRSKDJHV )RU H[DPSOH ZKHQ [ P/ RI&\DQRSKDJH 6%%6 ZHUH DGGHG WR [ P/ RI H[SRQHQWLDOO\ JURZLQJ Synechococcus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
IRU&\DQRSKDJH630IRUZKLFKDGVRUSWLRQNLQHWLFV ZHUH WKH VDPH RQ SKRVSKDWHUHSOHWH RU GHSOHWH Synechococcus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
& 5HSOLFDWLRQ /LJKW ,QWXLWLYHO\ UHSOLFDWLRQ VKRXOG EH VHQVLWLYH WR HQYLURQPHQWDO HIIHFWV EHFDXVH DQ\WKLQJ WKDW DIIHFWV KRVWFHOO SK\VLRORJ\ SRWHQWLDOO\ DIIHFWV YLUDO UHSOLFDWLRQ )RU H[DPSOH $73 LV UHTXLUHG IRU YLUDO UHSOLFDWLRQ 3DGDQ HW DO 6KHUPDQ DQG +DVHONRUQ $GROSKDQG +DVHONRUQ DQG FRQVHTXHQWO\ F\DQRSKDJH SURGXFWLRQ LV FXUWDLOHG LQ GDUNQHVV ,I Plectonema boryanum 87(; LV NHSWLQWKHGDUNWKURXJKRXWWKHSHULRG RIF\DQRSKDJH JURZWK WKHUH LV D ODJ LQ WKH DSSHDUDQFH RIIUHH YLUDO SDUWLFOHV DQG WKHEXUVW VL]HLV JUHDWO\ UHGXFHG3DGDQ HW DO 6KHUPDQ DQG +DVHONRUQ ,I LQIHFWHG FHOOV DUH UHWXUQHG WR WKH OLJKW HYHQ DIWHU PDQ\ KRXUV RI GDUNQHVV YLUDO UHSOLFDWLRQ SURFHHGV QRUPDOO\ DOWKRXJK WKH ILQDO EXUVW VL]H FDQ EH HYHQ ODUJHU ,Q FRQWUDVW LQ WKH QLWURJHQIL[LQJ F\DQREDFWHULXP Nostoc muscorum $GROSK DQG +DVHONRUQ DQG LQ XQLFHOOXODU F\DQREDFWHULD 0DF.HQ]LH DQG +DVHONRUQ F $OOHQ DQG
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ynechococcus VSZDV PLVLGHQWLILHG DV Microcystis aeruginosa LQ D QXPEHU
&KDSWHU &\DQRSKDJHV RI HDUOLHU VWXGLHV VHH Host Range ,W LV DOVR LQWHUHVWLQJ WKDW XOWUDVWUXFWXUDO FKDQJHV ZHUH IRXQG WR RFFXU LQ XQLFHOOXODU DQG ILODPHQWRXV F\DQREDFWHULD DIWHU LQIHFWLRQ ,Q ILODPHQWRXV F\DQREDFWHULD GLVSODFHPHQW RI WKH SKRWRV\QWKHWLF ODPHOODH ZDV DVVRFLDWHG ZLWK WKH SURGXFWLRQ RI YLUDO SDUWLFOHV 6PLWKHWDO ZKHUHDVLQWKHXQLFHOOXODUIRUPV WKHUH ZHUH QR REYLRXV FKDQJHV LQ WKH WK\ODNRLGV 6KHUPDQHWDO
1XWULHQWV 2EYLRXVO\SURWHLQDQG'1$V\QWKHVLVDUHDQLQWHJUDO SDUW RI YLUDO UHSOLFDWLRQ VR LW LV UHDVRQDEOH WR DQWLFLSDWH WKDW XQGHU WKH FRQGLWLRQV RI QLWURJHQ RU SKRVSKRUXV OLPLWDWLRQ DV H[LVW LQ PDQ\ QDWXUDO ZDWHUV YLUDO UHSOLFDWLRQ ZLOO EH DIIHFWHG
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
9,,, /\VRJHQ\ $ )LODPHQWRXV &\DQREDFWHULD %\ WKH HDUO\ V WKH H[LVWHQFH RI O\VRJHQLF DVVRFLDWLRQV EHWZHHQ F\DQRSKDJHV DQG ILODPHQWRXV F\DQREDFWHULD ZHUH ZHOO HVWDEOLVKHG HJ &DQQRQ HW DO 3DGDQHWDO 6LQJKDQG6LQJK .KXG\DNRY DQG *URPRY 5LPRQ DQG 2SSHQKHLP DOWKRXJK ZKHWKHU O\VRJHQ\ ZDV WUXO\ GHPRQVWUDWHG LQ VRPH RI WKH HDUO\ VWXGLHV 6KHUPDQ DQG %URZQ LV D PDWWHU RI GHEDWH 0XFK RI WKH HDUOLHU ZRUN IRFXVHG RQ O\VRJHQV RI Plectonema boryanum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
% 8QLFHOOXODU&\DQREDFWHULD ,Q FRQWUDVW WR WKH ZHDOWK RI VWXGLHV RQ WHPSHUDWH F\DQRSKDJHV RI ILODPHQWRXV F\DQREDFWHULD GDWD RQ O\VRJHQV RI XQLFHOOXODU F\DQREDFWHULD ZHUH VORZ WR PDWHULDOL]H ,Q IDFW ZLWK WKH H[FHSWLRQ RI VRPH SUHOLPLQDU\ HYLGHQFH IRU O\VRJHQ\ LQ Anacystis
nidulans %LVHQ HW DO O\VRJHQV ZKLFK LQIHFW XQLFHOOXODU F\DQREDFWHULD UHPDLQHG XQGRFXPHQWHG XQWLO WKH LVRODWLRQ RI D WHPSHUDWH F\DQRSKDJH ZKLFK LQIHFWHG PDULQH Synechococcus VS VWUDLQ 1.%* 6RGHHWDO
& 6LJQLILFDQFH &RQVLGHUDWLRQ RI WKH HFRORJLFDO DQG HYROXWLRQDU\ VLJQLILFDQFH RI O\VRJHQ\ LQ F\DQRSKDJHV ZDV QRW DGGUHVVHG +RZHYHU JLYHQ WKH LQFUHDVHG UHFRJQLWLRQ RI WKH LPSRUWDQFH RI F\DQREDFWHULD LQFOXGLQJ SURFKORURSK\WHV LQWKHZRUOG VRFHDQVDQGODNHV DQG WKHUHQHZHGLQWHUHVWLQWKHUROHRIF\DQRSKDJHV LQWKH HQYLURQPHQWLWLVDQDUHDWKDWLVULSHIRULQYHVWLJDWLRQ ,Q SDUWLFXODU O\VRJHQLF F\DQRSKDJHV PD\ EH LPSRUWDQW DV YHFWRUV RI JHQHWLF LQIRUPDWLRQ ZLWKLQ F\DQREDFWHULDO FRPPXQLWLHV $OWKRXJK WKH PRYHPHQW RIF\DQREDFWHULDO'1$E\ YLUXVHV WUDQVGXFWLRQ ZDV QRW GHPRQVWUDWHG WKHUH LV FOHDU HYLGHQFH WKDW O\VRJHQLFF\DQRSKDJHVLQWHJUDWHLQWRWKHKRVWJHQRPH HJ )UDQFKH 7KH WULJJHUV WKDW LQGXFH SURSKDJHV LQ F\DQREDFWHULD WR HQWHU WKH O\WLF F\FOH YDU\ )RU VRPH O\VRJHQL]HG F\DQREDFWHULD HQYLURQPHQWDO IDFWRUV LQFOXGLQJ 895 2KNL DQG )XMLWD WHPSHUDWXUH 5LPRQ DQG 2SSHQKHLP RU H[SRVXUH WR KLJK OHYHOV RIFRSSHU 6RGH HW DO WULJJHU LQGXFWLRQ ZKLOH LQ RWKHU FDVHV DWWHPSWV DW LQGXFWLRQ ZHUH XQVXFFHVVIXO HYHQ ZLWK PXWDJHQV VXFK DV PLWRP\FLQ & HJ 3DGDQ HW DO $W WKH SUHVHQW WLPH WKHUH DUH QR HVWLPDWHV RI WKH SURSRUWLRQ RI F\DQREDFWHULD WKDW DUH O\VRJHQL]HG LQ QDWXUH KRZHYHU HVWLPDWHV LQGLFDWH WKDW D UHODWLYHO\ VPDOO SURSRUWLRQ RI WKH HQWLUH PDULQH EDFWHULDO FRPPXQLW\ PD\ EH O\VRJHQL]HG -LDQJ DQG 3DXO ZHUH XQDEOH WR GHWHFW O\VRJHQLF YLUDO SURGXFWLRQLQDOOEXWDIHZRI VDPSOHVFROOHFWHGLQ ROLJRWURSKLF RIIVKRUH HQYLURQPHQWV EXW REWDLQHG HYLGHQFHRILQGXFWLRQDWRI HXWURSKLFVWDWLRQV,Q D VLPLODU VWXG\ :HLQEDXHU DQG 6XWWOH XVHG D PRUH VHQVLWLYH DSSURDFK IRU WKH GHWHFWLRQ RI SKDJH SURGXFWLRQ LQ VDPSOHV IURP WKH *XOI RI 0H[LFR 7KH\ HVWLPDWHG WKDW WR DYHUDJH RI WKH QDWXUDOEDFWHULDO FRPPXQLW\FRXOG EH LQGXFHG E\ WKH DGGLWLRQ RI0LWRP\FLQ & 7KHVH VWXGLHV VKRZHG WKDW D VPDOO SURSRUWLRQ RI PDULQH EDFWHULDO FRPPXQLWLHV DV DZKROH ZHUH LQGXFHGE\ PLWRP\FLQ & RU 89& UDGLDWLRQ VR LW LV UHDVRQDEOH WR VXSSRVH WKDW F\DQREDFWHULD UHVSRQG LQ D VLPLODU ZD\ 7KH JUHDWHVWGLIILFXOW\ZLWKLQWHUSUHWLQJWKHVHH[SHULPHQWV LQWKHFRQWH[WRIQDWXUDOPLFURELDOFRPPXQLWLHVLVWKH
&$6XWWOH HIIHFWLYHQHVV RI WKH LQGXFLQJ DJHQWV LQ VWLPXODWLQJ SKDJH SURGXFWLRQ LQ O\VRJHQV 0LWRP\FLQ & RU 89& UDGLDWLRQ DUH QRW HQYLURQPHQWDOO\ UHOHYDQW DQG PD\ QRWEHHIIHFWLYHLQGXFLQJDJHQWV ,Q WKH QDWXUDO HQYLURQPHQW LW LV QRW NQRZQ ZKDW IDFWRUV DUH LPSRUWDQW IRU WKH HVWDEOLVKPHQW RI D O\VRJHQLF DVVRFLDWLRQ LQ F\DQREDFWHULD QRU ZKDW WULJJHUV O\VRJHQV WR HQWHU WKH O\WLF SDWKZD\ 2QH SRVVLELOLW\ LV WKDW QXWULHQWV PD\ EH LQYROYHG :LOVRQ HW DO UHSRUWHG WKDW O\VLV RFFXUUHG LQ RQO\ RIWKH FHOOV ZKHQ LQIHFWLRXV F\DQRSKDJHV ZHUH DGGHG WR Synechococcus JURZQ XQGHU ORZ SKRVSKDWH FRQGLWLRQV ,QFRQWUDVW O\VLV RFFXUUHGLQ DOOFHOOVWKDW ZHUH JURZQ XQGHUSKRVSKDWH UHSOHWH FRQGLWLRQV 2QH LQWHUSUHWDWLRQ RI WKLV UHVXOW LV WKDW ORZ SKRVSKDWH FRQGLWLRQVUHVXOWHGLQWKHHVWDEOLVKPHQWRIDO\VRJHQLF DVVRFLDWLRQ KRZHYHU WKLV FRQFOXVLRQ DVVXPHV WKDW ORZ SKRVSKDWH FRQFHQWUDWLRQV GR QRW DIIHFW FHOO YLDELOLW\ )URP DQ HYROXWLRQDU\ SHUVSHFWLYH LW PDNHV VHQVHWKDWWKHQXWULWLRQDO VWDWXVRIWKHKRVWPD\ DIIHFW WKHHVWDEOLVKPHQWRIO\VRJHQ\
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
&KDSWHU &\DQRSKDJHV ZHUH QXPHURXV DGYDQFHV LQ UHFHQW \HDUV IRU WHFKQLTXHV WR LVRODWH DQG FXOWXUH F\DQRSKDJHV IURP WKHHQYLURQPHQW $OORIWKHVH IDFWRUV VXJJHVWWKDWWKH HFRORJLFDO LPSOLFDWLRQV RI F\DQRSKDJH LQIHFWLRQ VKRXOGEHDIUXLWIXODUHDIRUUHVHDUFKLQWKHIXWXUH
$FNQRZOHGJPHQWV , DP LQGHEWHG WR WKH LQGLYLGXDOV ZKR RYHUWKH \HDUV PDGHP\ODERUDWRU\DQH[FLWLQJHQYLURQPHQWWRVWXG\ DQG WKLQN DERXW YLUXVHV DQG WKHLU UROH LQ DTXDWLF HQYLURQPHQWV ,Q SDUWLFXODU WKH KDUG ZRUN DQG LQVLJKWV RI )HQJ &KHQ 0DWWKHZ &RWWUHOO +XGVRQ 'H
5HIHUHQFHV $FNHUPDQQ+:DQG'X%RZ06 D 9LUXVHV RI3URNDU\RWHV 9ROXPH 1DWXUDO *URXSV RI %DFWHULRSKDJHV &5& 3UHVV %RFD5DWRQ)ORULGD $FNHUPDQQ+:DQG 'X%RZ06 E 9LUXVHVRI3URNDU\RWHV 9ROXPH *HQHUDO 3URSHUWLHV RI%DFWHULRSKDJHV &5& 3UHVV %RFD5DWRQ)ORULGD $GDPV0+ %DFWHULRSKDJHV-RKQ:LOH\DQG6RQV1< $GROSK.:DQG+DVHNRUQ5 ,VRODWLRQDQGFKDUDFWHUL]DWLRQ RI D YLUXV LQIHFWLQJ WKH EOXHJUHHQ DOJD Nostoc muscorum. 9LURORJ\ $GROSK .: DQG +DVHONRUQ 5 3KRWRV\QWKHVLV DQG WKH GHYHORSPHQW RI WKH EOXHJUHHQ DOJDO YLUXV 1 9LURORJ\ $GROSK .: DQG +DVHONRUQ 5 D ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RI D YLUXV LQIHFWLQJ D EOXHJUHHQ DOJD RI WKH JHQXV Synechococcus. 9LURORJ\ $GROSK.: DQG+DVHONRUQ5E %OXHJUHHQDOJDOYLUXV 1, 3K\VLFDO SURSHUWLHV DQG GLVDVVHPEO\ LQWR VWUXFWXUDO SDUWV 9LURORJ\ $OOHQ00DQG+XWFKLQVRQ) (IIHFWRIVRPHHQYLURQPHQWDO IDFWRUVRQF\DQRSKDJH$6GHYHORSPHQWLQAnacystis QLGXODQV $UFK0LFURELRO $QGHUVRQ7) 7KHUHDFWLRQV RIYLUXVHVZLWKWKHLUKRVWFHOOV %RW5HY $POD '9 3KRWRUHDFWLYDWLRQ RIXOWUDYLROHW LUUDGLDWHG EOXH JUHHQ DOJD Anacystis nidulans DQG &\DQRSKDJH $6 $UFK 9LURO
$POD '9 &KHODWLQJ DJHQW VKRFN RI F\DQRSKDJH $6 LQIHFWLQJ XQLFHOOXODU EOXHJUHHQ DOJDH ,QGLDQ - ([S %LRO $POD'9 5RZHOO3DQG 6WHZDUW:'3 0HWDEROLF FKDQJHV DVVRFLDWHGZLWKF\DQRSKDJH1 LQIHFWLRQRIWKHF\DQREDFWHULXP Nostoc muscorum. $UFK0LFURELRO $VDWR < 8OWUDYLROHW OLJKW LQDFWLYDWLRQ DQG SKRWRUHDFWLYDWLRQ RI$6 F\DQRSKDJH LQ Anacysitis nidulans. %DFWHULRO %DJFKL 61 .DOR\D3 DQG %LVHQ 36 (IIHFW RIF\DQRSKDJH 1O LQIHFWLRQRQWKHV\QWKHVLV DQG VWDELOLW\RINostoc muscorum QLWUDWHUHGXFWDVH&XUU0LFURELRO %DQFURIW , DQG 6PLWK 5- 5HVWULFWLRQ PDSSLQJ RIJHQRPLF '1$ IURP F\DQRSKDJHV LQIHFWLQJ WKH KHWHURF\VWRXV F\DQREDFWHULD Nostoc DQG Anabaena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nacysris nidulans. 0LFURELRV/HWW %LVHQ 36 $XGKROLD 6 %KDWQDJDU $. DQG %DJFKL 61 E (YLGHQFH IRU O\VRJHQ\ DQG YLUDO UHVLVWDQFH LQ WKH F\DQREDFWHULXP Phormidium uncinatum. &XUU0LFURELRO %LVHQ36 $XGKROLD66KXNOD+' *XSWD$DQG6LQJK '3 (YLGHQFH IRU SKRWRV\QWKHWLF LQGHSHQGHQFH RI YLUDO PXOWLSOLFDWLRQ LQ F\DQRSKDJH/33 LQIHFWHG &\DQREDFWHULXP Phormidium uncinatum. )(060LFURELRO/HWW %LVHQ 36 %DJFKL 61 DQG $XGKROLD 6 D 1LWUDWH UHGXFWDVH DFWLYLW\ RI D F\DQREDFWHULXP Phormidium uncinatum DIWHU F\DQRSKDJH/33 LQIHFWLRQ)(060LFURELRO/HWW %RUEp\*.DUL&*XO\DV$DQG)DUNDV*/ %DFWHULRSKDJH LQIHFWLRQ LQWHUIHUHV ZLWK TXDQRVLQH GLSKRVSKDWH GLSKRVSKDWH DFFXPXODWLRQ LQGXFHG E\ HQHUJ\ DQG QLWURJHQ VWDUYDWLRQ LQ F\DQREDFWHULXP Anacystis nidulans. - %DFWHULRO %UDLWKZDLWH '. DQG ,WR - &RPSLODWLRQ DOLJQPHQW DQG SK\ORJHQHWLF UHODWLRQVKLSV RI'1$ SRO\PHUDVHV 1XFOHLF $FLGV 5HV %URZQ50-U $OJDO9LUXVHV$GY9LUXV5HV &DQQRQ 5( 6KDQH06DQG %XVK 91 /\VRJHQ\RIDEOXH JUHHQDOJDPlectonema boryanum. 9LURORJ\ &DQQRQ 5( 6KDQH 06 DQG :KLWDNHU -0 ,QWHUDFWLRQ RI Plectonemu boryanum &\DQRSK\FHDH DQG WKH /33 F\DQRSKDJHVLQFRQWLQXRXVFXOWXUH-3K\FRO &DURQ '$ 3LFN )5 DQG /HDQ '56 &KURRFRFFRLG F\DQREDFWHULD LQ /DNH 2QWDULR YHUWLFDO DQG VHDVRQDO GLVWULEXWLRQVGXULQJ -3K\FRO
&DUSHQWHU(- 3K\VLRORJ\DQGHFRORJ\RIPDULQHSODQNWRQLF Oscillatoria (Trichodesmium). 0DU%LRO/HWW &DUSHQWHU (- DQG 5RPDQV . 0DMRU UROH RI WKH F\DQREDFWHULXP Trichodesmium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icromonas pusilla. /LPQRO2FHDQRJU &RXORPEH $0 DQG 5RELQVRQ **& &ROODSVLQJ Aphanizomenon flos-aquae EORRPV 3RVVLEOH FRQWULEXWLRQV RI SKRWRR[LGDWLRQ 2 WR[LFLW\ DQG F\DQRSKDJHV &DQ - %RW &RZLH '% DQG 3UDJHU / &\DQRSK\WD DQG WKHLU YLUXVHV &DPHJLH ,QVWLWXWLRQ RI :DVKLQJWRQ
&$6XWWOH *DU]D'5DQG6XWWOH&$LQUHYLHZ 7KHHIIHFWRIF\DQRSKDJHVRQ WKHPRUWDOLW\RISynechococcus VSSDQGVHDVRQDOFKDQJHVLQWKH UHVLVWDQFHRIQDWXUDO YLUDOFRPPXQLWLHVWR89UDGLDWLRQ *LQ]EHUJ ' 3DGDQ ( DQG 6KLOR 0 (IIHFW RIF\DQRSKDJH LQIHFWLRQ RQ &2 SKRWRDVVLPLODWLRQ LQ Plectonema boryanum. 9LURO *LQ]EHUJ ' 3DGDQ ( DQG 6KLOR 0 0HWDEROLF DVSHFWV RI /33 F\DQRSKDJH UHSOLFDWLRQ LQ WKH F\DQREDFWHULXP Plectonema boryanum. %LRFKLP%LRSK\V $FWD *ROGVWHLQ'$%HQGHW,-/DXIIHU0$DQG6PLWK.0 6RPH ELRORJLFDO DQG SK\VLFRFKHPLFDO SURSHUWLHV RI EOXHJUHHQ DOJDO YLUXV/339LURORJ\ *RQ]iOH] -0 DQG 6XWWOH &$ *UD]LQJ E\ PDULQH QDQRIODJHOODWHV RQ YLUXVHV DQG YLUXVVL]HG SDUWLFOHV ,QJHVWLRQ DQGGLJHVWLRQ0DU(FRO3URJU6HU *URPRY%9 &\DQRSKDJHV$QQ0LFURELRO +HOGDO 0 DQG %UDWEDN * 3URGXFWLRQ DQG GHFD\ RIYLUXVHV LQDTXDWLFHQYLURQPHQWV0DU(FRO3URJU6HU +HQQHV.3&KDQ$0DQG6XWWOH&$ )OXRUHVFHQWO\ODEHOHG YLUXV SUREHV VKRZ WKDWQDWXUDOYLUXVSRSXODWLRQV FDQFRQWURO WKH VWUXFWXUH RI PDULQH PLFURELDO FRPPXQLWLHV $SSO (QYLURQ 0LFURELRO +RUQH $- DQG *ROGPDQ &5 /LPQRORJ\ 0F*UDZ+LOO 1HZ
&KDSWHU &\DQRSKDJHV .LP 0 DQG &KRL <. $ QHZ Synechococcus F\DQRSKDJH IURPDUHVHUYRLULQ.RUHD9LURORJ\ .R]\DNRY6
0LOOHU59DQG.RNMRKQ7$ *HQHUDOPLFURELRORJ\RIrec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yngbya wollei, Anabaena flosaquae DQG Anabaena circinalis. - $TXDW 3ODQW 0DQDJH 0XUDGRY 00 &KHUNDVRYD *9 $NKPHGRYD '8 .DPLORYD )' 0XNKDPHGRY 56 $EGXNDULPRY $$ DQG .KDOPXUDGRY $* &RPSDUDWLYH VWXG\ RI 13,7 F\DQRSKDJHV ZKLFK O\VRJHQL]H QLWURJHQIL[LQJ EDFWHULD RI WKH JHQHUD Nostoc DQG Plectonema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hormidium persicinurn. -3K\FRO 2OVRQ5- &KLVKROP6:=HWWOHU(5 $OWDEHW0$DQG'XVHQEHUU\ -$ 6SDWLDO DQG WHPSRUDO GLVWULEXWLRQV RISURFKORURSK\WH SLFRSODQNWRQ LQ WKH 1RUWK $WODQWLF 2FHDQ 'HHS 6HD 5HV 3DGDQ ( *LQ]EXUJ' DQG6KLOR 0 7KH UHSURGXFWLYH F\FOH RI F\DQRSKDJH /33,* LQ Plectonema boryanum DQG LWV GHSHQGHQFHRQSKRWRV\QWKHWLFDQGUHVSLUDWRU\V\VWHPV 9LURORJ\ 3DGDQ ( 5LPRQ $ *LQ]EHUJ ' DQG 6KLOR 0 $ WKHUPRVHQVLWLYH F\DQRSKDJH /33* DWWDFNLQJ WKH EOXHJUHHQ DOJD Plectonema boryanum. 9LURORJ\ 3DGDQ ( DQG 6KLOR 0 'LVWULEXWLRQ RI F\DQRSKDJHV LQ QDWXUDOKDELWDWV9HUK,QW9HUHLQ/LPQRO 3DGDQ(DQG6KLOR0 &\DQRSKDJHVYLUXVHVDWWDFNLQJEOXH JUHHQDOJDH%DFWHULRO5HY 3DGDQ ( 6KLOR 0 DQG 2SSHQKHLP $% /\VRJHQ\ RI WKH EOXHJUHHQ DOJD Plectonema boryanum E\ /3363, F\DQRSKDJH9LURORJ\ 3KOLSV (- 0RQHJXH 5/ DQG $OGULGJH )- &\DQRSKDJHV ZKLFK LPSDFW EORRPIRUPLQJ F\DQREDFWHULD - $TXDW 3ODQW 0DQDJH 3URFWRU /0 DQG )XKUPDQ -$ 9LUDO PRUWDOLW\ RI PDULQH EDFWHULDDQGF\DQREDFWHULD1DWXUH 3URFWRU /0 )XKUPDQ -$ DQG /HGEHWWHU 0& 0DULQH EDFWHULRSKDJHVDQGEDFWHULDOPRUWDOLW\(RV 3URFWRU /0 2NXER $ DQG )XKUPDQ -$ &DOLEUDWLQJ HVWLPDWHV RI SKDJHLQGXFHG PRUWDOLW\ LQ PDULQH EDFWHULD
8OWUDVWUXFWXUDO VWXGLHV RI PDULQH EDFWHULRSKDJH GHYHORSPHQW IURPRQHVWHSJURZWKH[SHULPHQWV0LFURE(FRO 5LPRQ $ DQG 2SSHQKHLP $% +HDW LQGXFWLRQ RIWKHEOXH JUHHQ DOJD Plectonemu boryanum O\VRJHQLF IRUWKH F\DQRSKDJH 63,FWV9LURORJ\ 5LSSND 5 DQG +HUGPDQ 0 3DVWHXU &XOWXUH &ROOHFWLRQ RI &\DQREDFWHULD&DWDORJXHDQG7D[RQRPLF+DQGERRN9ROXPH, &DWDORJXHRI6WUDLQV,QVWLWXW3DVWHXU3DULV 5RGGD .0 7HPSRUDO DQG 6SDWLDO '\QDPLFV RI Synechococcus VSS DQGMicromonas pusilla KRVWYLUDO V\VWHPV 0$7KHVLV0DULQH6FLHQFH8QLYHUVLW\RI7H[DVDW$XVWLQ 5RGGD .0 6XWWOH &$ &ODUN // DQG ,QJDOO ( LQ UHYLHZ ,QIHFWLRXV YLUXVHV SHUVLVW LQ PDULQH VHGLPHQWV IRU DW OHDVW \HDUV 6DIIHUPDQ 56 9LUXV GLVHDVHV LQ EOXHJUHHQ DOJDH ,Q -DFNVRQ')HG $OJDH 0DQDQG WKH(QYLURQPHQW SS 6\UDFXVH8QLYHUVLW\3UHVV1HZ
&$6XWWOH 6KHUPDQ /$ ,QIHFWLRQ RISynechococcus cedrorum E\ WKH F\DQRSKDJH $60 ,,, &HOOXODU PHWDEROLVP DQG SKDJH GHYHORSPHQW9LURORJ\ 6KHUPDQ /$ DQG %URZQ 50 -U &\DQRSKDJHV DQG YLUXVHV RI HXNDU\RWLF DOJDH ,Q )UDHQNHO&RQUDW + DQG :DJQHU 55 HGV &RPSUHKHQVLYH 9LURORJ\ SS 3OHQXP 3UHVV 1HZ
&KDSWHU &\DQRSKDJHV 6XWWOH&$ ,QKLELWLRQRISKRWRV\QWKHVLVLQSK\WRSODQNWRQ E\ WKH VXEPLFURQ VL]H IUDFWLRQ FRQFHQWUDWHG IURP VHDZDWHU 0DU (FRO3URJU6HU 6XWWOH &$ 7KH VLJQLILFDQFH RI YLUXVHV WR PRUWDOLW\ LQ DTXDWLF PLFURELDOFRPPXQLWLHV 0LFURE (FRO 6XWWOH &$ DQG &KDQ $0 0DULQH F\DQRSKDJHV LQIHFWLQJ RFHDQLF DQG FRDVWDO VWUDLQV RI Synechococcus: $EXQGDQFH PRUSKRORJ\ FURVVLQIHFWLYLW\ DQG JURZWK FKDUDFWHULVWLFV 0DU (FRO3URJU6HU 6XWWOH &$ DQG &KDQ $0 '\QDPLFV DQG GLVWULEXWLRQ RI F\DQRSKDJHV DQG WKHLU HIIHFW RQ PDULQH Synechococcus VSS $SSO(QYLURQ0LFURELDO 6XWWOH&$&KDQ$0 &KHQ)DQG*DU]D'5 &\DQRSKDJHV DQGVXQOLJKW$SDUDGR[,Q*XHUUHUR5DQG3HGURV$OLR&HGV 7UHQGVLQ0LFURELDO(FRORJ\SS 6SDQ6RF0LFURELRO %DUFHORQD 6XWWOH &$ &KDQ $0 DQG &RWWUHOO 07 ,QIHFWLRQ RI SK\WRSODQNWRQE\YLUXVHV DQGUHGXFWLRQ RISULPDU\SURGXFWLYLW\ 1DWXUH 6XWWOH &$ &KDQ $0 5RGGD .0 6KRUW 60 :HLQEDXHU 0* *DU]D '5 DQG :LOKHOP 6: 7KH HIIHFW RIF\DQRSKDJHV RQ Synechococcus VSS GXULQJ D EORRP LQ WKH ZHVWHUQ *XOIRI 0H[LFR(RVVXSSO 2626 6XWWOH &$ DQG &KHQ ) 0HFKDQLVPV DQG UDWHV RIGHFD\ RI PDULQH YLUXVHV LQ VHDZDWHU $SSO (QYLURQ 0LFURELRO 6XWWOH &$ &KHQ )DQG &KDQ $0 0DULQH YLUXVHV GHFD\ UDWHV GLYHUVLW\ DQG HFRORJLFDO LPSOLFDWLRQV ,Q 1DVK && HG ,QWHUQDWLRQDO0DULQH%LRWHFKQRORJ\&RQIHUHQFH ,0%& SS :%URZQ'XEXTXH 7HNOHPDULDP 7$ 'HPHWHU 6 'HN = 6XUQ\L * DQG %RUEÊO\ * $6 F\DQRSKDJH LQIHFWLRQ LQKLELWV WKH SKRWRV\QWKHWLF HOHFWURQIORZRISKRWRV\VWHPOOLQSynechococcus VS3&& DF\DQREDFWHULXP)(%6/HWW 7XUQHU6 0ROHFXODUV\VWHPDWLFVRIR[\JHQLFSKRWRV\QWKHWLF EDFWHULD 3ODQW6\VWHPDWLFVDQG(YROXWLRQ6 :DWHUEXU\ -% DQG 5LSSND 5 6XEVHFWLRQ , 2UGHU &KURRFRFFDOHV:HWWVWHLQ HPHQG 5LSSNDHWDO ,Q 6WDOH\-7%U\DQW033IHQQLJ1+ROW-%HGV %HUJH\ V0DQXDO RI 6\VWHPDWLF %DFWHULRORJ\ SS :LOOLDPV DQG :LONLQV%DOWLPRUH :DWHUEXU\ -% DQG 9DORLV ): 5HVLVWDQFH WR FRRFFXUULQJ SKDJHV HQDEOHV PDULQH Synechococcus FRPPXQLWLHV WR FRH[LVW ZLWK F\DQRSKDJHV DEXQGDQW LQ VHDZDWHU $SSO (QYLURQ 0LFURELRO :DWHUEXU\ -% :DWVRQ 6: 9DORLV ): DQG )UDQNV '* %LRORJLFDO DQG HFRORJLFDO FKDUDFWHUL]DWLRQ RI WKH PDULQH XQLFHOOXODU F\DQREDFWHULXP Synechococcus. ,Q 3ODWW 7 DQG /L :.: HGV 3KRWRV\QWKHWLF 3LFRSODQNWRQ &DQ %XOO )LVK $TXDW 6FL :HLQEDXHU 0* DQG 6XWWOH &$ 3RWHQWLDO VLJQLILFDQFH RI O\VRJHQ\ to EDFWHULRSKDJHSURGXFWLRQ DQG EDFWHULDO PRUWDOLW\ LQ FRDVWDO ZDWHUV RIWKH *XOIRI0H[LFR $SSO (QYLURQ 0LFURELRO :HLQEDXHU 0* :LOKHOP 6: 6XWWOH &$ DQG *DU]D '5 3KRWRUHDFWLYDWLRQ FRPSHQVDWHV IRU 89 GDPDJH DQG UHVWRUHV LQIHFWLYLW\ WR QDWXUDO PDULQH YLUDO FRPPXQLWLHV $SSO (QYLURQ 0LFURELRO :LJJLQV %$DQG $OH[DQGHU0 0LQLPXPEDFWHULDO GHQVLW\ IRU EDFWHULRSKDJH UHSOLFDWLRQ LPSOLFDWLRQV IRU VLJQLILFDQFH RI EDFWHULRSKDJHV LQ QDWXUDO HFRV\VWHPV $SSO (QYLURQ 0LFURELDO
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nacystis nidulans DQG VRPH RWKHU Synechococcus VWUDLQV - *HQ 0LFURELRO :LOVRQ :+ &DUU 1* DQG 0DQQ 1+ 7KH HIIHFW RI SKRVSKDWH VWDWXV RQ WKH NLQHWLFV RIF\DQRSKDJH LQIHFWLRQ LQ WKH RFHDQLF F\DQREDFWHULXP Synechococcus VS:+ - 3K\FRO :LOVRQ:+-RLQW,5&DUU1*DQG0DQQ1+ ,VRODWLRQDQG PROHFXODU FKDUDFWHUL]DWLRQ RI ILYH PDULQH F\DQRSKDJHV SURSDJDWHGRQSynechococcus VS VWUDLQ:+ $SSO(QYLURQ 0LFURELRO :RHVH&5 %DFWHULDOHYROXWLRQ0LFURELRO5HY :RPPDFN .( +LOO 57 0XOOHU 7$ DQG &ROZHOO 55 (IIHFWV RI VXQOLJKW RQ EDFWHULRSKDJH YLDELOLW\ DQG VWUXFWXUH $SSO(QYLURQ0LFURELRO :X-+/HZLQ5$DQG:HUELQ+ 3KRWRUHDFWLYDWLRQRI89 LUUDGLDWHGEOXHJUHHQDOJDYLUXV/339LURORJ\ :X -+ 6KXJDUPDQ DQG 30 (IIHFW RIYLUXV LQIHFWLRQ UDWH RQ SKRWRV\QWKHVLV DQG UHVSLUDWLRQ RI D EOXHJUHHQ DOJD Plectonema boryanum. 9LURORJ\
&KDSWHU &\DQREDFWHULDO 5HVSRQVHV WR 895DGLDWLRQ 5LFKDUG:&DVWHQKRO] DQG)HUUDQ*DUFLD3LFKHO 'HSW RI%LRORJ\ 8QLYHUVLW\RI2UHJRQ(XJHQH 2UHJRQ 86$ 'HSDUWPHQW RI0LFURELRORJ\$UL]RQD6WDWH8QLYHUVLW\ 7HPSH$UL]RQD86$
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hitton and M Potts (eds), The Ecology of Cyanobacteria, © Kluwer Academic Publishers. Printed in the Netherlands.
5:&DVWHQKRO]DQG )*DUFLD3LFKHO
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
, ,QWURGXFWLRQ $ 7KH'HWULPHQWDO(IIHFWVRI895RQ 0LFURRUJDQLVPV %HFDXVHRILWVSRWHQWLDOWRGLUHFWO\DIIHFWPROHFXOHVRI FHQWUDO ELRORJLFDO LPSRUWDQFH XOWUDYLROHW UDGLDWLRQ 895 LVFRQVLGHUHGGHWULPHQWDOWRDOOOLIHIRUPV%XW EHFDXVH :5 LVSDUWRIWKHWRWDO VRODU VSHFWUXP DWWKH (DUWKV VXUIDFH DYRLGDQFH LV GLIILFXOW $OWKRXJKWKH\ DUH RQO\ D VPDOO SURSRUWLRQ RI WKH WRWDO HQHUJ\ RU SKRWRQIOX[ WKHZDYHOHQJWKV ZKLFK DUHWHUPHG 895 89$ QP 89% QP89& QP EHFRPH SURJUHVVLYHO\ PRUH GHWULPHQWDO DV ZDYHOHQJWK GHFUHDVHV DQG H[SRQHQWLDO LQFUHDVHV LQ SRWHQWLDO GDPDJH WR FHOOV ZRXOG EH H[SHFWHG 9LUWXDOO\ DOO 89& DQG PRVW RI WKH 89% GR QRW FXUUHQWO\ UHDFK WKH (DUWKV VXUIDFH DOWKRXJK WKHVH EDQGV SUHVXPDEO\ SHQHWUDWHG WKH DWPRVSKHUH LQ VLJQLILFDQW DPRXQWV YLUWXDOO\ XQDWWHQXDWHG GXULQJ WKH$UFKHDQDQGHDUO\3URWHUR]RLF(RQVEHIRUHR]RQH DFFXPXODWHGLQWKHVWUDWRVSKHUH.DVWLQJHWDO ,QGHHG LW LV SRVVLEOH WKDW 895 SOD\HG D VLJQLILFDQW UROH GXULQJ WKH HYROXWLRQ RI F\DQREDFWHULD $ SODXVLEOHVFHQDULRIRUWKLVUROHZDVH[DPLQHGUHFHQWO\ *DUFLD3LFKHO 8QGHU YHU\ FOHDU VNLHV DW WHPSHUDWH WR HTXDWRULDO ODWLWXGHV WRWDO 89% WRGD\ PD\EHDV KLJKDV:P DQG89$DVKLJKDV :P DVFRPSDUHGWRYDOXHVXSWRa:P IRUYLVLEOHZDYHOHQJWKVa QP ([FHSWIRUD VPDOO SRUWLRQ RI QHDU 89$ 895 LV QRW SKRWRV\QWKHWLFDOO\ DFWLYH 7KHUHIRUH 895 LV FRQVLGHUHGWREHVRPHWKLQJWRWROHUDWHFRXQWHUDFW RU DYRLG +RZHYHU ERWK89$DQGYLROHWEOXHUDGLDWLRQ DUH SRVLWLYHO\ HIIHFWLYH LQ SLJPHQW V\QWKHVLV DQG LQ SKRWRUHDFWLYDWLRQ RI 89%LQGXFHG GLPHUL]DWLRQ EHWZHHQ DGMDFHQW S\ULPLGLQH EDVHV RI '1$ 2WKHU SRVVLEOH EHQHILFLDO HIIHFWV PD\ EH WKH XVH RI 89$ DQGRU89%DVDFXHLQSKRWRPRWLOLW\UHVSRQVHV
VXFK DV SKRWRSKRELF UHDFWLRQV DQG SKRWRWD[LV VHH ODWHU 6SHFLILF895³SKRWRUHFHSWRUV´DUHXQNQRZQ 7KH QHJDWLYH HIIHFWV RI 895 H[SRVXUH DUH PDQ\ DQG GLYHUVH $W WKH SKHQRPHQRQ OHYHO SKRWRV\QWKHVLV JURZWK PRWLOLW\ SKRWRPRYHPHQWV FHOO GLIIHUHQWLDWLRQ DQG RWKHU SURFHVVHV DUH DIIHFWHG 7DEOH %UDXQH DQG 'RKOHU
Abbreviations: 895 XOWUDYLROHWUDGLDWLRQ89% XOWUDYLROHW %89$ XOWUDYLROHW$0$$ P\FRVSRQQHOLNHDPLQRDFLG36 ,, SKRWRV\VWHP,,*\U \HDUV
&KDSWHU 5HVSRQVHVWR895
Table 1. 7\SLFDODGYHUVHHIIHFWVRI895RQPHWDEROLFSURFHVVHVLQF\DQREDFWHULD $EEUHYLDWLRQV )5IUDFWLRQRIFRQWUROPHWDEROLFDFWLYLW\ UHPDLQLQJDIWHUJLYHQWUHDWPHQW36,,SKRWRV\VWHP,,58%,6&2ULEXORVHELVSKRVSKDWHFDUER[\ODVHR[\JHQDVH DU\(7SULPDU\HOHFWURQ WUDQVSRUW &KO D FKORURSK\OO D 3%3 SK\FRELOLSURWHLQV 3%6 SK\FRELOLVRPH /33 3%6 OLQNHUSRO\SHSWLGH 3& SK\FRF\DQLQ *$$6 JOXWDPDWHV\QWKDVH 335SKRWRSKRELFUHVSRQVH A., Anacystis Synechococcus Anab, Anabaena; O., Oscillatoria N., Nostoc.
3URFHVV
Photosynthesis &DUERQIL[DWLRQ 2[\JHQHYROXWLRQ
'RVH UDWH 7LPH E )5 :P PLQ
3RVVLEOH WDUJHWD
:DYHOHQJWK
36,,58%,6&2
89%
F FRQVWDQW
,DU\(7
89$G
2UJDQLVP
5HIHUHQFH
Anab. flos-aquae O. amphigranulata
'RKOHUHWDO :LQJDUGHWDO
A. nidulans $&,$07 0 A. nidulans
+LURVDZDDQG0L\DFKL .XODQGDLYHOXHWDO .XODQGDLYHOXHWDO .XODQGDLYHOXHWDO
36,,
89%
A. niduluns
36,,
XYF
A. nidulans
3%6H[FLWDWLRQ WUDQVIHU
/33
89%$
Anab. VSDQGN. carmium
6LQKDHWDO
7HWUDS\UURO V\QWKHVLV
$/6
89$
FRQVWDQW
A. nidulans $&,$07 0
+LURVDZDDQG 0L\DFKL
&KURPDWLF DGDSWDWLRQ
"
89$
Fremyella diplosiphon 0
2KNLHWDO
3LJPHQWORVV
3&
89%
/600
'RKOHUHWDO
&KOD
89%
/600
DOO3%3 &KOD
89% 89%
0.2/ GD\V 0.2/ GD\V
S. leopoliensis 6$* S. leopoliensis 6$* O. amphigranulata O. amphigranulata
PXOWLSOH
89$
FRQVWDQW
A. nidulans $&,$07
FRQVWDQW
Oscillatoria %* Chlorogloeopsis 2 &JV Synechococcus 1.%* Oscillatoria %* Phormidium murrayi
+LURVDZDDQG 0L\DFKL :DFKLHWDO
Growth 'HOD\
FRQVWDQW 5DWHUHSUHVVLRQ
PXOWLSOH
89$
FRQVWDQW FRQVWDQW FRQVWDQW
PXOWLSOH
89%
F F FRQVWDQW FRQVWDQW FRQVWDQW
PXOWLSOH
XYF
FRQVWDQW
Synechococcus leopoliensis Anubuena cylindrica Nostoc commune '5+ Phormidium miirrayi Gloeocapsa &&DO G(2) Pliormidium miirrayi
'RKOHUHWDO :LQJDUGHWDO :LQJDUGHWDO
*DUFLD3LFKHOHWDO :DFKLHWDO :DFKLHWDO 9LQFHQW DQG 4XHVDGD 'RKOHUHWDO 'RKOHUHWDO (KOLQJ6FKXO] HW DO 9LQFHQW DQG 4XHVDGD *DUFLD3LFKHO HW DO 9LQFHQW DQG 4XHVDGD
>RYHU
5: &DVWHQKRO]DQG )*DUFLD3LFKHO
Table I FRQWLQXHG 3URFHVV
3RVVLEOH WDUJHWD
:DYHOHQJWK
'RVH UDWH 7LPH )5 E :P PLQ
2UJDQLVP
5HIHUHQFH
6XUYLYDO
PXOWLSOH
89%
N. spongiaeforme Anab. VSDQGN. carmium N. commune DQG Scytonema VS Aphanothece halophytica Agmenellum quadruplicatum
7\DJLHWDO 6LQKDHWDO
Anabaena flos-aquae Anabaena $os-aquae N. spongiaeforme N. calcicola Anab. cylindrica
1HZWRQHWDO 1HZWRQHWDO 7\DJLHWDO .XPDUHWDO 'RKOHUHWDO
Anab. variabilis e
%UDXQH DQG 'RKOHU
Anab. cylindrica
'RKOHUHWDO
Phormidium uncinatum Phormidium uncinatum
+DGHU
Anabaena aequalis
%ODNHILHOG DQG +DUULV %ODNHILHOG DQG +DUULV
XYF
6LQKDHWDO
Nitrogen metabolism 1LWURJHQIL[DWLRQ
QLWURJHQDVH"
89%
QLWUDWHXSWDNH
PHPEUDQH SURWHLQV" PHPEUDQH SURWHLQV"
89%
5 &
89%
*$$6
89%
"
89%
"
89%
89%
.-P ( 0 )
89%
.-P(0) I
DPPRQLXPXSWDNH DPLQRDFLG PHWDEROLVP 0RWLOLW\ DQG 7D[HV VWHSGRZQ335 FHOOPRWLOLW\ Cell differentiation KHWHURF\VW IRUPDWLRQ DNLQHWHLQGXFWLRQ
f
Anabaena aequalis
+DGHUHWDO
D 7KHDVVLJQPHQWRIDSRVVLEOHWDUJHWLVLQPRVWFDVHVRXUV E :KHQ SRVVLEOH WKHLUUDGLDWLRQ FRQGLWLRQV KDYHEHHQUHFDOFXODWHGIRUFRQVLVWDQFHLQXQLWV ,IH[SRVXUHWLPHH[FHHGHG GD\V LWKDV EHHQ UHSRUWHGDVFRQVWDQW F GDLO\H[SRVXUHVRIPLQWZLFH G VKRUWWHUPHIIHFWDFFOLPDWHGFHOOVVKRZQRRUPXFKGDPSHQHGGHSUHVVLRQV H JHUPLQDWLQJDNLQHWHVZHUHLUUDGLDWHG1RHIIHFWZDVVHHQLILUUDGLDWHGMXVWEHIRUHJHUPLQDWLRQ I 2QO\DEVROXWHGRVHUHFHLYHGZDVJLYHQH[FLVVLRQUHSDLUKDGEHHQDUWLILFLDOO\LQKLELWHGZLWKFDIIHLQ
% ([SRVXUHRI&\DQREDFWHULD WR895LQ 1DWXUDO+DELWDWV 3K\VLFDO([SRVXUH,QFLGHQW,UUDGLDQFH DQG2SWLFDO0RGLILFDWLRQVLQ WKH (QYLURQPHQW 7KH DOPRVW H[FOXVLYH SKRWRDXWRWURSKLF PRGH RI JURZWK RIF\DQREDFWHULD GHPDQGV WKDW WKH\ WKULYH LQ KDELWDWV H[SRVHG GLUHFWO\ RU LQGLUHFWO\ WR VRODU UDGLDWLRQ 0RUH RIWHQ WKDQ QRW WKLV UHTXLUHPHQW
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
5: &DVWHQKRO]DQG )*DUFLD3LFKHO
Fig. 1: (IIHFWV RI WKH KDELWDW RQ WKH SK\VLFDO H[SRVXUH RI F\DQREDFWHULD , 7KH VSHFWUDO VFDODU LUUDGLDQFH VXQ DQG VN\ UDGLDWLRQ LQFLGHQW DW JURXQG OHYHO DW QRRQ LQ D FOHDU PLGVXPPHU GD\ DW 1 /DW 7KH UHVW RI WKH SODWHV GHSLFW WKH LQ VLWX VFDODU LUUDGLDQFH H[SHULHQFHG E\ F\DQREDFWHULDOFHOOV WKULYLQJLQVHYHUDOKDELWDWVH[SRVHGWRWKH LQFLGHQWIOX[HV RI,QRWHGLIIHUHQWVFDOHV ,, D³VWURQJVKDGH´KDELWDW 1 IDFLQJVXUIDFHLOOXPLQDWHGE\H[WUHPHO\GLIIXVHVN\UDGLDWLRQRQO\ ZKHUHVFDODULUUDGLDQFHLVYHU\ORZEXWWKHUHODWLYHLPSRUWDQFHRI89LV HQKDQFHG ,,,DSODQNWRQLFKDELWDWXQGHU PRIFOHDURSHQRFHDQZDWHU ZKHUHDOOIOX[HVUHPDLQIDLUO\KLJK DQG 895% DQGYLVLEOHDUH PRUHVWURQJO\DWWHQXDWHG WKDQ 89$ ,9 WKH VXUIDFH RIEHDFK TXDUW] IHOGVSDU VDQG ZKHUH DOO 89% 89$ DQG YLVLEOHDUH KLJKHU WKDQ LQFLGHQWE\ DQGUHVSHFWLYHO\ GXHWROLJKWWUDSSLQJHIIHFWV 9 PGHHSLQDZHWWRSVRLOZKHUH89% DQG89$KDYH EHHQ DWWHQXDWHG EHORZ RILQFLGHQW EXW FD RIWKH YLVLEOH OLJKW UHPDLQV 9, VFDODU LUUDGLDQFH ZLWKLQ WKH WKDOOXV RIWKH WHUUHVWULDO F\DQREDFWHULDOOLFKHQ Collema VS 6HH&RORU3ODWHFGDQGH 7KHVSHFWUDZHUHFDOFXODWHGIURPWKHIROORZLQJVRXUFHV ) *DUFLD3LFKHO XQSXEOLVKHG *DUFLD3LFKHO %GHOHWDO 6PLWKDQG%DNHU
,Q WHUUHVWULDO KDELWDWV WZR IDFWRUV ORFDOO\ PRGLI\ WKHLQFLGHQW895 WKH RULHQWDWLRQ RIWKH VXUIDFHZLWK UHVSHFW WR WKH VRODU YHUWLFDO DQG WKH DOEHGR RI WKH VXSSRUWLQJVXUIDFH ,QVWDQWDQHRXVH[SRVXUH DQG GRVH DUH KLJKHVW LQ VXUIDFHV RULHQWHG RUWKRJRQDO WR WKH 6XQ¶V YHUWLFDO DQG ORZHVW LQ WKRVH LQ WKH RSWLFDO VKDGRZ RIVROLGV +RZHYHU GXH WR WKH SUHIHUHQWLDO DWPRVSKHULF 5D\OHLJK VFDWWHU RI WKH VKRUWHU ZDYHOHQJWKV LQ WKH DWPRVSKHUH WKH VSHFWUDO FRPSRVLWLRQ RIWKH GLIIXVH OLJKWZKLFK SHQHWUDWHV WKH VKDGH XQGHU VROLGV LV KLJKO\ HQULFKHG LQ 895 5RELQVRQ )UHGHULFN HW DO $W ZDYHOHQJWKVRIQPDSSUR[LPDWHO\RQHKDOIRIWKH LQFLGHQW UDGLDWLRQ LV GLIIXVH DQG RQHKDOI FRPHV GLUHFWO\ IURP WKH VRODU GLVF ,W IROORZV WKDW SRSXODWLRQV WKULYLQJ RQ VKDGHG VXUIDFHV ZLOO UHFHLYH ORZHU DEVROXWH 895 GRVHV EXW KLJKHU 895 GRVHV UHODWLYH WR WKH YLVLEOH WKDQ SRSXODWLRQV RQ VXQQ\ VXUIDFHV )LJ 7KLV LPSOLHV WKDW895PD\SOD\ D SDUWLFXODUO\ LPSRUWDQW UROH LQ WKH ELRORJ\ RI F\DQREDFWHULDH[SRVHGWRGLIIXVHOLJKW 7KHUDGLDWLRQ UHIOHFWHG IURP WKH VXUIDFH PD\ FKDQJH WKH DEVROXWH
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¡UJHQVHQ DQG 0DUDLV *DUFLD3LFKHO DQG &DVWHQKRO] .KO DQG-¡UJHQVHQ *DUFLD 3LFKHO DQG %HERXW &ORVH WR WKH VXUIDFH RI WKHVH OD\HUV OLJKW WUDSSLQJ SKHQRPHQD UHVXOW LQ ORFDOL]HG LUUDGLDQFH PD[LPD ZKLFK DUH KLJKHU WKDQ LQFLGHQW LUUDGLDQFH 7\SLFDOO\ WKHUH LV DQ RQVHW RI TXDVLH[SRQHQWLDO DWWHQXDWLRQRIOLJKWEHORZWKHOLJKW WUDSSLQJ ]RQHV 7KXV LQ VHGLPHQWDU\ KDELWDWV RQH ILQGV H[WUHPHO\ VWHHS JUDGLHQWV RI H[SRVXUH UDQJLQJ IURP ]RQHV RI LQFUHDVHG H[SRVXUH )LJ ,9 WR
&KDSWHU 5HVSRQVHVWR895 ]RQHV ZKLFK DUH YLUWXDO UHIXJHV IURP 895 )LJ 9 %HFDXVH OLJKWWUDSSLQJ HIIHFWV DUH JUHDWHVW DW ORQJHUZDYHOHQJWKV DQG DEVRUSWLRQE\SDUWLFXODWHV LV PRVWSURQRXQFHG DWVKRUWHVWZDYHOHQJWKV WKHUDWLR RI 895WRYLVLEOHOLJKWLQYDULDEO\GHFUHDVHV ZLWKGHSWK $ FRPSDULVRQ RI GDWD IURP VHYHUDO VHGLPHQWDU\ PLFURHQYLURQPHQWV VKRZHG WKDW WKH VSDFHDYHUDJHG 895 GRVDJH UDWHV ZLWKLQ WKH HXSKRWLF ]RQH UDQJHG EHWZHHQ DQG RIWKH LQFLGHQW D PXFK KLJKHU 895 H[SRVXUH WKDQ WKRVH LQ WKH HXSKRWLF ]RQHV RI QDWXUDOZDWHUV*DUFLD3LFKHODQG%HERXW
%LRORJLFDOO\(IIHFWLYH ([SRVXUH %LRORJLFDO :HLJKWLQJ )XQFWLRQV ,I RQH VWULYHV WR SUHGLFW WKH ELRORJLFDO LPSDFW RI H[SRVXUH WR 895 RQ WKH EDVLV RI WKH VSHFWUDO LUUDGLDQFH RU GRVHV UHFHLYHG DQG JLYHQ WKDW DOO ELRORJLFDO HIIHFWV RI895 SUHVHQW VWURQJ ZDYHOHQJWK GHSHQGHQFH DIXQFWLRQZKLFKUHODWHVERWKSDUDPHWHUV LV QHHGHG 7KHVH ELRORJLFDO ZHLJKWLQJ IXQFWLRQV %:) \LHOG XSRQ PXOWLSOLFDWLRQ E\ WKH VSHFWUDO LUUDGLDQFH DQ HVWLPDWH RI WKH ELRORJLFDOO\ HIIHFWLYH H[SRVXUH %LRORJLFDO ZHLJKWLQJ IXQFWLRQV DUH LQ SULQFLSOH HTXLYDOHQW WR WKH DFWLRQ VSHFWUD XVHG LQ SKRWRELRORJ\ WR HOXFLGDWH WKH VSHFWUDO FKDUDFWHULVWLFV RI SKRWRUHFHSWRUV RU WDUJHW PROHFXOHV &OD\WRQ 7KH\ GLIIHU IURP WKHP LQ WKDW DEVROXWH XQLWV DUHUHTXLUHG DQG WKHXVHRISRO\FKURPDWLFUDGLDWLRQLV SUHIHUUHGWRPRQRFKURPDWLFEDQGVLQRUGHUWRLQFOXGH DQ\ V\QHUJLVWLF HIIHFWV RIHLWKHU YLVLEOH OLJKW RU RWKHU 89 ZDYHOHQJWKV LH &XOOHQ HW DO 8QIRUWXQDWHO\ QR ULJRURXV F\DQREDFWHULDO %:) IRU DQ\WDUJHWSURFHVV LV \HW DYDLODEOH HYHQ WKRXJK WKHUH LV DPSOH HYLGHQFH RI WKH HIIHFW RI 895LQGXFHG JURZWK GHSUHVVLRQ DQG GDPDJH WR '1$ SKRWRV\QWKHVLV GLQLWURJHQ IL[DWLRQ DQG VHYHUDO RWKHU PHWDEROLF SURFHVVHV 7DEOH DV ZHOO DV VWURQJ ZDYHOHQJWK GHSHQGHQFH (YHQ WKRXJK WKH XVH RI %:)DVDSUHGLFWLYHWRROLVQRWZLWKRXWSUREOHPVWKH GHWHUPLQDWLRQ RI F\DQREDFWHULDO %:)V IRU VHYHUDO WDUJHW SURFHVVHV VKRXOG EH D SULRULW\ WDVN LI RQH ZLVKHV WR XQGHUVWDQG WKH LPSRUWDQFH RI895 RQ WKH DEXQGDQFH DQG GLVWULEXWLRQ RI F\DQREDFWHULD LQ QDWXUHDQGLIRQHZLVKHVWRSUHGLFWWKHSRVVLEOHDIIHFW RI D UHGXFWLRQ LQ VWUDWRVSKHULF R]RQH RQ F\DQREDFWHULDOSRSXODWLRQV
0HWDEROLFDOO\7LPHG ([SRVXUH &\DQREDFWHULD VHHP WR EH VXFFHVVIXO FRORQL]HUV RI KDUVK KDELWDWV ZKHUH PHWDEROLF DFWLYLW\ LV UHVWULFWHG
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cytonema VS Nostoc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
D +RW6SULQJV $OWKRXJK KRW VSULQJV PD\ RFFXU WRGD\ LQ QXPHURXV VLWHV DVVRFLDWHG ZLWK YROFDQLF RU WHFWRQLF DFWLYLW\ PRVWWHUUHVWULDOVSULQJVDUHH[SRVHGWRIXOOVXQVLPSO\ EHFDXVH PRVW YDVFXODU SODQWV LQ WKH LPPHGLDWH YLFLQLW\RIJHRWKHUPDODFWLYLW\DUHNLOOHG ,QDGGLWLRQ VRPH RI WKH PDMRU UHJLRQV RI KRW VSULQJV VXFK DV WKRVH RI
E ,QWHUWLGDODQG 2WKHU6KDOORZ0DULQH +DELWDWV ,QWHUWLGDOPDULQHKDELWDWVRI³QRUPDO´VDOLQLW\VXFKDV PXGIODWVDQGPDUVKHVDUHPRVWO\GHYRLGRIH[WHQVLYH RUSHUHQQLDOF\DQREDFWHULDGRPLQDWHGPLFURELDOPDWV SULQFLSDOO\ EHFDXVH RI D YDULHW\ RI HXNDU\RWLF FRPSHWLWRUV DQG KHUELYRUHV 1HYHUWKHOHVV VRPH LQWHUWLGDOIODWV DUHLQXQGDWHGE\ZDWHUVR LQIUHTXHQWO\ HJ /DJXQD *XHUUHUR 1HJUR 0H[LFR &KDSWHU WKDWPRVWHXNDU\RWHVDUH DEVHQWRYHUODUJH DUHDV DQG F\DQREDFWHULDOPDWV RIWHQGRPLQDWHGE\VF\WRQHPLQ ULFK Lyngbya FI aestuarii DQG Calothrix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
5:&DVWHQKRO]DQG )*DUFLD3LFKHO JHQHUDOO\ FOHDU VKDOORZ ZDWHUV RIXQSROOXWHG WURSLFDO DQGRFHDQLF ZDWHUV DQGDUH WKHUHIRUHH[SRVHGWRKLJK 895 OHYHOV 0DQ\ RI WKH ODWWHU IRUP ZLVSOLNH HSKHPHUDO FRYHUV RU WRXJKHU JHODWLQRXV WXIWV RQ YDULRXVVXEVWUDWHV
F %HQWKLF)UHVKZDWHU+DELWDWV &OHDU ZDWHU KDELWDWV RFFXU LQ ROLJRWURSKLF ODNHV WKURXJKRXW WKH (DUWK DQG WKHVH LQFOXGH D ODUJH SURSRUWLRQ RI DOSLQHVXEDOSLQH ODNHV $OWKRXJKPRVW RIWKHVHKDELWDWV DUH QRWGRPLQDWHGE\ F\DQREDFWHULD VRPH XOWUDROLJRWURSKLF ODNHV VXFK DV :DOGR /DNH 2UHJRQKDYHGRPLQDQWEHQWKLFFRYHUVRIVF\WRQHPLQ ULFK Stigonema DQG Scytonema WR GHSWKV RI VHYHUDO PHWHUV &DVWHQKRO] XQSXEOLVKHG GDWD 6RPH RI WKHVH ROLJRWURSKLF IUHVK ZDWHUV DUH GRPLQDWHG E\ F\DQREDFWHULD E\ YLUWXH RI DQ H[WUHPH GHILFLHQF\ RI FRPELQHG QLWURJHQ EXW QRW QHFHVVDULO\ RI RWKHU QXWULHQWV ,Q WKHVH HQYLURQPHQWV KHWHURF\VWRXV F\DQREDFWHULDPD\ EH FRQVSLFXRXV VXFK DV WKH KXJH VORZJURZLQJ VF\WRQHPLQFRQWDLQLQJ FRORQLHV RI Nostoc FI pruniforme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ostoc 3ODWHF
G 7HUUHVWULDO+DELWDWV ,Q H[SRVHG WHUUHVWULDO KDELWDWV WKDW DUH H[WUHPH E\ YLUWXH RI SHULRGLF RU ORQJWHUP GHVLFFDWLRQ F\DQREDFWHULD WKULYH E\ YLUWXH RI WKHLU WROHUDQFH WR GHVLFFDWLRQ EXW SUHVXPDEO\ DOVR EHFDXVH RI D KLJK WROHUDQFHWR895WR ZKLFK WKH\DUH VRRIWHQH[SRVHG 3RWWV 7KH WHUUHVWULDO KDELWDWV ZLWK FRQVSLFXRXV F\DQREDFWHULDO PDWV RU FUXVWV LQFOXGH
&KDSWHU 5HVSRQVHVWR895 PDQ\ RI WKH ZDUP DQG FROG GHVHUWV RI WKH (DUWK ZKHUH WKH F\DQREDFWHULD PD\ IRUP D JURXQG FRYHU RIWHQ EHWZHHQ DQG DURXQG GHVHUW VKUXEV VHH *DUFLD 3LFKHO DQG %HOQDS 0D]RU HW DO 3ODWH E 7KH KDUGHU WHUUHVWULDO VXEVWUDWHV VXFK DV FOLII IDFHV DUH RIWHQ FRYHUHG E\ SHULRGLFDOO\ZHWWHG F\DQREDFWHULDO FUXVWV ZKLFK DUH RIWHQ GDUN LQ FRORU GXH WR VF\WRQHPLQ RU ³JORHRFDSVLQ´ 3ODWHV G DQG H 7KHVH GDUN DUHDV DUH RIWHQ UHIHUUHG WR DV ³WLQWHQVWULFKH´ LQN VWUHDNV 3ODWH E &\DQROLFKHQVFRPPRQLQDODUJHYDULHW\RIWHUUHVWULDO KDELWDWVPD\DOVREHH[SRVHGWRKLJKVRODULUUDGLDQFH DQG DV D FRQVHTXHQFH FRQWDLQ VF\WRQHPLQ ZLWKLQ WKH WKDOOXV%GHOHWDO 3ODWHFGDQGH (YHQ PRUH H[WUHPH WHUUHVWULDO KDELWDWV VXFK DV WKH RXWHU VKHOOV RI KRW DQG FROG GHVHUW VDQGVWRQHV KDUERU FU\SWRHQGROLWKLF F\DQREDFWHULD HJ Chroococcidiopsis VSS3ODWHV DQG ,WLVSRVVLEOH WKDW WKHVH WRR PD\ EH H[SRVHG WR KLJK 895 IOXHQFH UDWHV 1LHQRZ DQG )ULHGPDQQ DOWKRXJK VSHFLILF895LQWHQVLWLHVLQWKHVHVLWXDWLRQVZHUHQRW PHDVXUHG
H 0DULQHDQG)UHVKZDWHU3ODQNWRQ 7KH F\DQREDFWHULD RI WKH SODQNWRQ DUH H[SRVHG DW OHDVW SHULRGLFDOO\ WR KLJK VRODU LUUDGLDQFH ZKHQ WXUEXOHQFH EULQJV WKHP QHDU WKH VXUIDFH XQGHU FRQGLWLRQV RI KLJK OLJKW &KDSWHU 3LFR SODQNWRQLF F\DQREDFWHULD VXFK DV PDULQH VSHFLHV RI Synechococcus DQG Prochlorococcus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richodesmium LQ ZDUPHU ZDWHUVDQGE\VSHFLHVRIAnabaena, Aphanizomenon, Oscillatoria, Limnothrix, Microcystis, Coelosphaerium DQG RWKHUV LQ IUHVK DQG EUDFNLVK ZDWHUV $OWKRXJK UHJXODWLRQ RIYHUWLFDO SRVLWLRQ LQ WKH ZDWHU FROXPQ RFFXUV LQ VRPH RIWKH JDV YDFXRODWH IRUPV VRPH IRUPV DUH VLPSO\ EXR\DQW :DOVE\ 7KHVH F\DQREDFWHULD PD\ VLPSO\ DFFXPXODWH RQ WKH
VXUIDFH GXULQJ FDOP SHULRGV RI ORZ WXUEXOHQFH ZLWK KLJK PRUWDOLW\ DV D UHVXOW RI H[SRVXUH WR KLJK VRODU LUUDGLDQFH 0DQ\ RI WKH ODUJHU SODQNWRQLF F\DQREDFWHULD VXFK DV Trichodesmium, Aphanizomenon, Dactylococcopsis, DQG RWKHUV DUH NQRZQ WR DFFXPXODWH ODUJH TXDQWLWLHV RI SXWDWLYH LQWUDFHOOXODUVXQVFUHHQV RIWKH0$$ W\SH EXW 0$$V GR QRW VHHP WR RFFXU LQ SLFRSODQNWRQLF IRUPV *DUFLD3LFKHODQG$SSHOXQSXEOLVKHGGDWD
,, 6WUDWHJLHVRI 8957ROHUDQFHE\ &\DQREDFWHULD $ 896FUHHQLQJ &RPSRXQGV 0$$VDQG2WKHU&RPSRXQGV 7KHXVH RI895DEVRUELQJ FRPSRXQGV DV VXQVFUHHQV LV NQRZQ LQ PDQ\ RUJDQLVPV $ VXQVFUHHQUHSUHVHQWV D SDVVLYH PHFKDQLVP IRU SURWHFWLRQ 895 UDGLDWLRQ LV WR VRPH H[WHQW DEVRUEHG E\ WKH VXQVFUHHQ DQG FDQQRW LQWHUDFW ZLWK SRWHQWLDO FHOOXODU WDUJHWV 7KH PHODQLQV RIWHJXPHQWDO DQLPDOV DUH SHUKDSV WKH EHVW NQRZQ H[DPSOH RI VXFK VXQVFUHHQV .ROOLDV HW DO ,QKLJKHUSODQWVSKHQ\OSURSDQRLGV DQGUHODWHG VXEVWDQFHVDQWKRF\DQLQVIODYRQHJO\FRVLGHV SURYLGH SURWHFWLRQ DJDLQVW 895 7DNDKDVKL HW DO ,Q PLFURRUJDQLVPV VXQVFUHHQ UROHV ZHUH VXJJHVWHG IRU RU DVFULEHG WR D YDULHW\ RI VXEVWDQFHV LQFOXGLQJ PHODQLQV DQG XVQLF DFLG LQ IXQJL SURGLJLRVLQH LQ Serratia marcensis DQG YLRODFHLQ LQ Chromobacterium violaceum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
895 DEVRUSWLRQ LV D QHFHVVDU\ EXWFHUWDLQO\ QRW D VXIILFLHQW FULWHULRQ WR GHWHUPLQH WKDW D FRPSRXQG PD\IXQFWLRQDVDVXQVFUHHQ $QLQWHUHVWLQJH[DPSOH LV WKDW RI WKH ELRSWHULQ JOXFRVLGH UHSRUWHG IURP D 89$WROHUDQWPDULQH Oscillatoria 0DWVXQDJDHWDO :DFKL HW DO %HFDXVH WKH FHOOXODU FRQWHQW RIWKLV FRPSRXQG FDQ EH UDLVHG E\ H[SRVXUH RIWKH FHOOV WR 89$ LQ D PDQQHU SURSRUWLRQDO WR LWV LQWHQVLW\ WKH FRPSRXQG ZDV FRQVLGHUHG WR EH SKRWRSURWHFWLYH 7KLV DQG RWKHU ELRSWHULQ JO\FRVLGHV ZHUH VXEVHTXHQWO\ LGHQWLILHG LQ RWKHU F\DQREDFWHULD HJ Synechococcus leopoliensis, Microcoleus chthonoplastes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ostoc commune 6FKHUHU HW DO ZDV VKRZQ WR EH D PL[WXUH RI ROLJRVDFFKDULGH0$$V 260$$ %RKP HW DO 7KH JO\FRV\ODWHG 0$$V RI N. commune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
&KDSWHU 5HVSRQVHVWR895
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icrocoleus chthonoplastes .DUVWHQ DQG *DUFLD 3LFKHO XQSXEOLVKHG GDWD 7KH DGDSWLYH DGYDQWDJH JDLQHG E\ DYRLGLQJ RVPRWLF OLPLWDWLRQV DQG E\ HQKDQFLQJ WKH RSWLFDO HIILFLHQF\ PD\ KDYH GULYHQ WKH HYROXWLRQ RI H[WUDFHOOXODU 0$$V 260$$ %H\RQG DQHFGRWDO REVHUYDWLRQV ILHOG GDWD RU FRUUHODWLRQV EHWZHHQ F\DQREDFWHULDO 0$$V DQG RWKHU HQYLURQPHQWDOSDUDPHWHUVDUHVWLOOODFNLQJ
6KHDWK3LJPHQWV
Fig.2. &KHPLFDOVWUXFWXUHRIVRPHVXQVFUHHQFRPSRXQGVLGHQWLILHG LQ F\DQREDFWHULD 7KH VWUXFWXUHV DUH VF\WRQHPLQ 3URWHDX HW DO 260$$%RKPHWDO P\FRVSRULQHJO\FLQH,WR DQG +LUDWD SRUSK\UD 7DNDQR HW DO VKLQRULQH 7VXMLQRHWDO DQGDVWHULQD1DNDPXUDHWDO
VS FRXOG QRW EH GLVFRXQWHG *DUFLD3LFKHO HW DO ,Q F\DQREDFWHULD D FRQVWLWXWLYH OHYHO RI 0$$V DSSHDUV WR EH SUHVHQW XQGHU DOO JURZWK FRQGLWLRQV EXW WKH VSHFLILF FRQWHQW FDQ EH VLJQLILFDQWO\ UDLVHG E\ H[SRVXUHWR 89%$UDGLDWLRQ ZLWK D PD[LPXP DW QP *DUFLD3LFKHO DQG &DVWHQKRO] ,QGloeocapsa VSWKHOHYHOVZHUH VKRZQWREH DGLUHFWIXQFWLRQ RIWKH LQFLGHQW89 LUUDGLDQFH DQG ORQJHUZDYHOHQJWKV ZHUHLQFDSDEOH RI HOLFLWLQJWKHUHVSRQVH 6LPLODUHYLGHQFHZDVJDWKHUHG IRU WKH 260$$V IURP WHUUHVWULDO Nostoc commune %RKPHWDO (KOLQJ6FKXO]HWDO 7KH VLQJOHFHOO VXQVFUHHQ HIIHFW RI LQWUDFHOOXODU 0$$V LQ F\DQREDFWHULD LV PRGHVW RI LQFLGHQW SKRWRQV ZHUH LQWHUFHSWHG LQ D IDLUO\ ODUJH FHOOHG Gloeocapsa VS DOWKRXJK FRORQ\IRUPDWLRQ DQG PDWW\SH JURZWK PD\ VXEVWDQWLDOO\ LQFUHDVH WKH HIILFLHQF\ 7KHFRQFHQWUDWLRQ RIWKH VXQVFUHHQ LV QRW WRRKLJKEHORZ RIWKHGU\ZHLJKWLQPRVWFDVHV DQG LW VHHPV WKDW WKH DFFXPXODWLRQ RI LQWUDFHOOXODU 0$$ PD\ SURYLGH VRPH EHQHILW IRU D VPDOO
&\DQREDFWHULDO VKHDWKV DUH RIWHQ FRORUHG
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ostoc commune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cytonema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³PHWDEROLFDOO\WLPHGH[SRVXUH´ 6HFWLRQ% /LWWOH LV NQRZQ DERXW WKH UHGYLROHW SLJPHQWV JORHRFDSVLQ RWKHU WKDQ WKHLU in situ UHDFWLRQ WR DFLGEDVH 1DJHOL DQG 6FKZHQGHUHU DQG WKHLU FRPPRQ RFFXUUHQFH LQ QDWXUDO SRSXODWLRQV RI F\DQREDFWHULD HJ Gloeocapsa VS 3ODWH F RQ YHUWLFDO IDFHV RIFOLIIV LQ PDQ\ PRXQWDLQRXV UHJLRQV ³7LQWHQVWULFKH´RULQNVWUHDNV-DDJ 3ODWHE
5:&DVWHQKRO]DQG )*DUFLD3LFKHO DQG LQ VRPH F\DQREDFWHULDO VRLO FUXVWV DQG ILODPHQWRXVWURSLFDO IRUPV HJ Porphyrosiphon VS $SSDUHQWO\ QR JORHRFDSVLQSURGXFLQJ VWUDLQV DUH LQ FXOWXUH3ODWHD E DQG F 7KH SHULSKHUDO SLJPHQW GLVWULEXWLRQ LQ FRORQLDO IRUPV SDUDOOHOV WKDW RI VF\WRQHPLQ ZLWK WKH LPSOLFDWLRQ WKDW H[SRVXUH WR VRODU LUUDGLDQFH FDXVHV LWV DFFXPXODWLRQ 2SWLFDO HYLGHQFH ZLWK QDWXUDO VDPSOHV VXJJHVWV WKDW WKH VXQVFUHHQ HIIHFW GXH WR JORHRFDSVLQ LV VXEVWDQWLDO *DUFLD3LFKHO XQSXEOLVKHGGDWD
% $YRLGDQFH 5HVSRQVHVE\0RWLOH &\DQREDFWHULD %HQWKLF 0LFURELDO0DW3RSXODWLRQV 7KHUH DUH QXPHURXV UHSRUWV RI WKH GRZQZDUG PRYHPHQW RI PRWLOH ³RVFLOODWRULDQ´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scillatoria VS DQGSpirulina FI subsalsa. 7KXV LW LV SRVVLEOH WKDW RSWLPL]DWLRQ RI SKRWRV\QWKHWLF LUUDGLDQFH ZDV DFKLHYHG DQG ZDV QRW VLPSO\ WKH DYRLGDQFH RI KDUPIXO LQWHQVLWLHV 1HYHUWKHOHVV WKH LQYROYHPHQWRI89LUUDGLDQFHZDV QRWLQYHVWLJDWHG LQ WKLVVWXG\
&KDSWHU 5HVSRQVHVWR895 *DUFLD3LFKHO DQG &DVWHQKRO] VXJJHVWHG WKDW : LUUDGLDQFH ZDV DSULPDU\ FXH IRU WKH YHUWLFDO PRYHPHQWV RI F\DQREDFWHULD ,Q DQ LQGLUHFW PHDVXUHPHQW RI YHUWLFDO PRYHPHQW SULPDULO\ RI Microcoleus FI chthonoplastes LQ D PDW IURP 6RODU /DNH(J\SW 89% ZDVSDUWLFXODUO\HIIHFWLYH%HERXW DQG*DUFLD3LFKHO $UWLILFLDOLOOXPLQDWLRQZDV XVHG DWLQWHQVLWLHVWKDWFRUUHVSRQGHGWR VWURQJQDWXUDO IOX[HV RIWKH 89% FRPSRQHQW EXW89$ DQG YLVLEOH OLJKWZHUHPXFKORZHUWKDQQDWXUDO ,QDPRUHUHFHQW RXWGRRU VWXG\ RI PRWLOH Spirulina DQG Oscillatoria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scillatoria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³RVFLOODWRULDQV´ LQ
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
3ODQNWRQLF$YRLGDQFH 5HVSRQVHV 6LQFH VRPH SODQNWRQLF JDV YDFXRODWH F\DQREDFWHULD DUHNQRZQWREHFDSDEOHRIYHUWLFDOSRVLWLRQLQJLQWKH ZDWHU FROXPQ :DOVE\ LW LV SRVVLEOH WKDW D VXEWOHUHJXODWLRQLQVRPHFDVHVPD\EHLQUHVSRQVHWR 89 ZDYHOHQJWKV ZKLFKSHQHWUDWHWKHZDWHU 7KHUHLV DPSOHHYLGHQFHWKDWPDULQHSK\WRSODQNWHUV LQFOXGLQJ F\DQREDFWHULD DUH QHJDWLYHO\ DIIHFWHG E\ 895 DW OHYHOVFRPSDUDEOHWRWKRVHLQQDWXUH+ROP+DQVHQHW DO EXW WKHUH LV QR LQIRUPDWLRQ DW SUHVHQW RQ VSHFLILF DYRLGDQFH UHVSRQVHV UHODWHG WR 89 LUUDGLDQFH
& 7ROHUDQFH WR 8955HTXLULQJ$FWLYH 5HSDLURU'H 1RYR 6\QWKHVLV 7KH DFFOLPDWLRQ RIFHOOV WR 895 PD\ LQYROYH DFWLYH UHSDLU PHFKDQLVPV LQ ZKLFK GDPDJHG WDUJHWV DUH SDUWLDOO\VXEVWLWXWHGRUWKHGDPDJHLVUHSDLUHGZLWKRXW WKHQHHGIRUde novo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
SK\VLRORJLFDO EDVLV RI WKLV SKHQRPHQRQ ZDV RQO\ H[SORUHGVXSHUILFLDOO\ ([SRVXUH RIF\DQREDFWHULD WR 895 HOLFLWHG de novo V\QWKHVLV RI VRPH NQRZQ 5HF$ FKDSHURQHV DQG PDQ\ XQLGHQWLILHG SHSWLGHV 2ZWWULP DQG &ROHPDQ &KLWQLV DQG 1HOVRQ 7KDWH[WUDFKURPRVRPDOJHQHWLFHOHPHQWVPD\ EH LQYROYHG LQ F\DQREDFWHULDO 89 UHVLVWDQFH LV DQ LQWHUHVWLQJSRVVLELOLW\.XPDUDQG.XPDU
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scherichia coli )LJ LQ-DJJHU :KLOH WKHUH LV GLUHFW HYLGHQFH IRU WKH SUHVHQFH RI GDUN H[FLVLRQ UHSDLU LQ F\DQREDFWHULD /HYLQH DQG 7KLHO 2 %ULHQ DQG +RXJKWRQ F\DQREDFWHULDVHHPWRUHO\KHDYLO\ RQ SKRWRUHDFWLYDWLRQ IRU '1$ UHSDLU /HYLQH DQG 7KLHO%ODNHILHOGDQG+DUULV
5HSDLURIWKH 3KRWRV\QWKHWLF0DFKLQHU\ $IWHU DFXWH H[SRVXUHV WR 895 LI VXEVHTXHQW WUHDWPHQWV DUH PRGHUDWH DW OHDVW D SDUWLDO UHFRYHU\ PD\ RFFXU SKRWRV\VWHP ,, DFWLYLW\ UHVXPHV DQG VSHFLILF FRQWHQWV RI EOHDFKHG SKRWRSLJPHQWV ULVH 7KLVUHVSRQVHPLJKWVLPSO\EHGXHWRDQLQFUHDVHGRU UHQHZHG H[SUHVVLRQ RI JHQHV WKDW HQFRGH UHSODFHPHQWVIRUWKHGDPDJHGPROHFXOHV +RZHYHUD VSHFLILF OLJKW UHJXODWHG UHSDLU V\VWHP IRU GDPDJH WR 36,, ZKLFK GRHV QRW LQYROYH SURWHLQ RU 51$
5:&DVWHQKRO]DQG )*DUFLD3LFKHO V\QWKHVLV ZDVFODLPHGIRUA. nidulans %KDWWDFKDUMHH DQG'DYLG %KDWWDFKDUMHHHWDO 2QWKH RWKHU KDQG LW ZDV VKRZQ WKDW WKH JHQHV LQ psbA PXOWLJHQHIDPLO\ FRGLQJIRU GLIIHUHQWIRUPVRIWKH SURWHLQ ' RI 36,, LQ Synechococcus 3&& ZHUH H[SUHVVHG GLIIHUHQWLDOO\ DFFRUGLQJ WR WKH OLJKW LQWHQVLW\ *ROGHQ $QDO\VLV RI WKH SHUIRUPDQFH RIPXWDQWV GHIHFWLYH LQ VRPH RIWKH ' IRUPV VKRZHG WKDW RQH RI WKHP DOORZHG LQFUHDVHG UHVLVWDQFH WR SKRWRLQKLELWLRQ FDXVHG E\ YLVLEOH OLJKW *ROGHQ 6LPLODUO\ PXWDJHQHVLV RI WKH psbA JHQH RISynechocystis UHVXOWHG LQ SKRWRWROHUDQW PXWDQWV6DWRK 6LQFH WKH PDLQ WDUJHW RI89 GDPDJH WR 36,, VHHPHG WR EH WKH ' SURWHLQ LW LV SRVVLEOHWKDWVSHFLILF' SRO\SHSWLGHVUHVLVWDQWWR89 GDPDJH HYROYHG LQ F\DQREDFWHULD *DUFLD3LFKHO 7KH SUHVHQFH RI D UHSDLU PHFKDQLVP IRU WKH SKRWRV\QWKHWLF DSSDUDWXV RWKHU WKDQ WKH de novo V\QWKHVLV RI DIIHFWHG SRO\SHSWLGHVFKURPRSKRUHV UHPDLQV WR EH H[SORUHG H[SHULPHQWDOO\ De novo V\QWKHVLVRIWKH' DQG'UHDFWLRQFHQWHUVXEXQLWVLV NQRZQ DV D NH\ VWHS RI WKH UHSDLU SURFHVV LQ Synechocystis VS3&&6DVVHWDO
3URWHFWLRQE\&DURWHQRLGV &DURWHQRLG DEVRUSWLRQ PD[LPD RFFXU PDLQO\ LQ WKH YLVLEOH VSHFWUDO UDQJH LH ! QP EXW RIWHQ H[WHQG PLQLPDOO\ LQWR WKH 89$ UHJLRQ QP *RRGZLQ .R\DPDDQG+DVKLPRWR DQG VRPH KDYH D VPDOO DEVRUSWLRQ SHDN LQ WKH 89% UHJLRQ 7KHLU IXQFWLRQDV895VFUHHQLQJSLJPHQWVLV PLQLPDO DOWKRXJKLQVRPHHXNDU\RWHVHJWKH JUHHQ DOJDH Haematococcus, Trentepohlia, DQG Dunaliella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
&KDSWHU 5HVSRQVHVWR895 GHSHQGHQW ELRV\QWKHWLF UHSDLU SURFHVVHV VHH 5RRV DQG 9LQFHQW ,Q DVWUDLQ RIChroococcidiopsis *DUFLD3LFKHO DQG 'RU XQSXEOLVKHG GDWD H[SRVXUH WR89% KSHUGD\DW :P OHGWRDQDEVROXWH IROG LQFUHDVH LQ FDURWHQRLG FRQWHQW 0\[R[DQWKRSK\OOV DQG [DQWKLQHV LQFUHDVHG ZKLOH FDURWHQH GHFUHDVHG DQG HFKLQHQRQH UHPDLQHG WKH VDPH $ RQHGD\ H[SRVXUH RI Nostoc commune WR 89% UDGLDWLRQ UHVXOWHG LQ D VPDOO LQFUHDVH LQ P\[R[DQWKRSK\OO DQG HFKLQHQRQH FRQWHQW (KOLQJ 6FKXOW] HW DO ,Q RQH FDVH D UHODWLYHO\ 895 UHVLVWDQW VWUDLQ RI Gloeocapsa alpicola SRVVHVVHG D KLJKHUFDURWHQRLGFRQWHQWWKDQWKHZLOGW\SH%XFNOH\ DQG+RXJKWRQ ,Q PDQ\ FDVHV WKH FKDQJH WR KLJK FDURWHQRLGFKORURSK\OOSK\FRELOLQ UDWLRV LQ F\DQREDFWHULDPD\ EH LQ UHVSRQVH WR ORZ DYDLODELOLW\ RI VHYHUDO QXWULHQWV LQFOXGLQJ QLWURJHQ &ROOLHU DQG *URVVPDQ ,WLV QRWNQRZQ LIVXFK FHOOV KDYH DQ LQFUHDVHG UHVLVWDQFH WR 895 ,Q VRPH QRQ SKRWRV\QWKHWLF PLFURRUJDQLVPV VXFK DV Neurospora crassa, FDURWHQRLGOHVV PXWDQWV RU ZLOGW\SH FHOOV JURZQ LQ WKH SUHVHQFH RI LQKLELWRUV RI FDURWHQRLG V\QWKHVLVZHUHPRUHVHQVLWLYHWR89% WKDQZLOGW\SH FHOOV ZKLFK SRVVHVVHG FDURWHQRLGV EXW WKH QDWXUH RI WKH FDURWHQRLG SURWHFWLRQ ZDV XQNQRZQ 0RUULV DQG 6XEGHQ 7KH89% SURWHFWLYHHIIHFWRIYDULRXV FDURWHQRLGV ZDV UHFHQWO\ HYDOXDWHG LQ E. coli WUDQVIRUPHG ZLWK VHYHUDO FDURWHQRJHQLF JHQHV 6DQGPDQQHWDO ,Q F\DQREDFWHULD DOVR LW DSSHDUV WKDW WKH UROH RI FDURWHQRLGVLQ895SURWHFWLRQLVLQGLUHFWIXQFWLRQLQJ DV DQWLR[LGDQWV TXHQFKHUV RI SKRWRVHQVLWL]DWLRQ SURGXFWV VXFK DV WULSOHW FKORURSK\OO VLQJOHW 2 SHUR[\UDGLFDOV DQG DOVR DV LQKLELWRUV RIIUHH UDGLFDO UHDFWLRQV .ULQVN\ %ULWWRQ +LJK LQWHQVLW\ YLVLEOH ZDYHOHQJWKV HVSHFLDOO\ EOXH DQG 89$ DUH HVSHFLDOO\ HIIHFWLYH LQ SURGXFLQJ WKH SKRWRSURGXFWV WKDW FDURWHQRLGV KDYH WKH DELOLW\ WR ³TXHQFK´ EXW WKH UHODWLYH LPSRUWDQFH RI 89$ UHPDLQV XQHYDOXDWHG ,W LV FRPPRQ WR ILQG DQHFGRWDO DOOXVLRQV LQ WKH VFLHQWLILF OLWHUDWXUH DV WR WKH 89 SURWHFWLYHUROHRIFDURWHQRLGVLQF\DQREDFWHULD
2WKHU8956WUHVV5HVSRQVHV 5DGLDWLRQLQWKHQPUDQJH89% DQG89$ DWDQLQWHQVLW\ZKLFKUHVXOWHGLQFHOOVXUYLYDO FKDQJHG WKH SDWWHUQ RI SURWHLQ V\QWKHVLV LQ WKH XQLFHOOXODUF\DQREDFWHULXPAnacystis nidulans 5 Synechococcus VS 6KLEDWD HW DO 6L[WHHQ ³89VKRFN´ SURWHLQV ZHUH V\QWKHVL]HG +HDW VKRFN
ZLWKRXW895LQGXFHG WKH V\QWKHVLV RIVHYHQSURWHLQV WKDWZHUHVLPLODU LQWHUPVRI6'63$*(PRELOLW\WR VHYHQ RI WKH ³89VKRFN¶ SURWHLQV 6L[ GLVWLQFW SURWHLQV ZLWK VLPLODULW\ WR WKH ³89VKRFN´ SURWHLQV ZHUH DOVR LQGXFHG VLPSO\ E\ R[LGDWLYH VWUHVV 0RUH UHFHQWO\ 6KLEDWD HW DO GHPRQVWUDWHG WKDW D IRUP RI SURR[LGDQW OXPD]LQH IURP F\DQREDFWHULDO FHOOV JHQHUDWHG VXSHUR[LGH DQLRQ XQGHU ³QHDU´ 895 6KLEDWDHWDO VXJJHVWHGWKDWH[SRVXUHWR895 VKLIWHG V\QWKHWLF PHWDEROLVP WR WKH SURGXFWLRQ RI HQ]\PHV WKDW VFDYHQJH UHDFWLYH R[\JHQ PRLHWLHV WKHUHE\ OLPLWLQJ 895 GDPDJH 3 1LFKROVRQ HW DO DOVR REVHUYHG WKH V\QWKHVLV RI VRPH DSSDUHQWO\XQLTXH ³VWUHVV´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ynechococcus cf. lividus ELRILOPV DW 2FWRSXV 6SULQJ LQ
606 LQKLELWLRQ RI VKRUWWHUP SKRWRV\QWKHVLV SDUWLFXODUO\DWQRQRSWLPDOWHPSHUDWXUHV0LOOHUHWDO &KDSWHU +RZHYHU WKHVH SRSXODWLRQV RI Synechococcus SHUVLVWHG HYHQ DW VXSUDRSWLPDO WHPSHUDWXUHV LQ VXPPHU XQGHU IXOO VRODU LUUDGLDQFH DOWKRXJK WKH\ ZHUH ZLWKRXW R[\JHQLF FRPSHWLWRUV DERYH DERXW & ,W LV OLNHO\ WKDW FHOO UHSDLU SURFHVVHV SRVVLEO\ PDLQO\ DWQLJKW DUH UHTXLUHG IRU VXUYLYDO $QDWXUDOZDUPVSULQJVWUHDPZLWKDF\DQREDFWHULDO ELRILOP FRPPXQLW\ FRPSRVHG DOPRVW HQWLUHO\ RI Calothrix, D VSHFLHV ULFK LQ WKH VKHDWK SLJPHQW VF\WRQHPLQ ZDV WKH VXEMHFW RI WKUHHPRQWK VXPPHU H[SHULPHQWV LQ
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ynechococcus VS ZDV ODUJHO\ XQDIIHFWHG 6XUSULVLQJO\ WKH JURZWK RI F\DQREDFWHULDO SHULSK\WRQ Merismopedia VS DQG Phormidium VS ZDV VWURQJO\ VWLPXODWHG E\ WKH LQFOXVLRQRI89$ 7KHUH LVUHFHQWHYLGHQFH WKDWWKH 89 FRPSRQHQW RI LUUDGLDQFH LV QHFHVVDU\ WR PDLQWDLQ WKH QRUPDO WKUHH GLPHQVLRQDO VWUXFWXUH RIF\DQREDFWHULDO PDWV DW OHDVW DHULDO PDWV RQ EODFN PDQJURYH LQ WKH WURSLFV 53 6KHULGDQ SHUVRQDO FRPPXQLFDWLRQ 6KHULGDQ KDV HYLGHQFHWKDWVSHFLHVPRUHUHVLVWDQWWR895QRUPDOO\ RFFXS\ WKH PDW VXUIDFH JLYLQJ SURWHFWLRQ WR PRUH VHQVLWLYHVSHFLHVEHORZ :LWKRXW895 RSSRUWXQLVWLF VSHFLHV RI F\DQREDFWHULD RYHUJUHZ WKH ODERUDWRU\ PDLQWDLQHG PDWV DQG ZLWK WKLV GLVRUJDQL]DWLRQ QLWURJHQIL[DWLRQZDV VHYHUHO\UHGXFHG 7KHDGGLWLRQ RI 895 LQ WKH ODERUDWRU\ PDLQWDLQHG D FRQGLWLRQ PRUHVLPLODUWRWKDWRIWKHILHOG
,9 &RQFOXVLRQV ,W LV REYLRXV WKDW DQ LQFUHDVLQJ QXPEHU RI UHVHDUFK VFLHQWLVWV DUH FRQVLGHULQJ WKH HIIHFWV RI 895 RQ YDULRXV SKRWRV\QWKHWLF EDFWHULD DQG PLFURDOJDH 1RW DOOKDYHEHHQFLWHGKHUH 0DQ\ DUHUHGXFWLRQLVWV DQG IDLO WR FRQVLGHU WKH YDULDEOH PRGLI\LQJ HIIHFWV RI IDFWRUV RWKHU WKDQ 895 WKDW DUH LQWLPDWHO\ LQYROYHG DQGHQYLURQPHQWDOO\UHOHYDQW +RZHYHU PRVWVWXGLHV SURJUHVVHGLQUHFHQW\HDUVIURPVLPSO\PHDVXULQJWKH HIIHFWV RI 89% RU 89$ DORQH LQ WKH ODERUDWRU\
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
$FNQRZOHGJPHQWV 5:& JUDWHIXOO\ DFNQRZOHGJHV WKH 86 1DWLRQDO 6FLHQFH )RXQGDWLRQ IRU VXSSRUW RIUHVHDUFK E\ ERWK DXWKRUVUHSRUWHG KHUH ) *3 DOVR DFNQRZOHGJHV WKH VXSSRUW RI WKH 0D[ 3ODQFN 6RFLHW\ DQG RI WKH %LRWHFKQRORJ\ 3URJUDP RI WKH (XURSHDQ &RPPLVVLRQ :H WKDQN 0 (KOLQJ6FKXO] 6 6FKHUHU ' 9LQHEURRNH DQG 3 /HDYLWW IRU PDNLQJ XQSXEOLVKHGGDWDDYDLODEOHWRXV
5HIHUHQFHV $GDPV 1 /DQG6FKLFN-0 0\FRVSRULQHOLNHDPLQRDFLGV SURYLGH SURWHFWLRQ DJDLQVW XOWUDYLROHW UDGLDWLRQ LQ HJJV RI WKH JUHHQ VHD XUFKLQ Strongylocentrotus droebachiensis. 3KRWRFKHP3KRWRELRO $USLQ 1 )DYUH%RPYLQ - DQG 7KLYHQG 6 6WUXFWXUH OD P\FRVSRULQH QRXYHOOH PROHFXOH LVROHH GH Botrytis cynerea. 7HWUDKHGURQ/HWW $VDWR < ,VRODWLRQ DQG FKDUDFWHUL]DWLRQ RIXOWUDYLROHW OLJKW VHQVLWLYH PXWDQWV RI WKH EOXHJUHHQ DOJD Anacystis nidulans. %DFWHULRO %HERXW %0 DQG *DUFLD3LFKHO ) 89%LQGXFHG YHUWLFDO PLJUDWLRQV RIF\DQREDFWHULD LQ D PLFURELDO PDW $SSO (QYLURQ 0LFURELRO %HQ$PRW]$ 6KDLVK $ DQG $YURQ 0 0RGH RIDFWLRQ RI WKHPDVVLYHO\DFFXPXODWHGFDURWHQHRIDunaliella burdawil LQ SURWHFWLQJ WKH DOJD DJDLQVW GDPDJH E\ H[FHVV LUUDGLDWLRQ 3ODQW 3K\VLRO
%HUQLOORQ- 3DUXVVVLQH( /HWRXEORQ5)DYUH%RQYLQ-DQG$USLQ 1 )ODYLQPHGLDWHG SKRWRO\VLV RI P\FRVSRQQHV 3K\WRFKHP %KDWWDFKDUMHH 6. DQG 'DYLG .$9 8QXVXDO UHVLVWDQFH WR XOWUDYLROHWOLJKW LQ GDUNSKDVHRIEOXHJUHHQEDFWHULXPAnacystis nidulans. 1DWXUH %KDWWDFKDUMHH 6. 0DWKXU 0 5DQH 66 DQG 'DYLG .$9 89VHQVLWLYLW\ RI F\DQREDFWHULXP Anacystis nidulans 3DUW , (YLGHQFH IRU SKRWRV\VWHP ,, 36,, DV D OHWKDO WDUJHW DQG FRQVWLWXWLYH QDWXUH RI D GDUN UHSDLU V\VWHP DJDLQVW GDPDJH WR 36,, ,QGLDQ-([SHU%LRO %MRUQ /2 DQG 0XUSK\70 &RPSXWHUFDOFXODWLRQ RIVRODU XOWUDYLROHWUDGLDWLRQDWJURXQGOHYHO3K\VLRO9HJ %ODNHILHOG0.DQG+DUULV '2 'HOD\ RIFHOOGLIIHUHQWLDWLRQ LQ$QDEDHQDDHTXDOLV FDXVHGE\89% UDGLDWLRQ DQG WKHUROHRI SKRWRUHDFWLYDWLRQ DQG H[FLVLRQUHSDLU 3KRWRFKHP3KRWRELRO %RKP*$ 3IOHLGHUHU: %RJHU3DQG 6FKHUHU6 6WUXFWXUH RI D QRYHO ROLJRVDFFKDULGHP\FRVSRULQHDPLQRDFLG XOWUDYLROHW $% VXQVFUHHQ SLJPHQW IURP WKH WHUUHVWULDO F\DQREDFWHULXP Nostoc commune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nabaena variabilis &\DQREDFWHULD - %DVLF 0LFURELRO %UHQRZLW]6 DQG&DVWHQKRO]5: /RQJWHUPHIIHFWVRI89 DQG YLVLEOH LUUDGLDQFH RQ QDWXUDO SRSXODWLRQV RI D VF\WRQHPLQ FRQWDLQLQJ F\DQREDFWHULXP Calothrix VS )(06 0LFURELRO (FRO %ULWWRQ * 6WUXFWXUH DQG SURSHUWLHV RI FDURWHQRLGV LQ UHODWLRQWRIXQFWLRQ)$6(%- %XFNOH\ &( DQG +RXJKWRQ -$ $ VWXG\ RI WKH HIIHFWV RI QHDU 89 UDGLDWLRQ RQ WKH SLJPHQWDWLRQ RI WKH EOXHJUHHQ DOJD Gloeocapsa alpicola. $UFK0LFURELRO %GHO % .DUVWHQ 8 DQG *DUFLD3LFKHO ) 8OWUDYLROHW DEVRUELQJ VF\WRQHPLQ DQG P\FRVSRULQHOLNH DPLQR DFLG GHULYDWLYHV LQ H[SRVHG URFNLQKDELWLQJ F\DQREDFWHULDO OLFKHQV 2HFRORJLD &DUU 1* DQG :\PDQ 0 &\DQREDFWHULD WKHLU ELRORJ\ LQ UHODWLRQWRRFHDQLFSLFRSODQNWRQ ,Q3ODWW7DQG/L:.:HGV 3KRWRV\QWKHWLF SLFRSODQNWRQ SS &DQ %XOO )LVK $TXDW 6FL &DUUHWR - , 'H0DUFR 6* DQG /XW] 9$ 89DEVRUELQJ SLJPHQWV LQ WKH GLQRIODJHOODWHV Alexandrium excavatum DQG Prorocentrum micans: (IIHFWV RIOLJKWLQWHQVLW\ ,Q 2NDLFKL 7 $QGHUVRQ '0 DQG 1HPRWR 7 HGV 5HG 7LGHV %LRORJ\ (QYLURQPHQWDO 6FLHQFH DQG 7R[LFRORJ\ SS 1HZ
&DVWHQKRO]5: 0LFURELDOPDWUHVHDUFK WKHUHFHQWSDVWDQG QHZ SHUVSHFWLYHV ,Q 6WDO /- DQG &DXPHWWH 3 HGV 0LFURELDO 0DWV SS 1$72 $6, 6HULHV 9RO 6SULQJHU9HUODJ %HUOLQ &DVWHQKRO] 5: -¡UJHQVHQ %% '¶$PHOLR ( DQG %DXOG - 3KRWRV\QWKHWLF DQG EHKDYLRUDOYHUVDWLOLW\ RIWKH F\DQREDFWHULXP Oscillatoria boryana LQ D VXOILGHULFK PLFURELDO PDW )(06 0LFURELRO (FRO &KLWQLV 35 DQG1HOVRQ1 0ROHFXODUFORQLQJ RIWKH JHQHV HQFRGLQJ WZR FKDSHURQH SURWHLQV RI WKH F\DQREDFWHULXP Synechocystis VS3&&-%LRO&KHP &OD\WRQ 5. /LJKW DQG /LYLQJ 0DWWHU 0F*UDZ+LOO 1HZ
5: &DVWHQKRO]DQG )*DUFLD3LFKHO )DYUH%RPYLQ - $USLQ 1 DQG %UHYDUG & 6WUXFWXUH GH OD P\FRVSRULQH3 &DQ-&KHP )DYUH%RPYLQ - %HUQLOORQ - 6DOLQ 1 DQG $USLQ 1 %LRV\QWKHVLV RI P\FRVSRULQHV P\FRVSRULQH JOXWDPLQRO LQ Trichotecium roseum. 3K\WRFKHPLVWU\ )UHGHULFN-( 6QHOO+(DQG+D\ZRRG(. 6RODUXOWUDYLROHW UDGLDWLRQ DWWKHHDUWK¶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hlorogloeopsis VSS 3KRWRFKHP3KRWRELRO *DUFLD3LFKHO ) :LQJDUG &( DQG &DVWHQKRO] 5: (YLGHQFHUHJDUGLQJWKH89VXQVFUHHQUROHRIDP\FRVSRULQHOLNH FRPSRXQG LQ WKH F\DQREDFWHULXP Gloeocapsa VS $SSO (QYLURQ0LFURELRO *DXWLHU&+H*DQG
&KDSWHU 5HVSRQVHVWR895 *UHWKHU70DLHU- .DUVWHQ8*DUFLD3LFKHO)DQG1LQQHPDQQ+ 89OLJKW LQGXFWLRQ RI ELRV\QWKHVLV RI SWHULGLQH JO\FRVLGHV RI F\DQREDFWHULD IURP PLFURELDO PDWV $EVWUDFW 89%OXH /LJKW 3HUFHSWLRQ DQG 5HVSRQVHV LQ 3ODQWV DQG 0LFURRUJDQLVP0DUEXUJ*HUPDQ\ +DGHU '3 (IIHFWV RI 89% RQ PRWLOLW\ DQG SKRWRRULHQWDWLRQ LQ WKH F\DQREDFWHULXP Phormidium uncinatum. $UFK0LFURELRO +DGHU'3DQG7HYLQL 0 *HQHUDO3KRWRELRORJ\ 3HUJDPRQ 3UHVV2[IRUG +DGHU '3 :DWDQDEH 0 DQG )XUX\D 0 ,QKLELWLRQ RI PRWLOLW\ LQ WKH F\DQREDFWHULXP Phormidium uncinatum, E\ VRODUDQGPRQRFKURPDWLF89LUUDGLDWLRQ 3ODQW&HOO3K\VLRO +DGHU'3:RUUHVW5&.XPDU+'DQG6PLWK 5& (IIHFWV RI LQFUHDVHG VRODU XOWUDYLROHW UDGLDWLRQ RQ DTXDWLF HFRV\VWHPV $PELR +DJHQ& %UDXQH:DQG %M¡UQ/2 )XQFWLRQDO DVSHFWV RI VHFRQGDU\FDURWHQRLGVLQHaematococcus lacustris 9ROYRFDOHV ,,,$FWLRQDVD³VXQVKDGH´ -3K\FRO +DUP : %LRORJLFDO (IIHFWV RI 8OWUDYLROHW 5DGLDWLRQ &DPEULGJH8QLYHUVLW\3UHVV&DPEULGJH +HOEOLQJ(: 9LOODIDxH9DQG+ROP+DQVHQ 2 (IIHFWVRI XOWUDYLROHW UDGLDWLRQ RQ $QWDUFWLF PDULQH SK\WRSODQNWRQ SKRWRV\QWKHVLV ZLWK SDUWLFXODU DWWHQWLRQ WR WKH LQIOXHQFH RI PL[LQJ ,Q :HLOHU &6 DQG 3HQKDOH 3$ HGV 8OWUDYLROHW 5DGLDWLRQ LQ $QWDUFWLFD 0HDVXUHPHQWV DQG %LRORJLFDO (IIHFWV 9RO SS $PHULFDQ *HRSK\VLFDO 8QLRQ :DVKLQJWRQ'& +LURVDZD 7 DQG 0L\DFKL 6 (IIHFW RI ORQJZDYHOHQJWK XOWUDYLROHW 89$ UDGLDWLRQ RQ WKH JURZWK RI Anacystis nidulans. 3ODQW6FL/HWW +ROP+DQVHQ /XELQ ' DQG +HLEOLQJ (: 8OWUDYLROHW UDGLDWLRQ DQG LWV HIIHFWV RQ RUJDQLVPV LQ DTXDWLF HQYLURQPHQWV ,Q
.DUHQW]'0F(XHQ)6/DQG0&DQG'XQODS:& 6XUYH\ RIP\FRVSRULQHOLNHDPLQRDFLG FRPSRXQGV LQ $QWDUFWLF PDULQH RUJDQLVPV SRWHQWLDO SURWHFWLRQ IURP XOWUDYLROHW H[SRVXUH 0DU %LRO .DUVWHQ 8 DQG *DUFLD3LFKHO ) &DURWHQRLGV DQG P\FRVSRULQHOLNH DPLQR DFLGV LQ PHPEHUV RI WKH JHQXV Microcoleus &\DQREDFWHULD DFKHPRV\VWHPDWLF VWXG\ 6\VWHP $SSO0LFURELRO .DVWLQJ -) +ROODQG +' DQG .XPS /5 $WPRVSKHULF HYROXWLRQ WKHULVHRIR[\JHQ ,Q 6FKRSI-:DQG.OHLQ& HGV 7KH 3URWHUR]RLF %LRVSKHUH D 0XOWLGLVFLSOLQDU\ 6WXG\ SS &DPEULGJH8QLY3UHVV&DPEULGJH .LUN-7 2SWLFV RI89%UDGLDWLRQLQQDWXUDO ZDWHUV $UFK +\GURELRO%HLK(UJHEQ/LPQRO .LUN -72 /LJKW DQG 3KRWRV\QWKHVLV LQ $TXDWLF (FRV\VWHPV&DPEULGJH8QLYHUVLW\3UHVV&DPEULGJH .ROOHU /5 8OWUDYLROHW 5DGLDWLRQ - :LOH\ DQG 6RQV 1HZ
0LOOHU 65 :LQJDUG &( DQG &DVWHQKRO] 5: (IIHFWV RI YLVLEOHOLJKWDQG89UDGLDWLRQRQSKRWRV\QWKHVLV LQDSRSXODWLRQ RIDKRWVSULQJ F\DQREDFWHULXP D Synechococcus VS VXEMHFWHG WR KLJK WHPSHUDWXUHVWUHVV $SSO (QYLURQ 0LFURELRO 0LORW5R\ 9 DQG 9LQFHQW :) 89 UDGLDWLRQ HIIHFWV RQ SKRWRV\QWKHVLV WKHLPSRUWDQFHRIQHDUVXUIDFHWKHUPRFOLQHVLQD VXEDUFWLFODNH $UFK+\GURELRO%HLK(UJEQ/LPQRO 0LWFKHOO '/ DQG .DUHQW] ' 7KH LQGXFWLRQ DQG UHSDLU RI '1$ SKRWRGDPDJH LQ WKH HQYLURQPHQW ,Q
5:&DVWHQKRO]DQG )*DUFLD3LFKHO 2ZWWULP *: DQG &ROHPDQ -5 5HJXODWLRQ RI H[SUHVVLRQ DQG QXFOHRWLGH VHTXHQFH RIWKH Anabaena variabilis recA JHQH -%DFWHULRO 3DHUO +: 7XFNHU - DQG %ODQG 37 &DURWHQRLG HQKDQFHPHQW DQG LWV UROH LQ PDLQWDLQLQJ EOXHJUHHQ DOJDO Microcystis aeruginosa VXUIDFH EORRPV /LPQRO 2FHDQ 3HQWHFRVW$ (IIHFWVRIVHGLPHQWDWLRQDQGOLJKWLQWHQVLW\RQ PDWIRUPLQJ 2VFLOODWRULDFHDH ZLWK SDUWLFXODU UHIHUHQFH WR Microcoleus lyngbyaceus *RPRQW - *HQ0LFURELRO 3HQWHFRVW $ )LHOG UHODWLRQVKLSV EHWZHHQ VF\WRQHPLQ GHQVLW\JURZWKDQGLUUDGLDQFHLQF\DQREDFWHULDRFFXUULQJLQORZ LOOXPLQDWLRQUHJLPHV 0LFURE(FRO 3LD]HQD + DQG +DGHU '3 3HQHWUDWLRQ RI VRODU 89 UDGLDWLRQ LQ FRDVWDO ODJRRQV RIWKH 6RXWKHUQ %DOWLF VHD DQG LWV HIIHFWRQSK\WRSODQNWRQFRPPXQLWLHV 3KRWRFKHP3KRWRELRO 3LHUVRQ%. 2HVWHUOH$ DQG0XUSK\ */ 3LJPHQWV OLJKW SHQHWUDWLRQ DQG SKRWRV\QWKHWLF DFWLYLW\ LQ WKH PXOWLOD\HUHG PLFURELDOPDWVRI*UHDW6LSSHZLVVHWW6DOW0DUVK 0DVVDFKXVHWWV )(06 0LFURELRO(FRO 3RVWLXV&(UQVW$ .HQWHU8DQG%RJHU3 3HUVLVWHQFHDQG JHQHWLF GLYHUVLW\ DPRQJ VWUDLQV RI SK\FRHU\WKULQULFK F\DQREDFWHULD IURP WKH SLFRSODQNWRQ RI /DNH &RQVWDQFH - 3ODQNWRQ5HV 3RWWV 0 'HVLFFDWLRQ WROHUDQFH RISURNDU\RWHV 0LFURELRO 5HY 3UH]HOLQ % % 1 3 %RXFKHU DQG 5 & 6PLWK 0DULQH SULPDU\ SURGXFWLRQ XQGHU WKH LQIOXHQFH RI WKH $QWDUFWLF R]RQH KROH ,FHFRORUV ,Q :HLOHU &6 DQG 3HQKDOH 3$ HGV 8OWUDYLROHW 5DGLDWLRQ LQ $QWDUFWLFD 0HDVXUHPHQW DQG %LRORJLFDO (IIHFWV 9RO SS $PHULFDQ *HRSK\VLFDO 8QLRQ:DVKLQJWRQ'& 3URWHDX 3- *HUZLFN :+ *DUFLD3LFKHO ) DQG &DVWHQKRO] 5: 7KHVWUXFWXUHRIVF\WRQHPLQDQXOWUDYLROHWVXQVFUHHQ SLJPHQW IURP WKH VKHDWKV RI F\DQREDFWHULD ([SHULHQWLD 5DPVLQJ 1% DQG 3UXIHUW%HERXW / 0RWLOLW\ RI Microcoleus chthonoplastes VXEMHFWHG WR GLIIHUHQW OLJKW LQWHQVLWLHV TXDQWLILHG E\GLJLWDO LPDJH DQDO\VLV ,Q 6WDO /- DQG &DXPHWWH3 HGV 0LFURELDO 0DWV 6WUXFWXUH 'HYHORSPHQW DQG (QYLURQPHQWDO 6LJQLILFDQFH SS 1$72 $6, 6HULHV 9RO 6SULQJHU9HUODJ %HUOLQ 5HQJHU * 9RONHU 0 (FNHUW +- )URPPH 5 +RKP9HLW 6 DQG *UlEHU 3 2Q WKH PHFKDQLVP RI SKRWRV\VWHP ,, GHWHULRUDWLRQE\89% LUUDGLDWLRQ 3KRWRFKHP3KRWRELRO 5HYVEHFK13 -¡UJHQVHQ %% %ODFNEXUQ 7+ DQG&RKHQ < 0LFURHOHFWURGH VWXGLHV RI WKH SKRWRV\QWKHVLV DQG 2 +6 DQG S+ SURILOHV RI D PLFURELDO PDW /LPQRO 2FHDQRJU 5LFKDUGVRQ // DQG &DVWHQKRO] 5: 'LHO YHUWLFDO PRYHPHQWV RIWKH F\DQREDFWHULXP Oscillatoria terebriformis LQ DVXOILGHULFKKRWVSULQJPDW $SSO(QYLURQ0LFURELRO 5RELQVRQ1 6RODU5DGLDWLRQ (OVHYLHU$PVWHUGDP 5RRV -& DQG 9LQFHQW:) 7HPSHUDWXUH GHSHQGHQFH RI89 UDGLDWLRQHIIHFWVRQ$QWDUFWLFF\DQREDFWHULD- 5R\&5*LHV+37RPOLQVRQ':DQG/XJJ'/ (IIHFWVRI R]RQH GHSOHWLRQ RQ WKH XOWUDYLROHW UDGLDWLRQ HQYLURQPHQW DW WKH $XVWUDOLDQVWDWLRQVLQ$QWDUFWLFD,Q:HLOHU&6DQG3HQKDOH3$ HGV 8OWUDYLROHW 5DGLDWLRQ LQ $QWDUFWLFD 0HDVXUHPHQWV DQG
&KDSWHU 5HVSRQVHVWR895 %LRORJLFDO (IIHFWV 9RO SS $PHULFDQ *HRSK\VLFDO 8QLRQ:DVKLQJWRQ'& 6DQGHUV % 6WUHVV SURWHLQV LQ DTXDWLF RUJDQLVPV DQ HQYLURQPHQWDOSHUVSHFWLYH &ULW5HY7R[LFRO 6DQGPDQQ * .XKQ 6 DQG %RJHU 3 (YDOXDWLRQ RI VWUXFWXUDOO\ GLIIHUHQW FDURWHQRLGV LQ Escherichia coli WUDQVIRUPDQWV DV SURWHFWDQWV DJDLQVW 89% UDGLDWLRQ $SSO (QYLURQ0LFURELRO 6DVV / 6SHWHD & 0iWp = 1DJ\ ) DQG 9DVV , 5HSDLU RI 89% LQGXFHG GDPDJH RISKRWRV\VWHP ,,YLD de novo V\QWKHVLV RIWKH ' DQG 'UHDFWLRQFHQWUH VXEXQLWV LQ Synechocystis VS 3&&3KRWRV\Q5HV 6DWRK . &UHDWLRQ RI SKRWRWROHUDQW PXWDQWV RI D F\DQREDFWHULXP Synechocystis VS 3&& E\ in vitro UDQGRPPXWDJHQHVLVRIWKHpsbA JHQH ,Q 6DWRK.DQG0XUDWD1 HGV 6WUHVV 5HVSRQVHV RI 3KRWRV\QWKHWLF 2UJDQLVPV SS (OVHYLHU6FLHQFH%9$PVWHUGDP 6DWRK . DQG 0XUDWD 1 HGV 6WUHVV 5HVSRQVHV RI 3KRWRV\QWKHWLF2UJDQLVPV(OVHYLHU6FLHQFH%9$PVWHUGDP 6FKHUHU 6 &KHQ 7: DQG %RJHU 3 $ QHZ 89$% SURWHFWLQJ SLJPHQW LQ WKH WHUUHVWULDO F\DQREDFWHULXP Nostoc commune. 3ODQW3K\VLRO 6FXOO\ 10 0F4XHHQ '- /HDQ '56 DQG &RRSHU :- +\GURJHQ SHUR[LGH IRUPDWLRQ WKH LQWHUDFWLRQ RI XOWUDYLROHW UDGLDWLRQ DQG GLVVROYHG RUJDQLF FDUERQ LQ ODNH ZDWHUV DORQJ D 1JUDGLHQW /LPQRO2FHDQRJU 6KLEDWD+ .DWVX\D% DQG 2FKLDL + 1HDU89 LUUDGLDWLRQ LQGXFHV VKRFN SURWHLQV LQAnacystis nidulans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nabaena VS DQG Nostoc carmium. %RWDQLFD$FWD 6LYDOLQJDP 30 ,NDZD 7
7\DJL 5 6ULQLYDV * 9\DV ' .XPDU $ DQG .XPDU +' 'LIIHUHQWLDOHIIHFWRIXOWUDYLROHW% UDGLDWLRQRQFHUWDLQPHWDEROLF SURFHVVHV LQ D FKURPDWLFDOO\ DGDSWLQJ Nostoc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
&KDSWHU &\DQRWR[LQV &UDZIRUG6'RZDQG8WKD\D.6ZRERGD %LRORJLFDO6FLHQFHV 8QLYHUVLW\RI:DUZLFN &RYHQWU\ 8. 6XPPDU\ , ,QWURGXFWLRQ ,, 3RLVRQLQJ,QFLGHQWV ,OO :KLFK 7D[D DUH 7R[LF" ,9 7\SHV RI &\DQREDFWHULDO7R[LQV $ 1HXURWR[LQV $QDWR[LQD +RPRDQRWR[LQD $QDWR[LQDV $SKDQRWR[LQV , DQG ,, 6D[LWR[LQDQG1HRVD[LWR[LQ % +HSDWRWR[LQV & &\WRWR[LQV 6F\WRSK\FLQV &\DQREDFWHULQ +DSDOLQGROH$ $FXWLSK\FLQ /\QJE\DWR[LQ (QGRWR[LQ /LSRSRO\VDFFKDULGH ' 2WKHU³7R[LQV´ 9 7R[LQ$QDO\VLV 0RXVH %LRDVVD\ $OWHUQDWLYH %LRDVVD\ 7LVVXH &XOWXUH&\WRWR[LFLW\ &KURPDWRJUDSKLF$QDO\VLV 0DVV 6SHFWURPHWU\ 3URWHLQ3KRVSKDWDVH,QKLELWLRQ$VVD\ ,PPXQRORJLFDO3URFHGXUHV 9, (FRORJLFDO,PSOLFDWLRQV $ +XPDQ+HDOWK 7R[LFRORJ\ 6KRUWWHUP(IIHFWV /RQJWHUP (IIHFWV 5HFUHDWLRQDO([SRVXUH % 7R[LFLW\ WR )LVK & 7R[LFLW\ WR ,QVHFWV 9,, &RQFOXGLQJ &RPPHQWV
B A Whitton and M Potts (eds). The Ecology of Cyanobacteria, Kluwer Acadenirc Publishers. Printed in the Netherlands
&6'RZDQG8.6ZRERGD
6XPPDU\ 7KH WR[LFLW\ RIVRPH F\DQREDFWHULD WR PDPPDOV KDV EHHQ NQRZQ IRU RYHU D FHQWXU\ EXW RQO\ UHFHQWO\ KDV WKH GLYHUVLW\ RIWR[LF RUJDQLVPV DQG WKH WR[LQV SURGXFHG E\ WKHP EHFRPH FOHDU 0RVW VWXGLHV KDYH EHHQ GRQH RQ IUHVKZDWHU DQG PDULQH EORRPIRUPLQJ VSHFLHV EXW EHQWKLF DQG IORDWLQJ IRUPV DUH VRPHWLPHV DOVR WR[LF 7KH WR[LQV LQFOXGH QHXURWR[LQV HJ DQDWR[LQD DQG VD[LWR[LQ KHSDWRWR[LQV RYHU YDULDQWV RIPLFURF\VWLQV DQG F\WRWR[LQV VF\WRSK\FLQ F\DQREDFWHULQ KDSDOLQGROH DFXWLSK\FLQ O\QJE\DWR[LQ 0RXVH ELRDVVD\V RI WR[LFLW\ DUH VWLOO FRQGXFWHG LQ PDQ\ FRXQWULHV EXW LQFUHDVLQJO\ RWKHU PHWKRGV DUH EHLQJ DGRSWHG WKHVH LQFOXGH VLPSOHU RUJDQLVPVRUWLVVXHVIRUWKHELRDVVD\V LPPXQRORJLFDOSURFHGXUHVYDULRXVFKURPDWRJUDSKLFWHFKQLTXHVDQGPDVV VSHFWURPHWU\ $OWKRXJK WKH WR[LQV DSSHDU LQ JHQHUDO WR EH PRUH WR[LF WR PDPPDOV LQFOXGLQJ KXPDQV WKDQ DTXDWLF RUJDQLVPV WKHUH LV H[SHULPHQWDO HYLGHQFH IRU WKHLU WR[LFLW\ WR YDULRXV DQLPDOV LQFOXGLQJ WKH \RXQJ VWDJHVRIVRPHILVKVSHFLHV ,QDGGLWLRQWKHWR[LFLW\RIPDQ\ ODNHSRSXODWLRQVRIF\DQREDFWHULDLVNQRZQWRKDYH DQHIIHFWRQ VRPHSRWHQWLDO ]RRSODQNWRQ JUD]HUV DQG WKH WR[LQV PD\ WKXV KDYH DQ LPSRUWDQW HIIHFW RQ WKH IRRG ZHELQDODNH
, ,QWURGXFWLRQ
,, 3RLVRQLQJ,QFLGHQWV
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rocambarus clarkii 9DVFRQFHORV 1HYHUWKHOHVV WKHUH LV HYLGHQFH IRUWKH LQIOXHQFH RIF\DQRWR[LQV RQ DUDQJH RI DTXDWLF RUJDQLVPV DQG VRPH H[DPSOHV DUH GHVFULEHG LQ WKLV FKDSWHU 7KH HFRORJLFDO DQG HFRQRPLF LPSDFWV RI WR[LQV VHHP OLNHO\ WR LQFUHDVH DV EORRPV LQFUHDVH LQ PDQ\ FRXQWULHV DV D UHVXOW RI QXWULHQWHXWURSKLFDWLRQ &DUPLFKDHO &RGG DQG %HDWWLH +XQWHU E &KRUXV DQG %DUWUDP
7R[LF F\DQREDFWHULDO EORRPV KDYH EHHQ UHSRUWHG IRU RYHU D FHQWXU\ RQH RI WKH ILUVW EHLQJ D Nodularia EORRP GHVFULEHG E\ )UDQFLV LQ Nature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icrocystis VFXPIURP 5XWODQG :DWHU (QJODQG ,Q DGGLWLRQ ZLQGVXUIHUV ZKR KDG FRPH LQWR FRQWDFW ZLWK WKLV VFXP FRPSODLQHG RIVNLQUDVKHV PRXWKEOLVWHUV DQG VHYHUH WKLUVW &RGG DQG %HDWWLH +XQWHU DQG 5REHUWV 3HDUVRQHWDO 6LPLODUO\VROGLHUVZKR KDG EHHQ RQ D FDQRHLQJ H[HUFLVH RQ 5XG\DUG /DNH (QJODQG DQG KDG FRPH LQWR FRQWDFW ZLWK D M. aeruginosa VFXP VXEVHTXHQWO\ VKRZHG VLJQV RI F\DQRWR[LFRVLV VRUH WKURDW KHDGDFKHV EOLVWHUHG PRXWK GLDUUKRHDDQG YRPLWLQJ 7ZR ZHUH DGPLWWHG WR KRVSLWDO ZLWK IHYHU DQG DW\SLFDO SQHXPRQLD 7XUQHUHWDO ,Q DW/RFK,QVK 6FRWODQG VHYHUDO GRJV GLHG DIWHULQJHVWLQJ DQHXURWR[LF Oscillatoria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
8QGHU WKHVH FRQGLWLRQV F\DQREDFWHULD RXWJURZ HXNDU\RWLF PLFURDOJDH 7KH IORDWLQJ PDVVHV IRUPHG E\ D QXPEHU RI VSHFLHV &KDSWHU FDQ EH FRQFHQWUDWHG E\ FXUUHQWV DQG OLJKW ZLQGV WR IRUP D WKLFN VFXP RQ WKH VKRUHV RI ODNHV DQG UHVHUYRLUV PDNLQJ WKH LQJHVWLRQ RI D IDWDO GRVH D UHDG\ SRVVLELOLW\ &DUPLFKDHO D E 0RVW SRLVRQLQJ LQFLGHQWV DUH WKHUHIRUH DVVRFLDWHG ZLWK EORRP IRUPLQJSODQNWRQLF VSHFLHV +RZHYHU WKH/RFK ,QVK LQFLGHQWV ZHUH FDXVHG E\ D QHXURWR[LF EHQWKLF Oscillatoria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aphnia VSHFLHV DQG WKH WR[LFLW\ RI Microcystis GRPLQDWHG EORRPV 7KH ODWWHU SURYLGHV IXUWKHU VXSSRUW IRU WKH K\SRWKHVLV WKDW F\DQRWR[LQV KDYH HYROYHG DV D GHIHQFH DJDLQVW JUD]HUV &DUPLFKDHO 'H0RWWHWDO /XNDFDQG$HJHUWHU 6LYRQHQHWDO +DQD]DWR 'LUHFW SURRI WKDW PLFURF\VWLQ LV UHVSRQVLEOH IRU DFXWH DQG ORQJ WHUPWR[LFRVLVWRDaphnia ZDV VKRZQ E\ FRPSDULVRQ RIWKHHIIHFWV RIDPLFURF\VWLQFRQWDLQLQJ VWUDLQ DQG D PXWDQW ODFNLQJ WKH PLFURF\VWLQ UHVSRQVLEOH IRU LWV IRUPDWLRQ 'LWWPDQQ HW DO 7KH Daphnia VKRZHG WKH VDPH GLJHVWLRQ UDWHV ZLWK WKH ZLOGW\SH DQGPXWDQWVWUDLQV 0DULQH F\DQREDFWHULD IURP LQWHUWLGDO DQG VXEOLWWRUDO ]RQHV Lyngbya, Schizothrix, Oscillatoria
&6 'RZ DQG 8. 6ZRERGD
Table 1. &RQILUPHGWR[LQSURGXFLQJVSHFLHVRIF\DQREDFWHULD 63(&,(6 Anabaena circinalis 5DEHQK A. flos-aquae /\QJE %UHE A. hassallii .W] :LWWU A. lemmermanni 5LFKWHU A. spiroides YDUFRQWUDFWD.OHE A. variabilis .W] Anabaenopsis milleri :RURQ Aphanizomenon flos-aquae / 5DOIV Coelosphaerium kuetzingianum 1DJ Cylindrospermum VS Cylindrospermopsis raciborskii :RORV 6HHQD\DHW6XEED5DMX Fischerella epiphytica *KRVH Gloeotrichia echinulata 5LFKWHU Gomphosphaeria lacustris &KRG G. naegliana 8QJHU /HPPHUP Hormothamnion enteromorphoides *UXQ Lyngbya majuscula +DUYH\ Lyngbya wollei Microcystis aeruginosa .W]
KDYH \LHOGHG D UDQJH RIWR[LF FRPSRXQGV DVVRFLDWHG ZLWKFRQWDFWGHUPDWLWLV 0RRUH EXWWKHH[WHQW WR ZKLFK RFHDQLF SODQNWRQLF F\DQREDFWHULD SURGXFH WR[LQV LV VWLOO XQFOHDU +RZHYHU EORRPV RI Trichodesmium KDYHEHHQLPSOLFDWHGLQWKHGHDWKVRI ILVK R\VWHUV DQG FUDEV DQG WKH DYRLGDQFH RI VXFK EORRPV E\ WXQD KDV EHHQ REVHUYHG &KLGDPEDUDP DQG 8QQ\ 1DJDEKXVKDQDP &KHOODP DQG$OJDUVZDPL (YDOXDWLRQRIWKHHFRORJLFDOLPSDFWRIWKHWR[LQVLV OLPLWHG +RZHYHU LW LV NQRZQ WKDW WKH\ FDQ LPSDFW XSRQ D ZLGH UDQJH RI DTXDWLF OLIH IRUPV ± ]RRSODQNWRQ 'H0RWW HW DO Daphnia /LJKWQHU Paramecium /DZWRQ DQG &RGG Artemia salina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
Microcystis wesenbergii .RPiUHNLQ.RQGUDWHYD Microcystis: RWKHUIRUPVRIXQFHUWDLQVWDWXVM. botrys, M. viridis) Nodularia spurnigena 0HUWHQV Nostoc linckia 5RWK %RUQHW)ODK N. paludosum .W] N. rivulare .W] N. zetterstedtii $UHVFKRXJ Oscillatoria acutissima .XII O. agardhiilrubescens JURXS Plunktothrix) O. formosa %RU\ O. nigroviridis 7KZDLWHV Oscillatoria VS Pseudanabaena catenata /DXWHUE Schizothrix calcicola $J *RP Scytonema pseudohofinunni %KDUDGZ Synechococcus VWUDLQV0LDPL%&,,6 $7&& Synechocystis VS Tolypothrix byssoidea +DVV .LUFKQ Trichodesmium erythraeum (KUE Umezakia natans +DUDGD
YDULDELOLW\ RI WR[LQ H[SUHVVLRQ GLVFXVVHG ODWHU ODERUDWRU\ DQG ILHOG GDWD VXSSRUW WKH YLHZ WKDW WKH PDMRULW\ RI F\DQREDFWHULDO VSHFLHV SURGXFH WR[LF FRPSRXQGV 7KH WR[LFRORJ\ RIWKHVH WKHLU PRGH RI V\QWKHVLV IXQFWLRQDO VLJQLILFDQFH DW WKH FHOOXODU DQG HFRORJLFDO OHYHO DQG WKH ULVN WR KXPDQ KHDOWK LQ SDUWLFXODUDUHDOOTXHVWLRQVDZDLWLQJDQVZHUV
,,, :KLFK7D[DDUH7R[LF" 7KRVH JHQHUD UHSRUWHG DV KDYLQJ WR[LF VSHFLHV DUH OLVWHGLQ7DEOH $WR[LJHQLF DQG WR[LJHQLF VWUDLQV KDYH EHHQ LVRODWHG IURP WKH VDPH EORRP EXW WKHUH LV OLWWOH LQIRUPDWLRQ DV WR WKH IDFWRUV WKDW UHJXODWH H[SUHVVLRQ LH D WR[LF VWUDLQ PD\ EHFRPH D QRQWR[LF VWUDLQ RU vice versa. 7R[LFLW\ QRW RQO\ YDULHV EHWZHHQ VWUDLQV EXW EHWZHHQ FORQHV RIWKH VDPH LVRODWH &DUPLFKDHO /XNDF DQG $HJHUWHU 8WNLOHQ DQG *MROPH ,Q DGGLWLRQ VRPH VWUDLQV SURGXFH WKUHH RU PRUH WR[LQV ZLWK WKH UHODWLYH SURSRUWLRQV EHLQJ LQIOXHQFHG E\ HQYLURQPHQWDO IDFWRUV VXFK DV OLJKW S+ QLWUDWH SKRVSKDWH PHWDO LRQV DQG WHPSHUDWXUH&DUPLFKDHO .HHYLO /XNDF DQG $HJHUWHU 6LYRQHQ :DWDQDEH DQG 2LVKL :LFNV DQG 7KLHO 8WNLOHQ DQG *MROPH
&KDSWHU &\DQRWR[LQV )LHOG GDWD RQ WKH LQIOXHQFH RI HQYLURQPHQWDO IDFWRUV RQ WR[LQ IRUPDWLRQ LQ SDUWLFXODU VSHFLHV KDYH WREHDVVHVVHGZLWKFDUH EHFDXVHDQ\REVHUYHGHIIHFW PD\ EH GXH WR YDU\LQJ LQIOXHQFHV RQ GLIIHUHQW VSHFLHV )RU LQVWDQFH D FRPSDULVRQ RI )LQQLVK ODNHV 9DLWRPDD HW DO LQGLFDWHG WKDW KLJKHU FRQFHQWUDWLRQV RI DTXHRXV 3 IDYRXUHG KHSDWRWR[LF Microcystis DQG DQDWR[LQD FRQWDLQLQJ Anabaena EORRPV ZKHUHDV Anabaena EORRPV ZLWK XQNQRZQ QHXURWR[LFLW\ ZHUH DVVRFLDWHG ZLWK ORZ 3 DQG KLJK QLWUDWH1
,9 7\SHVRI&\DQREDFWHULDO7R[LQ &\DQREDFWHULD DUHDULFKVRXUFH RIELRORJLFDOO\DFWLYH VHFRQGDU\ PHWDEROLWHV LH FRPSRXQGV WKDW DUH QRW XWLOLVHG E\ WKH RUJDQLVP V SULPDU\ PHWDEROLVP &DUPLFKDHO /XNDFDQG$HJHUWHU PDQ\ RI ZKLFK PD\ KDYH SKDUPDFHXWLFDO SRWHQWLDO 7KH PDMRULW\ RIF\DQRWR[LQV FDQSUREDEO\ EH UHJDUGHG DV DV VHFRQGDU\ PHWDEROLWHV 7KH\ PD\ EH FODVVLILHG LQWRWZRJHQHUDOJURXSV D 7KRVH FDXVLQJ DFXWH OHWKDO SRLVRQLQJ WKH neurotoxins DQG hepatotoxins; E 7KRVH ZKLFK DUH QRW KLJKO\ OHWKDO WR DQLPDOV EXW VKRZ PRUH VHOHFWLYH ELRDFWLYLW\ cytotoxins 7DEOH
$ 1HXURWR[LQV 1HXURWR[LQV DUH SURGXFHG E\ VHYHUDO JHQHUD HJ Anabaena, Aphanizomenon, Oscillatoria DQG Trichodesmium &DUPLFKDHO DE.HHYLO 7KH PDMRU QHXURWR[LQV DUH DQDWR[LQD KRPRDQDWR[LQD DQDWR[LQDV VD[LWR[LQ DQG QHRVD[LWR[LQDSKDQRWR[LQV,DQG,,
$QDWR[LQD $QDWR[LQDLVSURGXFHGE\VRPHVWUDLQV RIAnabaena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
+RPRDQDWR[LQD 7KLV WR[LQ KDV EHHQ SXULILHG IURP Oscillatoria formosa DQG FKDUDFWHULVHG DV D VHFRQGDU\ DPLQH DONDORLG PHWK\OHQHDQDWR[LQD 6LPLODU WR DQDWR[LQ DLWLVDSRWHQWQHXURPXVFXODUEORFNLQJDJHQWZLWKDQ LQWUDSHULWRQHDO /' LQ PLFH RI J NJ ERG\ ZHLJKW6NXOEHUJHWDO 7R[LFRVLVLQWKHOHWKDO GRVH UDQJH OHDGV WR VHYHUH ERG\ SDUDO\VLV FRQYXOVLRQVDQGGHDWKE\UHVSLUDWRU\IDLOXUH
$QDWR[LQDV $QDWR[LQDV WKH V GHQRWLQJ VDOLYDWLRQ LQ YHU WHEUDWHV LV SURGXFHG E\ VWUDLQV RI Anabaena flosaquae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phanizomenon flos-aquae 6DZ\HU HW DO &KURPDWRJUDSKLF DQG SKDUPDFRORJLFDO HYLGHQFH KDV
&6 'RZDQG8.6ZRERGD
Table 2. &\DQREDFWHULDOWR[LQV
72;,1
6758&785(
/'
Anabaena flos-aquae 15&&DQDGD 15&&DQDGD
$QDWR[LQ$ $QDWR[LQ$6
6HFRQGDU\DPLQHDONDORLG0: 1K\GUR[\JXDQDGLQH PHWK\OSKRVSKDWH HVWHU0:
Aphanizomenon flos-aquae 1+86$ 1+686$
$SKDQWR[LQQHRVD[LWR[LQ $SKDQWR[LQ,,VD[LWR[LQ
3XULQHDONDORLG0: 0:
0LFURF\VWLQV
+HSWDSHSWLGHV0:
&\DQRJLQRVLQV &\DQRJLQRVLQ 0LFURF\VWLQ 0LFURF\VWLQ 0LFURF\VWLQ 0LFURF\VWLQ 0LFURF\VWLQ &\DQRYLULGLQ 1RGXODULQ 0LFURF\VWLQV 0LFURF\VWLQV &\OLQGURVSHUPRSVLQ &\OLQGURVSHUPRSVLQ +RUPRWKDPQLQ$
+HSWDSHSWLGHV0: +HSWDSHSWLGH0: +HSWDSHSWLGH0: +HSWDSHSWLGH0: +HSWDSHSWLGH0: +HSWDSHSWLGH0: +HSWDSHSWLGH0:DQG +HSWDSHSWLGH0: 3HQWDSHSWLGH0: +HSWDSHSWLGHV0: +HSWDSHSWLGHV0:
63(&,(6 Neurotoxins
Hepatoxins Anabaena flos-aquae 6J&DQDGD Microcystis aeruginosa :56RXWK$IULFD :DWHUEORRP$XVWUDOLD :DWHUEORRP86$ 15&66 &DQDGD 6WUDLQ6FRWODQG :DWHUEORRP1RUZD\ 0-DSDQ Microcystis viridis Nodularia spumigena Oscillatoria agardhii YDU isothrix 1RUZD\ Oscillatoria agardhii 1RUZD\ Cylindrospermopsis raciborskii Aphanizomenon ovalisporum Hormothamnion enteronwrphoides Anabaena %4 Anabaena laxa
/D[DSK\FLQ$ /D[DSK\FLQ%
300-500
&\FOLFXQGHFDSHSWLGH &\FOLFGHFDSHSWLGHFRQWDLQLQJFKORULQH &\FOLFSHSWLGH0: &\FOLFSHSWLGH0:
Cytotoxins Scytonema pseudohofmanni %&+DZDLL
6F\WRSK\FLQ
Scytonema hofmanni 87(;7H[DV Hapalosiphon fontinalis 9 0DUVKDOO,VODQGV Tolypothrix byssoidea + Oscillatoria acutissima %O +DZDLL Fischerella muscicola 87(; Scytonema rnirabile %< Microcystis aeruginosa Microcystis wesenbergii
&\DQREDFWHULQ +DSDOLQGROH$ 7XEHUFLGLQ $FXWLSK\FLQ )LVFKHUHOOLQ
0HWK\OIRUPDPLGH$0: 0HWK\OIRUPDPLGH%0: &KORULQDWHGGLDU\OODFWRQH 6XEVWLWXWHGLQGROHDONDORLG 3\UURORS\ULPLGLQH 0DFUROLGH &\DQREDFWHULDOLQKLELWRU $QWLPLFURELDO
YRODWLOHVXOIXUFRPSRXQGV YRODWLOHVXOIXUFRPSRXQGV
JNJ ERG\ZHLJKW
HVWDEOLVKHG WKDW WKH DSKDQRWR[LQV VWURQJO\ UHVHPEOH VD[LWR[LQ DQG QHRVD[LWR[LQ 7KH DSKDQRWR[LQV LQWHUIHUH ZLWK QHXURWUDQVPLVVLRQ LH QHUYH LPSXOVHV E\ EORFNLQJ WKH QHXURQH VRGLXP FKDQQHOV DFURVV WKH D[RQ PHPEUDQH 6D[LWR[LQ KDV DOVR EHHQ UHFRUGHG IURP Lyngbya wollei &DUPLFKDHO HW DO DQ RUJDQLVPZKLFKIRUPV QXLVDQFH JURZWKV RQSRQGV LQ
VRXWKHDVW 86$ DQG D EORRPIRUPLQJ VWUDLQ RI Oscillatoria LVRODWHG IURP D EORRP LQ /DNH 9DUHVH ,WDO\ 3RPDWL HW DO 7KHVH WR[LQV DUH DOVR DVVRFLDWHG ZLWK D QXPEHU RI PDULQH GLQRIODJHOODWHV HJ Alexandrium tamarense, ZKLFK DUH UHVSRQVLEOH IRU SDUDO\WLF VKHOOILVK SRLVRQLQJ LQ KXPDQV DQG PDVVLYH GHDWKV RI PDULQH DQLPDOV DVVRFLDWHG ZLWK
&KDSWHU &\DQRWR[LQV
³UHG WLGHV´ LQ FRDVWDO ZDWHUV ZRUOGZLGH 7KHVH WR[LQV FDXVH V\PSWRPV VXFK DV LUUHJXODU EUHDWKLQJ ORVV RI FRRUGLQDWLRQ WZLWFKLQJ DQG GHDWK E\ UHVSLUDWRU\ IDLOXUH &DUPLFKDHO D +XQWHU D .HHYLO 7KH LQWUDSHULWRQHDO /' IRU PLFH XVLQJ EDWFKFXOWXUHG FHOOV LV DSSUR[LPDWHO\ PJNJ HDFKJUDPRIO\RSKLOL]HGFHOOV\LHOGV PJ DSKDQWR[LQ , DQG PJ DSKDQDWR[LQ ,, 0DKPRRG DQG&DUPLFKDHO
% +HSDWRWR[LQV &\FOLF KHSDWRWR[LF SHSWLGHV DUH WKH PRVW FRPPRQ RIIHQGHUV ZRUOGZLGH LQ FDVHV RIZDWHUERUQH GLVHDVH FDXVHG E\ WR[LF F\DQREDFWHULD 7KH PDMRU F\DQRWR[LQV FDXVLQJ GHDWK DQG LOOQHVV LQ DQLPDOV DUH WKH SHSWLGH KHSDWRWR[LQV OLYHU WR[LQV &DUPLFKDHO D .HHYLO /XXNNDLQHQHWDO 7KH\ DUH FODVVLILHG LQWR WKH F\FOLF KHSWDSHSWLGHV LH WKH PLFURF\VWLQV )LJ ZKLFK KDYH EHHQ LVRODWHG DQG FKDUDFWHULVHG IURP Microcystis DQGWKHILODPHQWRXV JHQHUDAnabaena (A. flos-aquae Nostoc N. rivulare DQG Oscillatoria O.
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
Fig 2. 7KH VWUXFWXUH RI QRGXODULQ 7KH ODUJHVW YDULDWLRQ LQ VWUXFWXUH IRU WKH F\FOLF KHSDWRSHSWLGHV KDV EHHQ VWXGLHG LQ WKH EUDFNLVK ILODPHQWRXV F\DQREDFWHULXP 1RGXODULD VSXPLJHQD &DUPLFKDHO +XQWHU D 5LQHKDUW HW DO DQG 6LYRQHQ HW DO KDYH FRQILUPHG WKH VWUXFWXUH IRU WKLV 1RGXODULD SHSWLGH DV D F\FOLF SHQWDSHSWLGH ILYH PHPEHUHG ULQJ 0: RI 'D DQG QDPHG LW QRGXODULQ 12'/1 7KH JHQHUDO VWUXFWXUH RIQRGXODULQ SURGXFHG E\ WKLV ILODPHQWRXV RUJDQLVP LV F\FOR '0H$VS/$UJ$GGD'*OX10H=GHK\GUREXW\ULQF
agardhii) &DUPLFKDHO D &RGG DQG %HDWWLH .HHYLO /XXNNDLQHQHWDO DQG WKH F\FOLF SHQWDSHSWLGH QRGXODULQ )LJ LVRODWHG IURP WKH EUDFNLVK ZDWHU F\DQREDFWHULXP Nodularia spumigena &RGG DQG %HDWWLH +XQWHU D .HHYLO /XXNNDLQHQ HW DO $ GHWDLOHG DFFRXQW RI WKH RFFXUUHQFH RI PLFURF\VWLQV LV JLYHQ E\ 3DUN DQG :DWDQDEH :DWDQDEH DQG RWKHU FKDSWHUV LQ WKH ERRN Toxic Microcystis HGLWHGE\:DWDQDEHHWDO $QRWKHU SHSWLGH WR[LQ F\OLQGURVSHUPRSVLQ KDV EHHQ LVRODWHG LQ $XVWUDOLD IURP Cylindrospermopsis raciborskii DQG DOVR IURP D UHFHQWO\ FKDUDFWHULVHG RUJDQLVP IURP -DSDQ Umezakia natans +DUDGD HW DO DQGAphanizomenon ovalisporum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manita phalloides, WKH JUHHQ GHDWK FDS )DXOVWLFK DQG :LHODQG /XNDF DQG $HJHUWHU .HHYLO :LHODQGDQG)DXOVWLFK 0LFURF\VWLQV KDYH WKH JHQHUDO VWUXFWXUH F\FOR' $OD/;HU\WKURPHWK\O'LVR$VS/=$GGD' LVR*OX1PHWK\OGHK\GUR$OD ZKHUH ; DQG = UHSUHVHQWFHUWDLQ/DPLQRDFLGV)LJ %RWHVHWDO &DUPLFKDHO HW DO /DQDUDV HW DO 5LQHKDUW HW DO 7KH SUHVHQFH RI$GGD > 6666 DPLQRPHWKR[\ WULPHWK\O SKHQ\OGHFDGLHQRLF DFLG@ LV WKH PRVW XQXVXDO VWUXFWXUDO IHDWXUH $GGD SOD\V DQ LPSRUWDQW UROH LQ KHSDWRWR[LFLW\ VLQFH K\GURJHQDWLRQ RU R]RQRO\VLV RI WKHGLHQHV\VWHPLQWKLVXQLWJLYHVDQLQDFWLYHSURGXFW +DUDGDHWDO D+DUDGDHWDO E 2YHU YDULDQWV RI PLFURF\VWLQV KDYH EHHQ FKDUDFWHULVHG 7DEOH VRPH RI WKH PRVW UHFHQW EHLQJ LVRODWHG IURP Oscillatoria agardhii >'KD@ 0&<67)5 /XXNNDLQHQ HW DO DQG >'$VS 'KE@ 0&<6755 6DQR DQG .D\D 'LPHWK\O PLFURF\VWLQV DSSHDU WR EH XQLTXH WR 6FDQGLQDYLDQF\DQREDFWHULDZLWK0&<67<5DQGLWV GLPHWK\O IRUPV KDYLQJ EHHQ IRXQG VR IDU RQO\ LQ Microcystis VSS /XXNNDLQHQ HW DO 'LIIHUHQFHVEHWZHHQWKHPLFURF\VWLQVDUHSULPDULO\LQ WKHW\SH RI/DPLQRDFLGSUHVHQW DQG LQ WKH SUHVHQFH RUDEVHQFHRID PHWK\O JURXS RQ RQH RIWKH 'DPLQR DFLGVRURQDGHK\GURDPLQRDFLGRUERWK 6WXGLHVRQ WKH ELRV\QWKHVLV RI PLFURF\VWLQV VXJJHVW D QRQ ULERVRPDO PHFKDQLVP RI V\QWKHVLV 7LOOHWW HW DO UHSRUWHG WKDW DNE JHQH FOXVWHUFORQHG DQG FKDUDFWHULVHG IURP Microcystis KDG VLJQLILFDQW VLPLODULW\ WR RWKHU SHSWLGH V\QWKHWDVHV DQG KDV D VSHFLILF UROH LQ PLFURF\VWLQ ELRV\QWKHVLV 7KH EDVLF GLIIHUHQFH EHWZHHQ KHSDWRWR[LF DQG QRQWR[LF VWUDLQV RI M. aeruginosa 3&& LV WKH SUHVHQFH RI RQH
&6 'RZDQG8.6ZRERGD PRUH JHQHV FRGLQJ IRU PLFURF\VWLQ V\QWKHWDVHV %RUQHUHWDO +\EULGL]DWLRQH[SHULPHQWVZLWK '1$IUDJPHQWV RIDNESHSWLGH V\QWKHWDVH RSHURQ RI KHSDWRWR[LF Anabaena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
& &\WRWR[LQV $OWKRXJK WKH QHXURWR[LQV DQG KHSDWRWR[LQV DUH WKH KLJKSURILOHF\DQRWR[LQVEHFDXVHRIWKHLUOHWKDOLW\DQG XELTXLW\ WKHUH DUH VHYHUDO OHVV ZHOO VWXGLHG F\DQRWR[LQV WKH PDMRULW\ RI ZKLFK DUH VHFRQGDU\ PHWDEROLWHV 2ILQWHUHVWLV WKH REVHUYDWLRQ WKDWPDQ\ KDYH DQWLDOJDO DQWLP\FRWLF RU DQWLEDFWHULDO DFWLYLW\ ZKLOVW RWKHUV DUH DFWLYH DJDLQVW FHOO WLVVXH OLQHV DQG KDYH PRGHUDWH DQWLWXPRXU DFWLYLWLHV *HUZLFN HW DO 3DWWHUVRQHWDO
6F\WRSK\FLQV 7KHVH DUH OLSRSKLOLF WR[LQV SURGXFHG E\ Scytonema pseudohofmanni, ZKLFKDUH PRGHUDWHO\WR[LF WRPLFH /' RI J NJ 7KH\ DOVR KDYH VWURQJ F\WRWR[LF DFWLYLW\ DJDLQVW FHOO FXOWXUHV HJ KXPDQ HSLGHUPRLG FDUFLQRPD DQG PRXVH ILEUREODVWV ,Q DGGLWLRQ WKH\ DUH DFWLYH DJDLQVW LQWUDSHULWRQHDOO\ LPSODQWHG O\PSKRF\WLF OHXNHPLD DQG OXQJ FDUFLQRPD 0RRUHHWDO &DUPLFKDHOHWDO
&KDSWHU &\DQRWR[LQV
Table 3. ([DPSOHV RI VWUXFWXUDOYDULDQWVRI PLFURF\VWLQDGDSWHGIURP&DUPLFKDHO
0:'D
)2508/$
Microcystis aeruginosa Microcystis aeruginosab Microcystis VSSF Microcystis aeruginosab Microcystis aeruginosab Anabaena flos-aquaeb Microcystis aeruginosab Oscillatoria agardhiic Microcystis VSSF Nostoc VSE Oscillatoria agardhiic Microcystis aeruginosab Oscillatoria agardhiic Microcystis VSSF Microcystis VSSF Microcystis aeruginosac Nostoc VSE Microcystis aeruginosab Microcystis viridisb Microcystis aeruginosab Microcystis viridisb Anabaena flos-aquaeb Anabaena flos-aquaeb Microcystis VSSF
&+12 &+12 &+12 &+12 &+12
&+12 &+12 &+12 &+12 &+12 &+12 &+12 &+12 &+126 &+126 &+12 &+12
&+12
&+12 &+12 &+12
Microcystis aeruginosab Anabaena flos-aquaeb Microcystis aeruginosab Anabaena flos-aquaeb Microcystis viridisc Microcystis aeruginosab Microcystis VSSF Nostoc VSE Nostoc VSE Microcystis VSSF Nostoc VSE Nostoc VSE Nostoc VSE Microcystis VSSF
&+12
&+12 &+12
&+12 &+12
&+12 &+12 &+12 &+12 &+12 &+12
0,&52&<67,1
25*$1,60
0&<67/$ 0&<67/DED 0&<67$5 0&<67<$ 0&<67/5 0&<67/< >'$VS'KD@ 0&<6755 >'KD@ 0&<67)5 >'$VS $'0$GGD¶@ 0&<67/KDU >'$VS@ 0&<6755 >'KD@ 0&<6755 >'KE@ 0&<6755 0&<67)5 0&<672 5 0&<67<02 >$'0$GGD@ 0&<67/KDU 0&<6755 0&<67<5 >'$VS@ 0&<67+W\5 0&<67+W\5 0&<67:5
b
MICROCYSTIN-LR 0&<67/5 >'$VS'KD@ 0&<67/5 >'$VS¶@ 0&<67/5 >'KD@ 0&<67/5 >'0$GGD@ 0&<67/5 >'$VS $'0$GGD@ 0&<67/5 >0VHU@ 0&<67/5 >$'0$GGD@ 0&<67/5 >'VHU $'0$GGD@ 0&<67/5 >$'0$GGD 0VHU@ 0&<67/5 >'*OX2&+2 @ 0&<67/5
.H\ D$ED DPLQRLVREXW\ULF DFLG 'KD GHK\GURDODQLQH '0$GGD OGLPHWK\O$GGD $'0$GGD ODFHW\OOGLPHWK\O$GGD 0VHU NPHWK\OVHULQH +DU KRPRDUJDQLQH 02 PHWKLRQLQH6R[LGH +W\ KRPRW\URVLQH E7R[LQV LVRODWHG IURP VWUDLQV RI FXOWXUH FROOHFWLRQV 7R[LQV LVRODWHG IURP EORRPV
&\DQREDFWHULQ 7KLV SURGXFW RI Scytonema hofmanni LV D FKORULQH FRQWDLQLQJ GLDU\OODFWRQH ZKLFK KDV DQWL F\DQREDFWHULDO DFWLYLW\ DV DFRQVHTXHQFH LW KDV EHHQ SURSRVHG DV D SRVVLEOH DOJLFLGH 0DVRQ HW DO *OHDVRQDQG3DXOVRQ
+DSDOLQGROH$ $F\WRWR[LF DONDORLGKDVEHHQLVRODWHG IURPVWUDLQVRI Hapalosiphon fontinalis. 7KLV OLSRSKLOLF FRPSRXQG KDV D EURDG UDQJH RI DQWLDOJDO DQG DQWLP\FRWLF DFWLYLW\0RRUHHWDO
$FXWLSK\FLQ 7ZR QRYHO PDFUROLGHV DFXWLSK\FLQ DQG GLGHK\GURDFXWLSK\FLQ KDYH EHHQ LVRODWHG IURP Oscillatoria acutissima %DUFKL HW DO 7KH\ DUH F\WRWR[LF WR WLVVXH FHOOV DQG KDYH DQWLWXPRXU SURSHUWLHVOXQJFDUFLQRPD
&6 'RZDQG8.6ZRERGD 7R[LFRORJLFDO VWXGLHV LQ WKH ODERUDWRU\ KDYH EHHQ FRQWUDGLFWRU\ VRPH KDYH VKRZQ QR HIIHFW :HLVHHWDO .HOHWLHWDO ZKLOVWRWKHUV KDYH EHHQ OHWKDO .HOHWL HW DO .HOHWL DQG 6\NRUD
' 2WKHU³7R[LQV´ &\DQREDFWHULD DOVR SURGXFH RWKHU VHFRQGDU\ PHWDEROLWHV IDOOLQJ ZLWKLQ WKH EURDG FDWHJRU\ RI F\DQRWR[LQV 7KHVH DUH FRPSRXQGV ZKLFK DUH WR[LJHQLF EXW ZKRVH IXQFWLRQ DQG HFRORJLFDO LPSRUWDQFH DUH LOOGHILQHG 'RXEWOHVV WKH OLVW RI WR[LJHQLF VXEVWDQFHV LGHQWLILHG IURP F\DQREDFWHULD ZLOO FRQWLQXH WR JURZ KRZHYHU WKHLU FRQWULEXWLRQ WR RU UROH LQ F\DQREDFWHULDO SK\VLRORJ\ JURZWK DQG VXUYLYDOLVDWSUHVHQWVNHWFK\
9 7R[LQ$QDO\VLV
/\QJE\DWR[LQ LV D SRWHQW WXPRXU SURPRWHU 1LVKL]XND DQG DFWLYDWHV FDOFLXPDFWLYDWHG SKRVSKROLSLGGHSHQGHQW SURWHLQ NLQDVH & FDXVLQJ FRQWUDFWLRQ RI YDVFXODU VPRRWK PXVFOH 5RELQVRQ HW DO ,W UHVHPEOHV WKH SKRUERO HVWHUV WXPRXU SURPRWLRQDQGDFWLYDWLRQRISURWHLQNLQDVH& DQGKDV EHHQ LVRODWHG IURPLyngbya majuscula )XMLNL HW DO DVSHFLHVIRUPLQJH[WHQVLYHIORDWLQJPDVVHVLQ VKDOORZ VKHOWHUHG ZDWHUV LQ ZDUPHU UHJLRQV /\QJE\DWR[LQ LV DOVR D SRWHQW LUULWDQW FDXVLQJ VZLPPHU¶V LWFK &DUGHOOLQD HW DO 0RRUH
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
(QGRWR[LQ/LSRSRO\VDFFKDULGH
0RXVH%LRDVVD\
&\DQRQREDFWHULDO OLSRSRO\VDFFKDULGHV LQ DGGLWLRQWR WKHLU HQGRWR[LF DFWLYLW\ KDYH EHHQ LPSOLFDWHG DV EHLQJ FRQWDFW LUULWDQWV ,Q FRPPRQ ZLWK *UDP QHJDWLYH EDFWHULD F\DQREDFWHULD IRUP OLSRSRO\ VDFFKDULGH DV DQ LPSRUWDQW SDUW RI WKHLU RXWHU FHOO HQYHORSH EXW XQOLNH WKH IRUPHU F\DQREDFWHULDO OLSRSRO\VDFFKDULGHODFNVDQ\SKRVSKDWHLQWKHOLSLG$ FRUH .HOHWL DQG 6\NRUD &RPSDUHG ZLWK HQWHUREDFWHULDO HQGRWR[LQV VXFK DV WKDW IURP Salmonella VSS F\DQREDFWHULDO OLSRSRO\VDFFKDULGH ZDV IRXQG WR EH WHQ WLPHV OHVV WR[LF LQ ELRDVVD\V &RGG 1HYHUWKHOHVVLWKDVEHHQLPSOLFDWHGLQ KXPDQ LOOQHVVHV %LOOLQJV &RGG DQG %HOO
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
/\QJE\DWR[LQ
&KDSWHU &\DQRWR[LQV GLVWLQFW IRU D FDWHJRU\ RI WR[LQ ,W LV GLIILFXOW WR GLIIHUHQWLDWHEHWZHHQFORVHO\UHODWHGWR[LQV
$OWHUQDWLYH %LRDVVD\V 7KH DPRXQW RI ]RRSODQNWRQ LV NQRZQ WR GHFUHDVH XQGHU F\DQREDFWHULDO EORRP FRQGLWLRQV /LJKWQHU FRQVHTXHQWO\ ELRDVVD\V KDYH EHHQ GHYLVHG XVLQJ Daphnia VS DQG Artemia salina .LYLUDQWD HW DO $ IHZVWXGLHV KDYHEHHQGRQH ZLWK RWKHU FUXVWDFHDQVDQGURWLIHUV*LOEHUW EXWQRWWDNHQ WR WKH H[WHQW ZKHUH WKH\ FDQ EH XVHG IRU ELRDVVD\V +RZHYHU D ELRDVVD\ KDV EHHQ GHYHORSHG IRU WKH IUXLWIO\ Drosophila melanogaster 6ZRGRGD DQG 'RZ DQG PLFURLQMHFWLRQ LQWR PRVTXLWRHV KDV DOVREHHQVXJJHVWHG7XUHOODQG0LGGOHEURRN 0DMRUGLIILFXOWLHVLQWKHXVHRIDOWHUQDWLYHELRDVVD\V VXFKDVWKRVHEDVHGRQArtemia salina DUHKLJKOLJKWHG E\ .LYLUDQWD HW DO 0RGHUDWH WR KLJK FRQFHQWUDWLRQV RI WR[LQ QHXUR DQG KHSDWRWR[LQV FDQ EH GHWHFWHG LQ EORRP VDPSOHV XVLQJ WKLV RUJDQLVP EXW GHWHFWLRQ RI WKH ORZHU FRQFHQWUDWLRQV IRXQG HDVLO\ E\ PRXVH ELRDVVD\ LV OHVV UHOLDEOH ,Q FRQWUDVW D ODERUDWRU\JURZQ VWUDLQ RI Oscillatoria agardhii ZDV WR[LF WR ODUYDH RIWKH PRVTXLWR Aedes aegyptii .LYLUDQWD DQG $EGHOKDPHHG Artemia salina DQG Daphnia pulex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
7LVVXH&XOWXUH&\WRWR[LFLW\ ,W LV ZHOO GRFXPHQWHG WKDW KHSDWRWR[LQV FDXVH GHIRUPDWLRQ DQG XOWUDVWUXFWXUDO FKDQJHV RI WKH F\WRVNHOHWRQ RI PRXVH OLYHU KHSDWRF\WHV SULPDULO\ GXH WR WKH LQKLELWLRQ RI SKRVSKDWDVH DFWLYLW\ 'DEKRONDUDQG &DUPLFKDHO (ULNVVRQ HW DO $XQH DQG %HUJ SURSRVHG WKH XVH RI IUHVKO\ SUHSDUHG UDW KHSDWRF\WHV WR VWXG\DVVHVV WKH WR[LFLW\ RIF\DQREDFWHULDO EORRPV 7KH\KDYHVKRZQ WKDW WKH in vitro WR[LFLW\ WR FHOOV FDQ EH FRUUHODWHG
ZLWK FRQWURO DQLPDO H[SHULPHQWV ZKHQ FUXGH DOJDO ELRPDVVLV XVHG 6LPLODU HIIHFWV KDYH EHHQ REVHUYHG ZLWK SHUPDQHQW FHOO OLQHV DQG HU\WKURF\WHV *UDERZ HWDO +RZHYHU GHVSLWH WKH UHPDUNDEOH WR[LF SRWHQWLDO RI KHSDWRWR[LQV in vivo, QR FHOO O\VLV OLEHUDWLRQ RIODFWDWH GHK\GURJHQDVH RUKDHPRO\VLV KDV EHHQ REVHUYHG DIWHU DSSOLFDWLRQ RI SXUH KHSDWRWR[LQ WR SULPDU\ RU SHUPDQHQW FHOO OLQHV 5XQQHJDU DQG )DOFRQHU (ULNVVRQHWDO 7KH DPELJXLW\ RIWKH GDWD IURP ELRDVVD\V PD\ EH LQFUHDVHGGXHWR VXEVWDQFHV RWKHUWKDQWKHUHFRJQLVHG F\DQRWR[LQVKDYLQJ F\WRWR[LFHIIHFWVRQFXOWXUHG FHOO OLQHV +HQQLQJHWDO IRXQG WKDWWKH F\WRWR[LF HIIHFWV RI FUXGH DTXHRXV H[WUDFWV IURP Microcystis aeruginosa 3&& DUH D FRQVHTXHQFH RI D N'D KHDWVHQVLWLYH VXEVWDQFH QRW PLFURF\VWLQ 'D KHDWUHVLVWDQW 7KLV VXEVWDQFH LV HIIHFWLYH DJDLQVW &KDQJ OLYHU &KLPSDQ]HH OLYHU FHOOV +H/D FHOOVDQG$IULFDQJUHHQPRQNH\FHOOV
&KURPDWRJUDSKLF$QDO\VLV ,QLWLDOO\ WKLQOD\HU FKURPDWRJUDSK\ 7/& ZDV XVHG WR SXULI\ WKH KHSDWRWR[LQV EXW ODFNHG VHQVLWLYLW\ DQG VSHFLILFLW\ 7KLV ZDV VXSHUFHGHG E\ KLJK SHUIRUPDQFH OLTXLG FKURPDWRJUDSK\ +3/& ZLWK ZKLFK LW KDV EHHQ SRVVLEOH WR LVRODWH KHSDWRWR[LQV IURP VHYHUDO F\DQREDFWHULDO VSHFLHV DQG HQYLURQPHQWDOELRPDVVZLWKKLJKVHQVLWLYLW\ %RWHVHW DO %URRNV DQG &RGG 6LYRQHQ +DUDGD 5HVROXWLRQ RIWKH WR[LF IUDFWLRQ IURP Microcystis aeruginosa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
0DVV6SHFWURPHWU\ 7KHVWUXFWXUDOFKDUDFWHULVWLFVRIWKHKHSDWRWR[LQV KDYH EHHQDVFHUWDLQHGSULPDULO\E\IDVWDWRPERPEDUGPHQW
&6 'RZDQG8.6ZRERGD
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
EORRPV EXW JDYH SRVLWLYH UHVSRQVHV ZLWK WR[LF Microcystis aeruginosa EORRPV %URRNV DQG &RGG 1DJDWD HW DO KDYH SURGXFHG PRQRFORQDO DQWLERGLHV DJDLQVW PLFURF\VWLQ/5 ZKLFK VKRZHG FURVVUHDFWLYLW\ ZLWK PLFURF\VWLQ55
3URWHLQ3KRVSKDWDVH,QKLELWLRQ$VVD\
$ +XPDQ+HDOWK
&RQILUPDWLRQ WKDW WKH KHSDWRWR[LF F\DQREDFWHULDO WR[LQV SURGXFH WKHLU WR[LF HIIHFWV WKURXJK WKH LQKLELWLRQRISURWHLQ SKRVSKDWDVHV DQG $ KDV ODLG WKH IRXQGDWLRQ IRU WR[LQ DVVD\V EDVHG RQ WKH LQKLELWLRQ RIWKHVH HQ]\PHV 6LP DQG 0XGJH 6ZRERGDHWDO 7KLVDVVD\LVDQHIIHFWLYHDQG UHOLDEOH PHDQV RI GHWHFWLQJ DOO KHSDWRWR[LF F\DQREDFWHULDO WR[LQV DQG KDV EHHQ HQKDQFHG E\ WKH GHYHORSPHQWRIDFRORULPHWULFSURFHGXUH:DUGHWDO
7R[LFRORJ\
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scillatoria RU Anabaena
9, (FRORJLFDO ,PSOLFDWLRQV
7KH 7ROHUDEOH 'DLO\ ,QWDNH 7', LH WKH GRVH OHYHO LQ KXPDQV WKDW LV FRQVLGHUHG WR EH ZLWKRXW DGYHUVH HIIHFWV ZKHQ WDNHQ GDLO\ RYHU D OLIHWLPH PXVW EH GHWHUPLQHG EHIRUH VDIH OHYHOV RI WR[LFDQWV RU FRQWDPLQDQWV FDQ EH DGYLVHG ,Q HVWDEOLVKLQJ WKHVH YDOXHV WKHUH DUH PDQ\ REVWDFOHV LQDGHTXDWH RU QRQ H[LVWDQW KXPDQ VWXGLHV WKH GLIIHUHQFH LQ VHQVLWLYLW\ EHWZHHQDQLPDOV DQGKXPDQV WKHIRUPHUEHLQJXVHG LQ ODERUDWRU\ VWXGLHV DQG FKDOOHQJHG ZLWK GLYHUVH SUHSDUDWLRQV RI WR[LQV WKH URXWHV RI H[SRVXUH PDQ\ WR[LQV DUH PRUH WR[LF ZKHQ DGPLQLVWHUHG YLD WKH LQWUDYHQRXV RU LQWUDSHULWRQHDO URXWH WKDQ E\ WKH RUDO URXWH WKH ODWWHU EHLQJ WKH PDLQ URXWH RI KXPDQ H[SRVXUH 7KH GDWD FXUUHQWO\ DYDLODEOH DUH LQVXIILFLHQW IRU WKH GHWHUPLQDWLRQ RI 7', YDOXHV IRU WKH JUHDW PDMRULW\ RI F\DQRWR[LQV $ IXUWKHU FRPSOLFDWLRQ LV LQWURGXFHG ZKHQ WKH VWUXFWXUDO YDULDQWV RI WKH KHSDWRWR[LQV DUH FRQVLGHUHG ZKLFK XVXDOO\ VKRZ YHU\ GLIIHUHQW WR[LFLWLHV LQ ODERUDWRU\ VWXGLHV (YLGHQFH IRUWKH DGYHUVHHIIHFWV RIF\DQRWR[LQV RQ KXPDQ KHDOWK FRPHV IURP WKUHH VRXUFHV HSLGHPLRORJ\SRLVRQLQJHYHQWV KXPDQDQG DQLPDO DQGWR[LFRORJLFDOVWXGLHV 6HYHUDO VWXGLHV RI KXPDQ SRSXODWLRQV KDYH SURGXFHG HSLGHPLRORJLFDO HYLGHQFH OLQNLQJ SRLVRQLQJ DQG RU LQMXU\ WR WKH SUHVHQFH RI F\DQRWR[LQV LQ GULQNLQJ RU UHFUHDWLRQDO ZDWHU +RZHYHU PRVW RI WKHVH VWXGLHV KDYH EHHQ UHWURVSHFWLYH DQG FRPSOHWH HSLGHPLRORJLFDO GDWD SDUWLFXODUO\ UHJDUGLQJ H[SRVXUH DUH UDUH 1HYHUWKHOHVV HSLGHPLRORJLFDO HYLGHQFH LV RI VSHFLDO LPSRUWDQFH LQ GHPRQVWUDWLQJ
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scillatoria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
WXPRXU SURPRWLRQ 7KH LPSOLFDWLRQV RI SURWHLQ SKRVSKDWDVH LQKLELWLRQ LQ KXPDQV GXH WR ORZ OHYHO FKURQLFH[SRVXUHWRPLFURF\VWLQVDUHQRWNQRZQ $W SUHVHQW WKH KXPDQ HYLGHQFH IRU PLFURF\VWLQ FDUFLQRJHQLFLW\LVLQDGHTXDWHDQGWKHDQLPDOHYLGHQFH LVOLPLWHG +RZHYHU WKH:+2 KDVSXEOLVKHG D SURYLVLRQDO 7', RI J NJ ERG\ ZW SHU GD\ DQG D SURYLVLRQDO JXLGHOLQH YDOXH RI J NJ ERG\ZWSHUGD\IRUPLFURF\VWLQ/5
5HFUHDWLRQDO([SRVXUH &RQWDFW ZLWK F\DQREDFWHULDLQ EDWKLQJ RUUHFUHDWLRQDO ZDWHUVFDQOHDGWRVNLQLUULWDWLRQ VZLPPHU V LWFK DQGLQFUHDVHGOLNHOLKRRGRIJDVWURLQWHVWLQDOV\PSWRPV 0RLNHKD DQG &KX +DVKLPRWR HW DO 3LORWWRHWDO 6HYHUHVNLQUHDFWLRQVKDYHEHHQ UHSRUWHG LQYROYLQJ WKH PDULQH F\DQREDFWHULXP Lyngbya majuscula, ZKLFK FDXVHV EOLVWHULQJ ZKHQ WUDSSHG EHQHDWK D EDWKLQJ FRVWXPH *UDXHU 7KLV RUJDQLVP H[SUHVVHV SRZHUIXO GHUPDO WR[LQV DSO\VLDWR[LQV DQG GHEURPRDSO\VLDWR[LQ 0\QGHUVH HW DO ZKLFK DUH SRWHQW WXPRXU SURPRWHUV DQG SURWHLQNLQDVH&DFWLYDWRUV)XMLNLHWDO Lyngbya majuscula PD\ JURZ HSLSK\WLFDOO\ RQ HGLEOHDOJDHHJAcanthophora spicifera. ,QDGGLWLRQ F\DQREDFWHULDKDYHDOVREHHQVXVSHFWHGWREHDVRXUFH RI WR[LQV LQ WKH FLJXDWHUD IRRG FKDLQ OHDGLQJ WR KXPDQSRLVRQLQJ +DKQ DQG&DSUD (QGHDQHW DO DQG Lynbya PLJKW WKHUHIRUH EH WKH DJHQW LQYROYHGKHUH
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ilapia DQG WURXW *DUFLD *DHWH HW DO %XU\HWDO +RZHYHU JLOO GDPDJH PD\ DOVR EH FDXVHG E\ WKH KLJK S+ YDOXHV DVVRFLDWHG ZLWK F\DQREDFWHULDO SKRWRV\QWKHVLV DQG WKH KLJK DPPRQLD FRQFHQWUDWLRQV DULVLQJ IURP WKH GHFRPSRVLWLRQ RI F\DQREDFWHULDO FHOOV ,UUHVSHFWLYHO\ RI WKLV JLOO GDPDJHDOPRVWFHUWDLQO\HQKDQFHVPLFURF\VWLQXSWDNH OHDGLQJWROLYHUQHFURVLV
&6'RZDQG8.6ZRERGD 7KH HIIHFWV RIPLFURF\VWLQV RQ WKH (XURSHDQ FDUS Cyprinus carpio, ZHUHDVVHVVHGE\FRPSDULVRQRIILVK IURP /DNH :HOOLQJWRQ QR HYLGHQFH RI VLJQLILFDQW JURZWK RI Microcystis aeruginosa) ZLWK ILVK IURP /DNH 0RNRDQ YLVLEOH M. aeruginosa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
& 7R[LFLW\WROQVHFWV 7KHUH DUH QR SXEOLVKHG GDWD RQ WKH HIIHFW RI F\DQREDFWHULDO EORRPV RQ LQVHFW SRSXODWLRQV LQ WKH QDWXUDO HQYLURQPHQW +RZHYHU WKH VHQVLWLYLW\ RI Daphnia %DLUG HW DO /DXUpQ0llWWl HW DO Locusta migratoria +LULSL HW DO Schistocerca gregaria 0FHOKLQH\ HW DO DQG RWKHU LQVHFWV WR D UDQJH RI F\DQREDFWHULDO WR[LQV SRLQWV WR HFRORJLFDO FRQVHTXHQFHV ,QVHFWFKDOOHQJHV DUH E\ LQJHVWLRQ RI F\DQREDFWHULDO FHOOV ZLWK PRUWDOLW\ VKRZLQJ D GRVH UHVSRQVH FXUYH .RWDN HW DO 7KH LPSOLFDWLRQV ZLWK UHVSHFW WR WKH DTXDWLFIRRGFKDLQKDYH\HWWREHDGGUHVVHG
&KDSWHU &\DQRWR[LQV 9,, &RQFOXGLQJ &RPPHQWV 'HVSLWH F\DQREDFWHULD KDYLQJ EHHQ UHFRJQLVHG IRU VRPHFRQVLGHUDEOHWLPHLWLVRQO\RYHUWKHSDVWWZHQW\ \HDUV WKDW WKHUH KDYH EHHQ VLJQLILFDQW DGYDQFHV LQ FKDUDFWHULVDWLRQ RIWKH WR[LF VSHFLHV DQG WKHLU WR[LQV per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
5HIHUHQFHV $QGHUVRQ 5- /XX +$ &KHQ '=; +ROPHV &)% .HQW 0 /H%ODQF07D\ORU)5-DQG:LOOLDPV'( &KHPLFDODQG ELRORJLFDO HYLGHQFH OLQNV PLFURF\VWLQV WR VDOPRQ ³1HWSHQ OLYHU 'LVHDVH´ 7R[LFRQ $VWUDFKDQ 1% DQG $UFKHU %* 6LPSOLILHG PRQLWRULQJ RI DQDWR[LQD E\ UHYHUVHSKDVH KLJK SHUIRUPDQFH OLTXLG FKURPDWRJUDSK\ DQG WKH VXEDFXWH HIIHFWV RIDQDWR[LQD LQ UDWV ,Q &DUPLFKDHO :: HG 7KH :DWHU (QYLURQPHQW $OJDO 7R[LQVDQG+HDOWK 3OHQXP3UHVV1HZ
%LOOLQJV :+ :DWHUDVVRFLDWHG KXPDQ LOOQHVV LQ QRUWKHDVW 3HQQV\OYDQLDDQGLWVVXVSHFWHGDVVRFLDWLRQZLWKEOXHJUHHQDOJDH EORRPV ,Q &DUPLFKDHO :: HG 7KH :DWHU (QYLURQPHQW $OJDO7R[LQVDQG+HDOWK 3OHQXP3UHVV1HZ
&6'RZDQG8. 6ZRERGD &KDLYLPRO-6ZRERGD8.DQG'RZ &6 D &KDUDFWHULVDWLRQ RI KHSDWRWR[LF IUHVKZDWHU Oscillatoria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richodesmium thiebautii DQG WKHLU HIIHFW RQ H[SHULPHQWDO SHDUO FXOWXUH DW 9HSSDORGDL ,QG-)LVK &KHQJDSSD003DFH/:DQG0FODXJKOLQ%* %OXHJUHHQ DOJDH Anabaena spiroides) WR[LFRVLV LQ SLJV - $P 9HW 0HG $66 &KLGDPEDUDP.DQG 8QQ\ 00 1RWH RQ WKH VZDUPLQJRI WKH SODQNWRQLF DOJD Trichodesmium erythraeum LQ WKH 3DPEDX DUHDDQGLWVHIIHFWRQWKHIDXQD &XUUHQW6FLHQFH &KRUXV , DQG %DUWUDP - HGV 7R[LF &\DQREDFWHULD LQ :DWHU $ *XLGH WR WKHLU 3XEOLF +HDOWK &RQVHTXHQFHV 0RQLWRULQJ DQG 0DQDJHPHQW ( )1 6SRQ /RQGRQ 1HZ
&KDSWHU &\DQRWR[LQV 6HSWHPEHU 7KH 5R\DO 6RFLHW\ RI &KHPLVWU\ &DPEULGJH (QGHDQ 5 0RQNV 6$ *ULIILWK -. DQG /OHZHOO\Q /( $SSDUHQW UHODWLRQVKLS EHWZHHQ WR[LQV HODERUDWHG E\ WKH F\DQREDFWHULXP Trichodesmium erythraeurn DQGWKRVHSUHVHQWLQ WKHIOHVKRIWKHQDUURZEDUUHGVSDQLVKPDFNHUHOScomberomorus commersoni. 7R[LFRQ (QYLURQPHQW $JHQF\ 7KH 'HWHUPLQDWLRQ RI 0LFURF\VWLQ $OJDO 7R[LQV LQ5DZ DQG7UHDWHG:DWHUV E\ +LJK 3HUIRUPDQFH /LTXLG&KURPDWRJUDSK\ 0HWKRGVIRUWKH([DPLQDWLRQ RI :DWHUV DQG $VVRFLDWHG 0DWHULDOV (QYLURQPHQW $JHQF\ RI (QJODQGDQG:DOHV%ULVWRO%68' SS (ULNVVRQ -( +DJHUVWUDQG + DQG ,VRPDD % &HOO VHOHFWLYH F\WRWR[LFLW\ RI D SHSWLGH IURP WKH F\DQREDFWHULXP Microcystis aeruginosa. %LRFKHP%LRSK\V (ULNVVRQ-(0HULOXRWR-$DQG/LQGKROP7 $FFXPXODWLRQ RISHSWLGH WR[LQ IURP WKH F\DQREDFWHULXP Oscillatoria agardhii LQWKHIUHVKZDWHUPXVVHOAnadonta cygnea. +\GURELRORJLD (ULNVVRQ-(7RLYROD' 0HULOXRWR-$2 .DUDNL + +DQ<* DQG +DUWVKRUQH ' +HSDWRF\WH GHIRUPDWLRQ LQGXFHG E\ F\DQREDFWHULDOWR[LQVUHIOHFWV LQKLELWLRQRISURWHLQSKRVSKDWDVHV %LRFKHP%LRSK\V5HV&RPP )DOFRQHU,5 7XPRXUSURPRWLRQDQGOLYHULQMXU\FDXVHGE\ RUDO FRQVXPSWLRQ RI F\DQREDFWHULD - (QYLURQ 7R[LFRO :DWHU 4XDO )DOFRQHU,5 +DUG\6- +XPSDJH$5)URVFLR 60 7R]HU*- DQG +DZNLQV35 +HSDWLFDQGUHQDOWR[LFLW\RIWKHEOXHJUHHQ DOJD F\DQREDFWHULXP Cylindrospemopsis raciborskii LQ PDOH 6ZLVV $OELQR PLFH - (QYLURQ 7R[LFRO :DWHU 4XDO LQ SUHVV )DOFRQHU,56PLWK-9-DFNVRQ$5%-RQHV$DQG5XQQHJDU07& 2UDO WR[LFLW\ RI D EORRP RI WKH F\DQREDFWHULXP Microcystis aeruginosa DGPLQLVWHUHG WRPLFHRYHUSHULRGV XSWR \HDU -7R[LFRO(QYLURQ+HDOWK )DXOVWLFK+DQG:LHODQG7 0XVKURRPSRLVRQV ,Q7X$7 HG )RRGSRLVRQLQJV+DQGERRNRI1DWXUDO7R[LQV 1HZ
*DUFLD *2 7R[LFLW\ RI WKH F\DQREDFWHULXP Microcystis aeruginosa VWUDLQ WR WURXW DQG WLODSLD D FOLQLFDO DQG KLVWRSDWKRORJLFDOVWXG\ 06F7KHVLV8QLYHUVLW\RI6WLUOLQJ8. *HUZLFN :+ 5REHUWV 3- 3URWHDX 3- DQG &KHQ -/ 6FUHHQLQJ FXOWXUHG PDULQH PLFURDOJDH IRU DQWLFDQFHUW\SH DFWLYLW\ -$SSO3K\FRO *LOEHUW -- 6XVFHSWLELOLW\ RI SODQNWRQLF URWLIHUV WR D WR[LF VWUDLQRIAnabaena flos-aquae. /LPQRO2FHDQRJU *OHDVRQ ). DQG 3DXOVRQ -3 6LWH RI DFWLRQ RIWKH QDWXUDO DOJLFLGH F\DQREDFWHULQLQWKHEOXHJUHHQDOJD Synchococcus VS $UFK0LFURELRO *UDERZ:2.'X5DQGW:&3UR]HVN\2:DQG6FRWW:( Microcystis aeruginosa WR[LQ FHOO FXOWXUH WR[LFLW\ KHPRO\VLV DQG PXWDJHQLFLW\ DVVD\V $SSO (QYLURQ 0LFURELRO *UDXHU) 6HDZHHGGHUPDWLWLV $UFK'HUPDWRO *XQQ *- 5DIIHUW\ $*5DIIHUW\ *& &RFNEXUQ 1 (GZDUGV & %HDWWLH .$ DQG &RGG *$ )DWDO FDQLQH QHXURWR[LFRVLV DWWULEXWHG WR EOXHJUHHQ DOJDH F\DQREDFWHULD 9HW5HF +DKQ 67 DQG &DSUD0) 7KH F\DQREDFWHULXP Oscillatoria erythraea – D SRWHQWLDO VRXUFH RI WR[LQ LQ WKH FLJXDWHUD IRRG FKDLQ )RRG$GG&RQWDP +DQD]DWR 7 7R[LF F\DQREDFWHULD DQG WKH ]RRSODQNWRQ FRPPXQLW\ ,Q :DWDQDEH0) +DUDGD+ &DUPLFKDHO::DQG )XMLNL + HGV 7R[LF Microcystis. &5& 3UHVV /RQGRQ +DUDGD . &KHPLVWU\ DQG GHWHFWLRQ RI PLFURF\VWLQV ,Q :DWDQDEH 0) +DUDGD + &DUPLFKDHO :: DQG )XMLNL + HGV 7R[LFMicrocystis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mezakia natans DQG LWV VFUHHQLQJ PHWKRG 7R[LFRQ +DVKLPRWR < .DPL\D +
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chizothrix calcicola $J *RPRQW F\DQREDFWHULD $SSO(QYLURQ0LFURELRO .ILU5 -RKDQQVHQ (DQG %RWHV '3 0RQRFORQDO DQWLERG\ VSHFLILF IRU F\DQRJLQRVLQ/$ SUHSDUDWLRQ DQG FKDUDFWHULVDWLRQ 7R[LFRQ .LYLUDQWD-DQG$EGHOEDPHHG$ 7R[LFLW\RIWKHEOXHJUHHQ DOJD Oscillatoria agardhii WR WKH PRVTXLWR Aedes aegyptii DQG WKH VKULPS Artemia salina. :RUOG - 0LFURELRO %LRWHFKQRO .LYLUDQWD - 1DPLNRVKL 0 6LYRQHQ . (YDQV :5 &DUPLFKDHO :: DQG 5LQHKDUW ./ 6WUXFWXUH GHWHUPLQDWLRQ DQG WR[LFLW\ RI D QHZ PLFURF\VWLQ IURP Microcystics aeruginosu. 7R[LFRQ .LYLUDQWD - 6LYRQHQ . DQG 1LHPHOD 6, 'HWHFWLRQ RI WR[LFLW\RIF\DQREDFWHULDE\Artemia salina ELRDVVD\ -(QYLURQ 7R[LFRO:DWHU4XDO .RWDN %* =XUDZHOO 5: 3UHSDV (( DQG +ROPHV &)% 0LFURF\VWLQ/5FRQFHQWUDWLRQLQDTXDWLFIRRGZHEFRPSDUWPHQWV IURPODNHV RIYDU\LQJWURSKLFVWDWXV &DQ- )LVK $TXDW6FL .XLSHU*RRGPDQ 7 )DOFRQHU , DQG )LW]JHUDOG - +XPDQ KHDOWKDVSHFWV ,Q &KRUXV ,DQG%DUWUDP-HGV 7R[LF &\DQREDFWHULD LQ :DWHU $ *XLGH WR 7KHLU 3XEOLF +HDOWK &RQVHTXHQFHV0RQLWRULQJDQG0DQDJHPHQW ( )1 6SRQ /RQGRQ 1HZ
&6 'RZDQG8.6ZRERGD /XXNNDLQHQ5 6LYRQHQ . 1DPLNRVKL 0 )DUGLJ 0 5LQHKDUW ./ DQG 1LHPHOD 6, ,VRODWLRQ DQG LGHQWLILFDWLRQ RI PLFURF\VWLQVIURP Oscillatoria agardhii VWUDLQVDQG VWUXFWXUH RIDQHZPLFURF\VWLQ $SSO(QYLURQ0LFURELRO 0DF.LQWRVK & %HDWWLH .$ .OXPSS 6 &RKHQ 3 DQG &RGG *$ &\DQREDFWHULDO PLFURF\VWLQ/5 LV DSRWHQWDQG VSHFLILF LQKLELWRURISURWHLQSKRVSKDWDVHV DQG $ IURPERWK PDPPDOV DQGKLJKHUSODQWV )HG(XU%LRFKHP62& /HWW 0DKPRRG 1$ DQG &DUPLFKDHO :: 3DUDO\WLF VKHOOILVK SRLVRQV SURGXFHG E\ WKH IUHVKZDWHU F\DQREDFWHULXP Anabaena flos-aquae 1+ 7R[LFRQ 0DVRQ &3 (GZDUGV .5 &DUOVRQ 5( 3LJQDWHOOR - *OHDVRQ ). DQG:RRG-0 ,VRODWLRQRIFKORULQHFRQWDLQLQJDQWLELRWLF IURP WKH IUHVKZDWHU F\DQREDFWHULXP Scytonema KRIPDQQL 6FLHQFH 0DWVXQDJD 6 0RRUH 5( 1LHPF]XUD :3 DQG &DUPLFKDHO :: $QDWR[LQDV D SRWHQW DQWLFKROLQHVWHUDVH IURP Anabaenaflos-aquae. -$PHU&KHP62& 0F(OKLQH\ - /DZWRQ /$ (GZDUGV & DQG *DOODFKHU 6 'HYHORSPHQW RI D ELRDVVD\ HPSOR\LQJ WKH GHVHUW ORFXVW (Schistocera gregaria) IRUWKHGHWHFWLRQ RIVD[LWR[LQ DQGUHODWHG FRPSRXQGVLQF\DQREDFWHULDDQGVKHOOILVK 7R[LFRQ 0HULOXRWR-$26DQGVWURP$(ULNVVRQ-(5HPDXG**UH\&UDLJ $ DQG &KDWWRSDGK\D\D - 6WUXFWXUH DQG WR[LFLW\ RI D SHSWLGH KHSDWRWR[LQ IURP WKH F\DQREDFWHULXP Oscillatoria agardhii. 7R[LFRQ 0RKDPPHG$0 -DPHO$O/D\ .6DQG-DPDO$O/DLO 66 7R[LF HIIHFWV RI WKH IUHVKZDWHU F\DQREDFWHULXP Oscillatoria agardii EOXHJUHHQ DOJDH RQ WKH VQDLO Melanoides tuberculata 0XOOHU -(J\SW*HU62& =RRO 0RLKHND 61 DQG &KX *: 'HUPDWLWLVSURGXFLQJ DOJD Lyngbya majuscula *RPRQWLQ+DZDLL ,, %LRORJLFDOSURSHUWLHV RIWKHWR[LFIDFWRU -3K\FRO 0RRUH 5( &RQVWLWXHQWV RI WKH EOXHJUHHQ DOJDH ,Q 6FEHXHU 3- HG 0DULQH 1DWXUDO 3URGXFWV &KHPLFDO DQG %LRORJLFDOSHUVSHFWLYHV$FDGHPLF3UHVV1HZ+@ O\QJE\DWR[LQ $ DQG >+@ GHEURPRSO\VLDWR[LQ WR WKH SKRUKROHVWHUUHFHSWRULQDPRXVHHSLGHUPDO SDUWLFXODWH IUDFWLRQ &DUFLQRJHQHVLV 7: 0\QGHUVH -6 0RRUH 5( .DVKLZDJL 0 DQG 1RUWRQ 75 $QWLOHXNHPLD DFWLYLW\ LQ WKH 2VFLOODWRULDFHDH LVRODWLRQ RI GHEURPRDSO\VLDWR[LQIURPLyngbya. 6FLHQFH 1DJDEKXVKDQDP$. 2QDQXQXVXDOO\GHQVHSK\WRSODQNWRQ EORRPDURXQG0LQLFR\LVODQG$UDELDQVHD DQGLWVHIIHFWVRQWKH ORFDOWXQDILVKHULHV &XUUHQW6FLHQFH 1DJDWD 6 6RXWRPH + 7VXWVXPL 7 +DVHJDZD $ 6HNLMLPD 0 6XJDPDWD 0 +DUDGD ., 6XJDQXPD 0 DQG 8HQR < 1RYHO PRQRFORQDO DQWLERGLHV DJDLQVW PLFURF\VWLQ DQG WKHLU SURWHFWLYHDFWLYLW\IRUKHSDWRWR[LFLW\ 1DWXUDO7R[LQV 1DPLNRVKL 0 5LQHKDUW ./ Sakai 5 6WRWWV 55 'DKOHP $0 %HDVOH\ 95 &DUPLFKDHO :: DQG (YDQV :5 ,GHQWLILFDWLRQ RI KHSDWRWR[LQV IURP D +RPHU /DNH EORRP RI WKH F\DQREDFWHULD Microcystis aeruginosa, Microcystis viridis DQG Microcystis wesenbergii: QLQH QHZ PLFURF\VWLQV - 2UJ &KHP 1LVKLZDNL0DWVXVKLPD 5 2KWD 1LVKLZDNL 6 6XJDZDPD 0 .RK\DPD. ,VKLNDZD7 &DUPLFKDHO :: DQG )XMLNL +
&KDSWHU &\DQRWR[LQV /LYHUWXPRXU SURPRWLRQ E\ WKH F\DQREDFWHULDO F\FOLF SHSWLGH WR[LQPLFURF\VWLQ/5 -&DQFHU5HV&OLQ2QFRO 2EHUHPP$%HFNHU- &RGG*$DQG 6WHLQEHUJ& (IIHFWV RI F\DQREDFWHULDO WR[LQV DQG DTXDHRXV FUXGH H[WUDFWV RI F\DQREDFWHULD RQ WKH GHYHORSPHQW RI ILVK DQG DPSKLELDQV (QYLURQ7R[LFRO 3DUN +' DQG :DWDQDEH 0) 7R[LF Microcystis LQ HXWURSKLFODNHV ,Q :DWDQDEH0) +DUDGD+&DUPLFKDHO:: DQG )XMLNL + HGV 7R[LF Microcystis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rtemia salina DQG Daphnia pulex. $UFK +\GURELRO 5HSDYLFK:0 6RQ]RJQL :& 6WDQGULGJH-+ :HGHSRKO 5( DQG 0HLVQHU /) &\DQREDFWHULD EOXHJUHHQDOJDH LQ :LVFRQVLQ :DWHUV ± DFXWH DQG FKURQLF WR[LFLW\ :DW5HV 5LQHKDUW ./ +DUDGD ., 1DPLNRVKL 0 &KHQ & +DUYLV &$ 0XQUR 0*+ %OXQW -: 0XOOLJDQ 3( %HDVOH\ 95 'DKOHP $0 DQG &DUPLFKDHO :: 1RGXODULQ PLFURF\VWLQ DQG WKHFRQILJXUDWLRQRI$GGD -$PHU&KHP6RF 5LQHKDUW ./ 1DPDNRSVKL DQG &KRL %: 6WUXFWXUH DQG ELRV\QWKHVLV RIWR[LQV IURPEOXHJUHHQ DOJDH F\DQREDFWHULD $SSO3K\FRO 5RELQVRQ &3 )UDQ] '5 DQG %RQGXUD 0( (IIHFWV RI O\QJE\DWR[LQ$ IURPWKHEOXHJUHHQDOJDLyngbya majuscula RQ UDEELWDRUWDFRQWUDFWLRQV 7R[LFRQ 5RGJHU +' 7XUQEXOO 7 (GZDUGV & DQG &RGG *$ &\DQREDFWHULDO EORRP DVVRFLDWHG SDWKRORJ\ LQ EURZQ WURXW Salmo trutta /LQ/RFK/HYHQ6FRWODQG -)LVK'LV 5RXKLDLQHQ/6LYRQHQ.3DXOLQ/ %XLNHPD:DQG+DVHONRUQ5 3HSWLGH V\QWKHWDVH JHQHV LQ Anabaena F\DQREDFWHULXP ,Q &DUPLFKDHO::DQG3DHUO+:HGV $EVWUDFWVWK ,QWHUQDWLRQDO &RQIHUHQFH RQ 7R[LF &\DQREDFWHULD %HDXIRUW 1RUWK&DUROLQD 5XQQHJDU 0 %HPGW 1 DQG .DSORZLW] 1 0LFURF\VWLQ XSWDNH DQG LQKLELWLRQ RI SURWHLQ SKRVSKDWDVH HIIHFWV RI FKHPRSURWHFWDQWVDQGVHOILQKLELWLRQLQUHODWLRQWRNQRZQKHSDWLF WUDQVSRUWHUV 7R[LFRO$SSO3KDUPDFRO 5XQQHJDU07& DQG )DOFRQHU,5 7KH in vivo DQG in vitro ELRORJLFDOHIIHFWVRIWKHSHSWLGHKHSDWRWR[LQIURPWKHEOXHJUHHQ DOJD Microcystis aeruginosa. 6RXWK$IU-6FL 5XQQHJDU07& DQG)DOFRQHU ,5 (IIHFWRIWR[LQ IURP WKH F\DQREDFWHULXP Microcystis aeruginosa RQ XOWUDVWUXFWXUDO
PRUSKRORJ\ DQG DFWLQ SRO\PHUL]DWLRQ LQ LVRODWHG KHSDWRF\WHV 7R[LFRQ 5XQQHJDU07& )DOFRQHU ,5 DQG 6LOYHU- 'HIRUPDWLRQ RI LVRODWHGUDWKHSDWRF\WHV E\ DSHSWLGHKHSDWRWR[LQ IURPWKH EOXH JUHHQDOJDMicrocystis aeruginosa. 1DXQ\Q6FKPLHGHEHUJV$FK 3KDUPDFRO 5XQQHJDU 07& .RQJ 60 DQG %HPGW 1 3URWHLQ SKRVSKDWDVH LQKLELWLRQ DQG in vivo KHSDWRWR[LFLW\ RI PLFURF\VWLQV$P-3K\VLRO** 6DQGVWURP $ *OHPDUHF & 0HULOXRWR -$2 (ULNVVRQ -( DQG &KDWWRSDGK\D\D - 6WUXFWXUH RI D KHSDWRWR[LF SHQWDSHSWLGH IURP WKH F\DQREDFWHULXP Nodularia spumigena. 7R[LFRQ 6DQR7DQG.D\D . $ DPLQREXWHQRLF DFLG '+% FRQWDLQLQJ PLFURF\VWLQ LVRODWHG IURP Oscillatoria agardhii. 7HWUDKHGURQ/HWWV 6DZ\HU3-*HQWLOH-+DQG6DVQHU---U 'HPRQVWUDWLRQRID WR[LQ IURP Aphanizomenon flos-aquae / 5DOIV &DQ - 0LFURELRO 6HDZULJKW $$ 1RODQ && 6KDZ *5 &KLVZHOO 5. 1RPV 5/ 0RRUH05DQG6PLWK0- 7KH RUDO WR[LFLW\ IRU PLFH RI WKH WURSLFDO F\DQREDFWHULXP Cylindrospermopsis raciborskii :RORV]\QVND (QY7R[LFRO:DWHU4XDOLQSUHVV 6LP $75 DQG 0XGJH /0 'HWHFWLRQ RI KHSDWRWR[LQV E\ SURWHLQ SKRVSKDWDVH LQKLELWLRQ DVVD\ DGYDQWDJHV SLWIDOOV DQG DQRPDOLHV ,Q&RGG*$-HIIULHV70.HHYLO&: DQG3RWWHU( HGV 'HWHFWLRQ 0HWKRGV IRU &\DQREDFWHULDO 7R[LQV 3URFHHGLQJV RIWKH )LUVW ,QWHUQDWLRQDO 6\PSRVLXP RQ 'HWHFWLRQ 0HWKRGV IRU&\DQREDFWHULDO7R[LQV 6HSWHPEHU 7KH5R\DO6RFLHW\RI&KHPLVWU\&DPEULGJH 6LYRQHQ . 7R[LF &\DQREDFWHULD LQ )LQQLVK )UHVK :DWHUV DQG7KH%DOWLF6HD 3K'7KHVLV8QLYHUVLW\RI+HOVLQNL 6LYRQHQ.DQG-RQHV * &\DQREDFWHULDOWR[LQV ,Q&KRUXV ,DQG%DUWUDP- HGV 7R[LF&\DQREDFWHULDLQ:DWHU $ *XLGH WR WKHLU 3XEOLF +HDOWK &RQVHTXHQFHV 0RQLWRULQJ DQG 0DQDJHPHQW ( )16SRQ/RQGRQ 1HZ
6WHYHQV '. DQG .ULHJHU 5, $QDO\VLV RI DQDWR[LQD E\ *&('& -$QDO\W7R[LFRO 6ZRERGD 8.DQG 'RZ&6 ([SUHVVLRQ RIF\DQRWR[LQV LQ HQYLURQPHQWDO ELRPDVV FRQWDLQLQJ VSHFLHV RI Oscillatoria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icrocystis aeruginosa SHSWLGH WR[LQ WR \HDUOLQJ UDLQERZ WURXW Oncorhynchus mykiss $TXD7R[LFRO 7LOOHWW''LWWPDQQ(%|UQHU7DQG1HLODQ%$ 6WUXFWXUDO DQDO\VLV RIWKHMicrocystis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icrocystis aeruginosa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icrocystis. &5&3UHVV/RQGRQ :DWDQDEH 0) +DUDGD + &DUPLFKDHO :: DQG )XMLNL + HGV 7R[LFMicrocystis. &5&3UHVV/RQGRQSS
&6 'RZDQG8.6ZRERGD :DWDQDEH0)DQG2LVKL6 (IIHFWVRIHQYLURQPHQWDOIDFWRUV RQ WR[LFLW\RIDF\DQREDFWHULXP (Microcystis aeruginosa) XQGHU FXOWXUHFRQGLWLRQV $SSO(YLURQ0LFURELRO :HLVH* 'UHZV *-DQQ% DQG-DQQ. ,GHQWLILFDWLRQDQG DQDO\VLV RI OLSRSRO\VDFFKDULGHV LQ FHOO ZDOOV RI WKH EOXHJUHHQ DOJDH Anacystis nidulans. $UFK0LNURELRO :+2 5HSRUW RI WKH :RUNLQJ *URXS RQ &KHPLFDO 6XEVWDQFHV LQ 'ULQNLQJ :DWHU *HQHYD $SULO 6HFWLRQ 0LFURF\VWLQ/5 :RUOG +HDOWK 2UJDQL]DWLRQ *HQHYD :+2 *XLGHOLQHV IRU 'ULQNLQJZDWHU 4XDOLW\ 6HFRQG (GLWLRQ $GGHQGXP WR 9ROXPH +HDOWK &ULWHULD DQG 2WKHU 6XSSRUWLQJ,QIRUPDWLRQ :RUOG+HDOWK2UJDQL]DWLRQ*HQHYD :LFNV 5- DQG 7KLHO 3* (QYLURQPHQWDO IDFWRUV DIIHFWLQJ WKH SURGXFWLRQ RI SHSWLGH WR[LQV LQ IORDWLQJ VFXPV RI WKH F\DQREDFWHULXP Microcystis aeruginosa LQ D K\SHUWURSKLF $IULFDQUHVHUYRLU (QYLURQ6FL7HFKQRO :LHODQG 7 DQG )DXOVWLFK + $QDWR[LQV SKDOORWR[LQV SKDOORO\VLQ DQG DQWDPDQLGH WKH ELRORJLFDOO\ DFWLYH FRPSRQHQWV RISRLVRQRXV Amanita PXVKURRPV &5& &ULW 5HY %LRFKHP :RQJ6+DQG+LQGLQ( 'HWHFWLQJDQDOJDOWR[LQE\ KLJK SUHVVXUHOLTXLG FKURPDWRJUDSK\ - $P:DWHU:RUNV $VVRF
2UJDQLVP ,QGH[ A Acanthophora spicifera, Acetohalobium arabaticum, Aedes aegyptii, Agapetus celatus, Agmenellum, Agmenellum quadruplicatum, Agrobacterium tumefaciens, Agrostis palustris, Alcaligenes eutrophus, Alcaligenes eutrophus &+ Alexandriuni tamarense, Amanita phalloides, DPRHED Plate 2d Anabaena, Plate 2f, 11a, 11b Anabaena aequalis, Anabaena azollae, Anabaena azollae $D/ Anabaena azollae $D1 Anabaena azollae $D6 Anabaena catenula, Anabaena circinalis, Plate l l f Anabaena cylindrica, Anabaena doliolum, Anabaena fertilissima, Anabaenaflos-aquae, Anabaenaflos-aquae 15& Anabaena flos-aquae 15& &DQDGD Anabaena flos-aquae 6J Anabaena hassallii, Anabaena OD[D Anabaena lemmermanni, Anabaena minutissima, Anabaena oscillarioides, Anabaena siamensis $V
Anabaena solitaria, Anabaena spiroides, Anabaena spiroides YDU contracta, Anabaena variabilis, Anabaena variabilis $7&& Anabaena variabilis $7&& Anabaena $% Anabaena $7&& Anabaena $7&& Anabaena $7&& Anabaena $7&& Anabaena %4 Anabaena & Anabaena &3% Anabaena 3&& Anabaena 3&& Anabaena 3&& Anabaena 3&& Anabaena 3&& Anabaenopsis, Anabaenopsis arnoldi, Anabaenopsis milleri, Anabaenopsis tanganyikae, Anacystis, Anacystis dimidiata, Anacystis halophytica, Anacystis marina, Anacystis montana, Anacystis nidulans, Anacystis nidulans $&,$070 Anacystis thermalis, Anthoceros, Anthoceros laevis, Anthoceros punctatus, $QWKRFHURWDH Anulus australis, Aphanizomenon, Aphanizomenon elenkinii, Aphanizomenonflos-aquae, Aphanizomenonflos-aquae IR gracile,
2UJDQLVP ,QGH[
Aphanizomenon flos-aquae 1+ 86$ 1+ 86$ Aphanizomenon gracile, Aphanizomenon ovalisporum, Aphanocapsa, Plate la Aphanocapsa concharum, Aphanocapsa delicatissima, Plate 12a Aphanocapsa elachista, Aphanocapsa endolithica, Aphanocapsa grevillei, Aphanocapsa halophytica, Plate 17c Aphanocapsa holsatica, Plate 12b Aphanocapsa incerta, Aphanocapsa nubilum, Aphanocapsa planctonica, Aphanocapsa Aphanacapsaopsis, Aphanomycopsis cryptica, Aphanothece, Aphanothece bachmannii, Plate 13f Aphanothece clathrata, Plate 13d Aphanothece halophytica, Plate 17c Aphanothece litoralis, Aphanothece microscopica, Aphanothece minutissima, Aphanothece pallida, Aphanothece smithii, Aphanothece stagnina, Arabidopsis, Arabidopsis thaliana, Archaeocillatoriopsis disciformis, Artemia, Artemia salina, Arthrobacter, Arthrospira, 3ODWH VHHDOVRSpirulina Arthrospirafusiformis, Arthrospira geitleri, Arthrospira gomontiana, Arthrospira indica, Arthrospira jenneri, Arthrospira khannae, Arthrospira. massarti, Arthrospira. maxima, Arthrospira okensis,
Arthrospira platensis, Arthrospira platensis 3&& Arthrospira platensis YDU minor, Arthrospira spirulinoides, Arthrospira tenuis, Arthrospira 0 Asterionella formosa, Asterionella glacialis, Aulacoseira, Aulacoseira (Melosira) granulata, Aulosira, Azolla, Azolla caroliniana, Azolla filiculoides, Azolla pinnata, Plate 15e, 28c Azospirillum brasilense, Azotobacter VS Azotobacter vinelandii,
B Bacillus, Bacillus subtilis, Bacillus thuringiensis VVS israeliensis, Balanion planktonicum, Blasia, Plates 29b, 29c, 29d Blasia pusilla, Plate 28d Boeckella, Bonellia fuliginosa, Bordetella pertussis, Bosmina, Plate 14e %RWU\RFRFFDFHDH Botryococcus, Botryococcus braunii, Brachionus, Brachytrichia, Bradyrhizobium japonicum,
C Calamoecia, Calothrix, Plates 5b, 8c, 30a Calothrix desertica, Calothrix gypsophila, Calothrix marchica,
2UJDQLVP,QGH[ Calothrix parietina, Calothrix ' Calothrix 3&& Culyptothrix alternata, Calyptothrix geminata, Candida nowegensis, Caudiculophycus acuminatus, Cerutium, Cerutodon, Ceriodaphnia, Chamaesiphon, 5, Chamaesiphon fuscus, Plate 1g Chlamydomonas, Chlamydomonas nivalis, Chlamydomonas reinhardtii, Chlamydomonas ulvaensis, Chlorella, Chlorella pyrenoidosa, &KORURELDFHDH Chlorobium, Chlorobium vibrioforme, Chlorococcum, &KORURIOH[DFHDH Chlorojlexus, Plate 5a Chloroflexus aurantiacus, Chlorogloeu (Chlorogloeopsis) fritschii, Chlorogloea microcystoides, Chlorogloeopsis, Chlorogloeopsis fritschii, Chlorogloeopsis (HTF), Chlorogloeopsis 2&JV Chondrilla nucula, &KURPDWLDFHDH Chromatium, Plate 17c Chromobacterium violaceum, &KURRFRFFDFHDH &KURRFRFFDOHV Chroococcidiopsis, Plate 21a Chroococcidiopsis kashaii, Chroococcus, Plate 2a, 13b Chroococcus aphanocapsoide, Chroococcus microscopicus, Plate 13b Chroococcus minimus, Chroococcus minutus, Chroococcus turgidus, Chroococcus turgidus YDU spelaeus, Chroococcusturgidus87(; Chroococcopsis, Citharistes,
Citharistes apsteinii, Coccochloris, Coccochloris aeruginosa, Coccochloris elabens, Coccochloris peniocystis, Coccochloris stagnina, Cocos nucifera, Codium, Coelosphaerium, Coelosphaerium kuetzingianum, Coelosphaerium minutissimum, Coelosphaerium punctiferum, Coelosphaerium subarcticum, Coenocystis, Collema, Collema coccophorum, Plate 31c, 31d 31e Collema cristatum, Plate 31b Collema tenax, Conophyton, Crepidula aculeata, Cricotopus fuscatus, Cricotopus nostocicola, Crinalium episammum, Cryptococcus, Cyanidium, Cyanobium, Cyanocatena planctonica, Cyanodictyon, Cyanodictyon filiforme, Plate 12e Cyanodictyon imperfectum, Plate 12c Cyanodictyon planctonicum, Plate 12d Cyanodictyon reticulatum, Cyanodictyon tubiforme, Cyanogranis ferruginea, Cyanonephron, Cyanonephron styloides, Plate 13c Cyanophora, Cyanophora paradoxa, Cyanosarcina, Cyanostylovirus, Cyanothece, Cyanothece aeruginosa, Cyanothece cedrorum, Cyanothece dianae, Cyanothece halobia, Cyanothece (Synechococcus) minewae, Cyanothece 3&& Cycas, Cycas circinalis, 550 Cycas revoluta, Cyclidium, Cyclotella meneghiniana, Cylindrospermopsis,
2UJDQLVP,QGH[
Cylindrospermopsis raciborskii, Cylindrospermum, Cyprinus carpio,
D Dactylococcopsis, Dactylococcopsis acicularis, Dactylococcopsis salina, Daphnia, Daphnia carinata, Daphnia galeata mendotae, Daphnia pulex, Daucus carota, Deinococcus, Deinococcus radiodurans, Dermocarpa, Dermocarpella, Desmococcus, Desulfomonas, Desulfotomaculum, Desulfovibrio, Desulfovibrio vulgaris, Diaphanosoma, Dichothrix, Dichothrix baueriana, Dichothrix calcarea, Dichothrix compacta, Dichothrix gypsophila, Dicranella, 'LGHPQLGDH Didemnum molle, Drepanocladus, Drosophila melanogaster, Dunaliella, Dunaliella parva, Dunaliella tertiolecta, Dysidea herbacea,
E Ectothiorhodospira, Plate 17c Emiliania huxleyi, Encephalartos, Encephalartos ferox, (QWRSK\VDOLGDFHDH Entophysalis, Plate 4A Eoastrion, Eoentophysalis belcherensis, Plate 4B Eomarginata striata, Eomicrocoleus, Eomycetopsis,
Epithemia turgida, (SLWKHPLDFHDH Erythrospira, Escherichia (E.) coli, Eubosmina, Eucalyptus, Plate 28b Eucapsis, Eudiaptomus, Euglena, Euglena gracilis, (XVSLUXOLQD
F Festuca octoflora, Filiconstrictosus diminutus, Filiconstrictosus majusculus, Fischerella, Fischerella epiphytica, Fischerella laminosus, Fischerella muscicola, Fischerella muscicola 87(; Flavobacterium odoratum, Fremyella diplosiphon, Fremyella diplosiphon 0 Funaria,
G Gallionella ferruginea, Geitleria, Geitleria calcarea, Geochelone gigantea, Geodia papyraceae, Geosiphon, Geosiphon pyriforme, Plate 28f Gloeocapsa, Gloeocapsa alpicola, Gloeocapsa alpina, Gloeocapsa biformis, Gloeocapsa calcarea, Gloeocapsa compacta, Gloeocapsa kuetzingiana, Gloeocapsa muralis,
2UJDQLVP ,QGH[ Gloeocapsa punctata, Gloeocapsa ralfsiana, Gloeocapsa rupestris, Gloeocapsa sanguinea, Plates 2c, 32b Gloeocapsa &&DO* Gloeodiniopsis uralicus, Gloeothece, Gloeothece rupestris, Gloeothece $7&& Gloeotrichia, Plate 15c, 15d Gloeotrichia echinulata, Gloeotrichia natans, Plate 15f Gloeotrichia pisum, Glomus, Glycine max, Gomphosphaeria, Gomphosphaeria aponina, Gomphosphaeria lacustris, Gomphosphaeria naegeliana, Gunnera, Gunnera chilensis, Plate 28a Gunnera manicata, Gunnera monoica, Gymnodinium, Gymnostomum recuwirostrum, Plate 16e
H Haeinatococcus, Haemophilus influenzae, Haloarcula, Haloarcula japonica, Halorubrum, Halorubrum distributum, Halteria grandinella, Hapalosiphon, Hapalosiphon fontinalis, Hapalosiphon fontinalis 9 Hapalosiphon intricatus, Hemiaulus, Hemichloris, Hemichloris antarctica, +HSDWLFDH Heterococcus endolithicus, Histioneis, Histoneis elongata, Histoneis highleyi, Homoeothrix, Homoeothrix balearica, Homoeothrix crustacea, Plate 16d
Homoeothrix janthina, Homoeothrix rivularis, Hormathonema, Hormothamnion enteromorphoides, Hyella, Plate 16b Hyella fontana, Hyella immanis, Hypheothrix,
I Ikedosoma gogoshimense, Iyengariella endolithica,
J Johannesbaptista pellucida, K Klebsiella pneumoniae, Klebsormidium flaccidum, Kyrtuthrix, Kyrtuthrix dalmatica,
L Leiothrichoides tenuitunicatus, Lemmermanniella pallida, Plate 13e Lemmermanniella pawa, Lemna, Lempholemma botryosum, Leptogium, Leptogium schraderi, Leptolyngbya, Lichina confinis, Limnothrix, Limnothrix amphigranuluta, Lissoclinum patella, Lissoclinum punctatum, Lissoclinum voeltzkowi, Listeria monocytogenes, Littoraria scabra, Locusta migratoria, Lyngbya, Plate 3A, 15b, 32a Lyngbya aestuarii, Plate 8f, 31a Lyngbya circumcreta, Lyngbya confewoides,
2UJDQLVP ,QGH[
Lyngbya digueti, Lyngbya majuscula, Lyngbya martensiana, Lyngbya murrayi, /\QJE\D valderianum, Lyngbya vandenberghenii, Lyngbya wollei,
M Macrozamia, Macrozamia communis, Mucrozamia lucida, Macrozamia moorei, Mucrozamia riedlei, Mastigocladus, Mastigocladus (Fischerella) laminosus, Mastigocladus +7) Mastigocoleus testarum, Melosiragranulata, Melosira nyassensis YDU victoriae, Mentzeliamultiflora, Mercenaria mercenaria, Merismopedia, Plate 2a Merismopedia tenuissima, Merismopedia warmingiana, Mesocyclops, Metallogenium, Microchaete, Microcoleus, Microcoleus chthonoplastes, Microcoleus chthonoplastes 3&& Microcoleus chthonoplastes 6$* Microcoleus lyngybyaceus, Microcoleus paludosus, Microcoleus paludosus YDU acuminatus, Microcoleus sociatus, Microcoleus vaginatus, Microcystis, Microcystis aeruginosa,
Plate 2d, 11c Microcystis aeruginosa IR aeruginosa, Microcystis aeruginosa 0 Microcystis aeruginosa 15& 66 Microcystis aeruginosa 3&& Microcystis aeruginosa 3&& Microcystis aeruginosa 87(; Microcystis aeruginosa :5 Microcystis aeruginosa Microcystis botrys, Microcystis firma, Microcystisflos-aquae, Microcystis stagnalis, Microcystis viridis, Microcystis wesenbergii, Micromonas pusilla, Mycoplasma genitalium, 0\RYLULGDH Myxobactron salinum, Myxococcus fulvus %*2 Myxococcus xanthus, Myxosarcina,
N Nassula aurea, Nephroma arcticum, Neurospora crassa, Nitzschia closterium, Nodularia, Nodularia haweyana, Nodularia spumigena, Plate 11e Nostoc, &KDSWHU Plate 29d, 29e, 29f Nostoc calcicola, Nostoc carneum, Nostoc commune, Plate 24b, 24c, 24d, 24e
2UJDQLVP,QGH[ Nostoc commune YDU flagelliforme, Plate 1i Nostoc commune &+(1 Nostoc commune '5+ Nostoc commune 87(; Nostoc cordubensis, Nostoc ellipsosporum, Nostoc flagelliforme, Plate 1i: VHHDOVRNostoc commune YDU flagelliforme Nostoc fuscescens YDU mixta, Nostoc linckia, Nostoc microscopicum, Nostoc minutum, Nostoc muscorum, Nostoc muscorum $ Nostoc paludosum, Nostoc parmelioides, Nostoc pruniforme, 3ODWHD Nostoc punctiforme, Nostoc punctiforme $7&& Nostoc rivulare, Nostoc sphaericum, Nostoc sphaeroides, Nostoc spongiaeforme, Nostoc verrucosum, Nostoc zetterstedtii, Nostoc $7&& Nostoc $ Nostoc %DOL Nostoc &F Nostoc ' Nostoc (IO Nostoc *LEVRQ Nostoc *P Nostoc 0$& Nostoc 0 Nostoc 0PO Nostoc 081 Nostoc 1,9$&<$ Nostoc 3&& Nostoc 3&& Nostoc 3&& Anabaena 3&& Nostoc 3&& Nostoc 3&& Nostoc 7 Nostoc 8$0 Nostoc 8&' Nostoc 6% 1RVWRFDFHDH 1RVWRFDOHV
Nostochopsis lobatus, Nostocomorpha, O
Ornithocercus, Ornithocerus magnificus, Ornithocerus quadratus, Ornithocerus steinii, Oscillatoria, 55, Plate 17a, 17b Oscillatoria acutissima, Oscillatoria acutissima % Oscillatoria agardhii, Oscillatoria agardhii YDU isothrix, Oscillatoria amphigranulata, Oscillatoria chlorina, Oscillatoria formosa, Oscillatoria irrigua, Oscillatoria limnetica, Oscillatoria limosa, Oscillatoria limosa 6WUDLQ Oscillatoria lutea, Oscillatoria nigroviridis, Oscillatoria perornuta, Plate 11d Oscillutoria priestleyii, Oscillatoria princeps, Oscillatoria redekei, Oscillatoria rubescens, Oscillatoria salina, Oscillatoria spongelia, Oscillatoria submembranacea, Oscillatoria tenuis, Oscillatoria terebriformis, 55, Plate 5a Oscillatoria %* 2VFLOODWRULDFHDH Plate 23e 2VFLOODWRULDH 2VFLOODWRULDOHV Oscillatoriopsis,
Oscillatoriopsis majuscula, Oscillatorites, RVWUDFRG Plate 5c Ostreobium, Ostreobium quekettii,
P Palaeolyngbya, Palaeolyngbya barghoorniana, Palaeolyngbya helva, Plate 3B Palaeolyngbya minor, Palaeonostoc, Palaeopleurocapsa fusiforma, Palaeopleurocapsa kelleri, Palaeopleurocapsa reniforma, Palaeovaucheria elevata, Pannus spumosus, Paramecium, Paramecium caudatum, Parasponia andersonii, Partitiofilum gongyloides, Pediastrum, Peltigera, Peltigera aphthosa, Plate 29g Peltigera canina, Peltigera membranacea, Phaeoceros, Plate 29a Phaeophila dendroides, Phalaropus, Phoenicopterus minor, Phormidium, Plate 1h, 6c Phormidium ambiguum, Phormidium angustissimum, Phormidium autumnale, Phormidium calcicola, Phormidium corium, Phormidium crossbyanum, Phormidium favosum, Phorrnidium foveolarum, Phormidium fragile, Phormidium frigidum, Phormidium hendersonii, Plate 1d, 1e Phormidium hypersalinum, Phormidium hypolimneticum, Phormidium incrustatum,
2UJDQLVP ,QGH[ Phormidium laminosum, Phormidium martensiana, Phorrnidium murrayi, Phormidium persicinum, Phormidium priestleyi, Phormidium tenue, Phormidium uncinatum, Phormidium valderianum, Pinguicula vulgaris, Plate 16e Placopsis contortuplicata, Placynthium, Placynthium nigrum, Planktothrix, Planktothrix perornata, Plate 11d Plectonema, Plate 23b, 23c, 23f Plectonema boryanum, Plectonema boryanum 87(; Plectonema nostocorum, Plectonerna polymorphum, Plectonema terebrans, Plectonema tomasianum, Plectonema Pleomorpha, Pleurocapsa, Plate 1a Pleurocapsa minor, 3OHXURFDSVDFHDH 3OHXURFDSVDOHV 3RGRYLULGDH Porites, Porites lobata, Porphyrosiphon, Porphyrosiphon fuscus, Porphyrosiphon notarisii, Portulaca (oleracea), Plate 24b Potarnocypris, Plate 5b 3UDVLQRSK\FHDH Prasiococcus, Primaevifilum amoenum, Primaevifilum conicoterminatum, Primaevifilum laticellulosum, Procambarus clarkii, Prochlorococcus, Prochlorococcus marina, Prochloron, Prochloron didemni, Prochlorothrix, Prochlorothrix hollandica, Prorocentrum minimum,
2UJDQLVP,QGH[ Prototheca zopfi, Pseudanabaena, Pseudanabaena catenata, Pseudanabaena galeata, Pseudanabaena 3&& Pseudomonas, Pseudomonas aeruginosa, Pseudomonas pelfectomarina, Pseudotrebouxia, 3WHULGRSK\WD Pyrenocollema, 3\UHQRSVLGDFHDH Pyricularia oryzae, Pythium torulosum, Pyura cancellata, Pyura carnea,
R Rhizosolenia, Rhizosolenia clevei, Rhodococcus, Rhodomonas lens, 5KRGRVSLULOODFHDH 5KRGRVSLULOOXP Rhopalodia gibba, Richelia, Richelia intracellularis, Rivularia, Plate 16c, 26a, 26c, 26d Rivularia atra, Plate 26e, 26f Rivularia biasolettiana, Rivularia haematites, Rivularia minutula, 5LYXODULDFHDH Plate 26 Romeria elegans, Romeria leopoliense,
S Saccharomyces cerevisiae, Salmonella, Salmonella typhimurium, Sargassum, Scenedesmus, Scenedesmus brasiliensis, Scenedesmus protuberans, Schistocerca gregaria, Schizothrix, Plate 2e, 9e, 23d Schizothrix adunca,
Schizothrix affinis, Schizothrix antarctica, Schizothrix arenaria, Schizothrix atacamensis, Schizothrix calcicola, Schizothrix coriacea, Schizothrix fasciculata, Schizothrix fasciculata IR semiglobosa, Schizothrix lardacea, Schizothrix lateritia, Schizothrix pelforans, Schizothrix pulvinata, Schizothrix rubella, Sclerotinia sclerotiorum, Scytonema, Plate 25, 31b Scytonema %6F\M Plate 32c Scytonema crustaceum, Scytonema endolithicum, Scytonema hofmanni, Scytonema hofmanni 87(; Scytonema julianum, Scyyonema mirabile, Scytonema mirabile %< Scytonema myochrous, Plates 2h, 25e Scytonema pseudohofmanni, Scytonema pseudohofmanni %& Selenastrum, Serratia marcensis, Siphonochalina tabernacula, Siphonophycus costatus, 6LSKRYLULGDH Skeletonema costatum, Snowella, Snowella atomus, Solentia, Sphaerocystis, Sphagnum, Sphagnum cuspidatum, Spirulina, Plate 27: VHHDOVRArthrospira Spirulina fusiformis, Spirulina geitleri, Spirulina jenneri IR platensis, Spirulina labyrinthiformis, Spirulina laxissima, Spirulina major, Spirulina maxima, Spirulina minosa,
2UJDQLVP,QGH[
Spirulina platensis, Spirulina subsalsa, Sporobolus virginicus, Staphylococcus aureus, Staphylococcus xylosus, Stichococcus, Stichococcus bacillaris, Stigmatella aurantiaca, Stigonema, Stigonema mamillosum, Plate 2i Stigonema minutum, Stigonema ocellatum, 6WLJRQHPDWDFHDH 6WLJRQHPDWDOHV Streptococcus pneumoniae, Streptomyces lividans, Strobilidium hexachinetum, 6W\ORYLULGDH Symploca, Synechococcus, Plate 5e, 5f, 6c, 7, 10c, 13, 14b, 17c, 23f Synechococcus aeruginosa, Synechococcus bacillaris, Synechococcus cedrorum, Synechococcus cedrorum 3&& Synechococcus cedrorum 87(; Synechococcus elongatus, Synechococcus elongatus 87(; 3&& Synechococcus gaarderi, Synechococcus leopoliensis, Synechococcus leopoliensis 6$* Synechococcus lividus, Synechococus lividus +7) Synechococcus minervae, Synechococcus nidulans, Synechococcus plancticus, Synechococcus vulcanus, Synechococcus $7&& Synechococcus & Synechococcus &
Synechococcus '& Synechococcus '81 Synechococcus +7) Synechococcus 0LDPL%&,,6 Synechococcus 0LDPL%* Synechococcus 1 Synechococcus 1.%* Synechococcus 3&& Synechococcus 3&& Synechococcus 3&& Synechococcus 3&& Synechococcus 3&& Synechococcus 3&& Synechococcus 3&& Synechococcus 5) Synechococcus 55,03 1, Synechococcus 61&O 87(; Synechococcus 6<1 Synechococcus 7; Synechococcus 87(; Synechococcus :+ Synechococcus :+ Synechococcus :+ Synechococcus :+ Synechococcus :+ Synechococcus :+ Synechococcus <FV Synechococcus %* Synechocystis,4, Synechocystis aquatilis, Synechocystis '81 Synechocystis 3&& Synechocystis 3&&
T
2UJDQLVP,QGH[ Tetrahymena, Tetrahymena pyriformis, Tetraphycus bistratosus, Tetrarcus ilsteri, Plate 12f Thalassiosira weissflogii, Theonella swinhoei, Thermothrix, Thermotogu maritima, Thiobacillus ferrooxidans, Thiobacillus neopolitanus, Thiocapsa, Plate 17b Thiocapsa roseopersicina, Tilapia, Tolypothrix, Tolypothrix brevis, Tolypothrix byssoidea, Plate 1c Tolypothrix byssoidea + Tolypothrix tenuis, Plate 2g Tortunema, Trebouxia, Tremella, Trentepohlia, Tribonema, Trichodesmium, Trichodesmium erythraeum, Trichodesmium thiebautii, Plate 10a, 10b Trichodesmium ,06 Trichormus, Trichormus azollae,
U Umezakia natans, Urotricha,
V Veteronostocale, Veteronostocale amoenum, Vibrio fischeri, Vibrio harveyii, Vibrio parahaemolyticus, Vitreoscilla, 9ROYRFDOHV Vorticella, Plate 14c Vorticella aquadulcis,
W Wollea, Woronichinia,
X Xenococcus, Xenococcus acervatus, Xenococcus kerneri,
Z Zamia, Zamia skinneri, Zea mays, Zygnema, Zymomonas mobilis,
*HQH DQG *HQH 3URGXFW ,QGH[ A $S$ arsA arsR arsS
B %LI$
C &DG& ccmA ccmK ccmKLMNO ccmL ccmM ccmN ccmO cemA che &KH$ &KH% &KH: &KH< &KH= chrA &OS% &OS& cmpA &PS$ cmpABCD &RD5 cobH cotA &27 cpcBACDEF cpcB1A1 cpcB2A2 cpcB3A3 cpcB-lacZ FKLPHULFJHQH cpcE cpcF cpeBA cpeCDE ctaA
&WD$ &\D$ &\D% &\D% cyaC &\D& cyal cynS cysA &\V$ cysP cysR &\V5 cysT &\V7 cysW &\V: czc &]F%$ czcD
D 'HU+ dnaG devR dspA
F )G%N )L[. frz )U]$ )U]( fur
G glbN *OE1 glnA glnB *OQ% glnD glnE glnN
H hatR hrmA hrmU hrmUA hupAB
I icfA 412, 421 ,SK3 ,W]$
K .DL$ kaiABC .DL&
*HQHDQG*HQH 3URGXFW,QGH[ 1LIO 1LI 1L5 nirA nirB QLU, QLU,, 1LU% nod nodEF nodMN nrtA 1UW$ nrtB nrtC nrtD 1WF$ ntcB 1WF%
L
O
lacZ lux luxAB JHQHV /\V5
2SF$ opcA orfR orfS
M
P
0HU5 0WQ%
pacL 3DF/ pacS 3DF6 S$1/SODVPLG patA petE pheT phn phoA phoB SKR% phoBR phoH SKR5 phoR SKR5 phoU phoV SKR9 phy pil 3LO*+ 3LO,
N narB natA natB nblA 1EO$ nblB 1EO5 nblR ndhB ndhC ndhE nifD nifH 1LI+ nif HDK nifLA nzjU nifH
*HQHDQG*HQH 3URGXFW,QGH[ 3LO- plpA psaI psaL psbA psbAI psbAII psbAIII psbD pstA SVW$ SVW% SVW& pstS SVW6 pstSCAB 3WU$ purK 3]S, 3,,
R
VOU VOU VOU VOU VOU VOU VOU VOU VOU smt smtA 6PW$ smtB 6PW% sphR 6SK5 sphS 6SK6 6SR2) 6SR7 spoT 6US$ 6US* 6US+ srrA srrB sucA 6ZP$
rbcLS rbcLX rcaC rcaE UFD( rcaF rec recA 5HO$ relA rhdA 5KG$ rpoB rpoC rpoD rpoS rpsU
:VS
S
Z
sbpA 6ES$ VHPOO sigA sigB sigC VOO VOO VOO VOO
ziaA =LD$ ziaR =LD5 =QW5 =QX$ =5&O =XU ]ZI
U ugpQ uidA ureABC ureDEF
W
6XEMHFW ,QGH[ A $EODWLRQ3RLQW$QWDUFWLFD DEUDVLRQ DEVFLVLFDFLG $EX *DEDUD /DNH(J\SW $FH /DNH $QWDUFWLF DFHWDOGHK\GH DFHWDQLOLGH DFHWDWH DFHW\OFKROLQH HVWHUDVH DFHW\OSKRVSKDWH DFHW\OHQHUHGXFWLRQDFWLYLW\ DVVD\ DFLGLF VHH also S+ HIIOXHQW ORZS+ODNHVDQGSRROV DFWLQ DFXWLSK\FLQ DGDSWDWLRQ VWHSZLVH $GDV]WHYHO2URVKD] +XQJDU\ $''$ 6 6 6 6 -DPLQRPHWKR[\ WULPHWK\O -SKHQ\OGHFDGLHQRLF DFLG DGHQ\OWUDQVIHUDVH DGHQ\ODWHF\FODVH DGKHVLRQ DGRQLWRO $'3 $GULDWLFFRDVW DHURWD[LV DJJUHJDWLRQ RI KRUPRJRQLDRUQRUPDOWULFKRPHV VRLO DJULFXOWXUH DLU IORUD DLUZDWHULQWHUIDFH DNLQHWH Plate 2d DODQLQH DODQLQH $ODVND DOEHGR $OGDEUDDWROO Plate 1a,1b, 1c, 1d, 1e, 24b, 24c DOGRODVHV $OH[DQGHU,VODQG$QWDUFWLFD
DOIDOID DOJDH DOJDOL]DWLRQ DOJLQDWH EHDGV $OJHULD $OKDPEUD *UDQDGD6SDLQ $OLFDQWH 6SDLQ DONDOLQH ODNHV &KDSWHU see also :DGL 1DWUXQ DONDOLQH SKRVSKDWDVH see also SKRVSKDWDVHSKRVSKRPRQRHVWHUDVH DONDQHV &KDSWHU HVS QDONDQRLF DFLGV $O.KLUDQ.XZDLW DONDOLSKLOH DONDORLG $OO ,QGLD &RRUGLQDWHG 5LFH,PSURYHPHQW3URMHFW DOOHUJLF UHVSRQVHV DOOHORSDWKLF UHVSRQVH DOORSDWULF VSHFLDWLRQ DOORSK\FRF\DQLQ DOSLQH ODNH $O] 5LYHU %DYDULD DPLQRDFLGV WUDQVSRUW DPLQRLVREXW\ULF DFLG DPLQROHYXOLQLF DFLG 6 6 6 6 DPLQRPHWKR[\ WULPHWK\O SKHQ\OGHFDGLHQRLF DFLG $GGD SDPLQRSKHQRO DPLQRWUDQVIHUDVHV DPPRQLD DPPRQLXP DVVLPLODWLRQ H[FUHWLQJPXWDQW UHSUHVVLRQ WUDQVSRUW YRODWLOLVDWLRQ DPRHEDPlate 2d DPRHERLG &$03 DPSKLELDQ DPSLFLOOLQ DQDWR[LQD DQDHURELFUHVSLUDWLRQ DQDHURELF DQDHURELRVLV
6XEMHFW ,QGH[
$QGURV ,V %DKDPDV Plate 25a, 25b, 25c DQJLRVSHUPV DQK\GURELRVLV DQLOLQH DQLPDO IHHG SDQLVLGHQH Anopore ILOWHUV DQRUH[LD DQR[LD DQR[LF DQR[\JHQLF SKRWRV\QWKHVLV DQWDJRQLVP $QWDUFWLF &KDSWHU &KDSWHU VHH also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
$UUKHQLXV SORW DUVHQLF DVFLGLDQ DVHULDWH PRUSKRORJ\ $VLU 0RXQWDLQV 6DXGL $UDELD Plate 1h DVSDUDJLQH DVSDUWDWH DVSDUWLF DFLG 550 SKRVSKRU\ODWLRQ DVSKDOWHQH DVWD[DQWKLQ DVWHULQD $VWRYH DWROO $WDFDPD 'HVHUW DWKDODVVRKDOLQH $WODQWLF FRDVW 2FHDQ DWPRVSKHULF 1 GHSRVLWLRQ SUHFLSLWDWLRQ DWROO VHH $OGDEUD $VWRYH )DUTXKDU 6W 3LHUUH $73 UHTXLUHPHQW IRU YLUDO UHSOLFDWLRQ V\QWKDVH $73DVH DWWDFKPHQW $XURPDLQH DXULFOH $XVWUDOLD Plate 1f, 11a, 11b,11c, 11d, 11e, 11f: see also $\HUV 5RFN (DUO\ $UFKHDQ $SH[ &KHUW +DZNHVEXU\ 5LYHU /DNH 'DUWPRXWK /DNH +D\ZDUG /DNH +XPH 0XUUXPELGJHH 5LYHU 3HHO +DUYH\ (VWXDU\ 6KDUN %D\ 6SHQFHU *XOI $XVWULD VHH also *VFKQLW]EDFK ,VDU DXWRIOXRUHVFHQFH $YGDW 1HJHY D[HQLF FXOWXUH $\HUV 5RFN $XVWUDOLD D]DVHULQH D]DWU\SWRSKDQ
B EDFWHULRFKORURSK\OO a, EDFWHULRFKORURSK\OO F EDFWHULRFLGDO EDFWHULRKRSDQRLG EDFWHULRO\WLF EDFWHULRSKDJH EDFWHULRSODQNWRQ EDFWHULRVWDWLF
6XEMHFW ,QGH[ EDHRF\WH EDJDVVH %DKDPDV %DMD &DOLIRUQLD %DOWLF 6HD %DQGHG ,URQ )RUPDWLRQV %,) %DQJODGHVK Plate 25d, 26b %DQNV - %DUZLFN 9DOOH\ %DUGDZLO ODJRRQ (J\SW EDULXP EDWKLQJ FRVWXPH %DWWOHVKLS 3URPRQWRU\ $QWDUFWLF Plate 20b %DXW]HQ 5HVHUYRLU %DYDULD see )LVFKNDOWHUVHH %&,3 EURPRFKORURLQGRO\O SKRVSKDWH %HDFRQ +HLJKWV $QWDUWLF Plate 20a %HDFRQ VDQGVWRQH $QWDUFWLF %HDJOH +06 VKLS %HDXIRUW 1RUWK &DUROLQD EHHW %HOL]H EHQ]HQH EHWDLQH VHH JO\FLQH EHWDLQH ELFDUERQDWH &KDSHU WUDQVSRUW XSWDNH %,) %DQGHG ,URQ )RUPDWLRQV ELOODERQJ ELQGLQJ ELRDVVD\V ELRILOP ELROXPLQHVFHQW ELRPDUNHU ELRSWHULQ JOXFRVLGH %LUG 6KRDO 1RUWK &DUROLQD ELUGV VHH DOVR IODPLQJR ELYDOYH EODFN ERG\ DEVRUSWLRQ EODFN OD\HU %ODFN 6HD EODGGHU IXQJDO see also Geosiphon EORRP IUHVKZDWHU &KDSWHU 3ODWH 5 PDULQH Plate 10a, 10b, 10d EOXH OLJKW %RJXH 6RXQG 1RUWK &DUROLQD ERULF DFLG ERULQJ EUDFNLVK
EUDQFKLQJ IDOVH WUXH %UDXQ%ODQTXHW SK\WRVRFLRORJ\ %UHWDJQH FRDVW )UDQFH %ULWLVK &ROXPELD &KLONR /DNH WUHH QXUVHU\ EURPLQDWHG SURGXFWV EURPRFKORURLQGRO\O SKRVSKDWH %&,3 EU\RSK\WH see also Dicrunella, OLYHUZRUW PRVV Sphagnum EXLOGLQJV EXR\DQF\ &KDSWHU HVS EXUQLQJ %XUUHQ ,UHODQG EXW\UDWH %\ORW ,VODQG $UFWLF
C FDGPLXP RFHDQLF FDIIHLQ &DKHU 5LYHU ,UHODQG FDOFLILFDWLRQ IUHVKZDWHU &KDSWHU PDULQH FDOFLWH &KDSWHU HVS VXSHUVDWXUDWLRQ FDOFLXP LRQV FDOFLXP FDUERQDWH &KDSWHU FDOFLXP SKRVSKDWH &DOLIRUQLD FDOPRGXOLQ &DOYLQ F\FOH FDO\FXOLQ FDO\SWUD &DPDUJXH )UDQFH &DPEULDQ &DQDGD see also %ULWLVK &ROXPELD +XURQLDQ 6XSHUJURXS /DNH +XURQ 4XpEHF &DQDGD *ODFLHU $QWDUFWLF FDQWKD[DQWKLQ &DSH &RG 0DVVDFKXVHWWV FDUEDPR\O SKRVSKDWH FDUEDU\O FDUERK\GUDWH DFFXPXODWLRQ UHVHUYH
652 FDUERQ GLR[LGH GHSOHWLRQ OLPLWDWLRQ IL[DWLRQ DQG IUHVKZDWHU SODQNWRQ PRWLOLW\ UHVSRQVH RI 2VFLOODWRULD VS WUDQVSRUW XSWDNH FDUERQLF DQK\GUDVH &DUERQLIHURXV OLPHVWRQH FDUER[\IOXRUHVFHLQ FDUER[\VRPH FDUFLQRJHQ FDUFLQRJHQLF &DUJLOOH LPPHUVLRQ RLO &DULEEHDQ FDURWHQH FDURWHQRLG
FDUS FDUURW &DVD $XUHXP 5RPH FDWDODVH FDWHFKRO FDWWOH FDYHV VHH DOVR VSHOHRWKHP )ORULGD LQIOXHQFH RI OLJKW FHOO GLYLVLRQ ORFDOL]HG FHOORELRVH FHOOXORVH FHSKDORGLD &)8 FRORQ\IRUPLQJ XQLW &KDG FKDON FKDSHURQLQ FKDVPRHQGR OLWK FKHODWLRQ
FKHPRDWWUDFWDQW FKHPRDWWUDFWLRQ FKHPRDXWRWURSKV FKHPRWD[LV FKHUW &KLOH : seeDOVR$WDFDPD'HVHUW &KLONR /DNH %ULWLVK &ROXPELD &KLQD &KLQHVH 1HZ
see DOVR
FKORURSK\OO b, SURFKORURSK\WH
6XEMHFW ,QGH[ FKORURSK\OO c, FKORURSODVW FKORURVLV FKORUSURPD]LQH FKRDQRIODJHOODWH FKURPDWH FKURPDWLF DGDSWDWLRQ -
FKURPRSKRUH FKURPRVRPH FKU\VRSK\WHV FK\PRWU\SVLQ FLJXDWHUD FLOLDWH FLOLRSKRUDQ FLQJXOXP FLUFDGLDQ F\FOH UK\WKP -
FLV GLK\GUR[\GLK\GURQDSKWKDOHQH
FLWUDWH FLWUXOOLQH FODGRFHUDQ &ODUHQV 6DQGVWRQH 6RXWK $IULFD FOHDUDQFH UDWH &OHDUZDWHU 6SULQJV
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
F\OLQGULFDO ERG\ F\OLQGURVSHUPRSVLQ F\VWHLQH F\VWHLQHWKLRO JURXSV F\VWLQ F\WRFKURPH EI F R[LGDVH F\WRSODVPLF PHPEUDQH F\WRVRO F\WRWR[LQ &]HFK 5HSXEOLF
D 'GLFKORURSKHQR[\ DFHWLF DFLG 'DQDUHDVHZDJH WUHDWPHQWSODQW,VUDHO 'DQ :DVWHZDWHU 5HFODPDWLRQ 3URMHFW,VUDHO GDUN UHSDLU 'DUZLQ 'DYLV 6WDWLRQ$QWDUFWLF '&08 GLFKORURSKHQ\O GLPHWK\OXUHD 'HDG 6HD GHEURPRDSO\VLDWR[LQ GHFDQHV GHFD\ GHFRPSRVLWLRQ GHHSZDWHUULFH Plate 15c, 15d, 25d GHK\GURDODQLQH 'HQPDUN GH5HDPXU GHQLWULILFDWLRQ GHSKRVSKRU\ODWLRQ GHSRVLWLRQ DWPRVSKHULF GHUPDOWR[LQ GHVHUW&KDSWHU $WDFDPD FHQWUDO$XVWUDOLDQ *REL KRWYHUVXVFROG 1DPLE 1HYDGD 1HJHY 6DKDUD 6RQRUDQ 'HVHUW 0LFURIORUD 3URJUDP GHVHUWYDUQLVK GHVLFFDWLRQ &KDSWHU &KDSWHU
GHVPLG GHZ GLDUUKRHD GLDGHQRVLQH WHWUDSKRVSKDWH GLDJHQHVLV GLDU\OODFWRQH GLDWRPV GLD]RR[R/QRUOHXFLQH '21 GLD]RWURSK VHH QLWURJHQIL[DWLRQ GLFKORURSKHQ\O GLPHWK\OXUHD'&08 GLFKORURSKHQR[\ DFHWLFDFLG' GLGHK\GURDFXWLSK\FLQ GLGHPQLG 'LGLP 7XUNH\ GLHOFKDQJH VHH also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
6XEMHFW ,QGH[
E (DUO\ $UFKHDQ $SH[ &KHUW %DVDOW $XVWUDOLD (DUWKULVH)DUPV&DOLIRUQLD Plate 27 (DVW %HDFRQ $QWDUFWLF HFKLQHQRQH HFKLQRGHUP HFKLXURLG ZRUP HFRV\VWHP (FXDGRU ('7$ HWK\OHQHGLDPLQHWHWUDDFHWDWH (GZDUG 9,,3HQLQVXOD$QWDUFWLF HIIOXHQW VHZDJH WDSLRFD 5 (IRULH 8NUDLQH (JZ\V 1\Q\GG :DOHV (J\SW see also $EX *DEDUD /DNH %DUGDZLO ODJRRQ6RODU/DNH:DGL1DWURQ (LODW Plate 25a (LQHVEHUJHU =HQWUDOH *HUPDQ\ Plate 13f HODVWDVH HOHFWURQ GRQRU VXOILGH (O+DPLUD%D\6LQDL (O 1LxR6RXWKHUQ2VFLOODWLRQ (OOHVPHUH ,VODQG HPEU\RJHQHVLV (QGHDYRXU +06 VKLS HQGRJDODFWRVLGDVH HQGROLWK HQGRVXOIDQ HQGRV\PELRQW HQGRWR[LQ (QJODQG see (QYLURQPHQW $JHQF\ (VWKZDLWH:DWHU)UHVKZDWHU%LRORJLFDO $VVRFLDWLRQ 0DOKDP 5XWODQG :DWHU 6XIIRON :DWHU%RDUG6ZDOH 5LYHU 6XQELJJLQ8SSHU 7HHVGDOH :DWHUIDOO %HFN (QJOLVK ODNHV QRUWKHUQ VHHDOVR (VWKZDLWH HQXPHUDWLRQ RISLFRSODQNWRQ HQWHULF EDFWHULD (QYLURQPHQW$JHQF\(QJODQGDQG:DOHV HSLGHPLRORJ\ HSLIOXRUHVFHQFH HSLOLPQLRQ HSLOLWKLF&KDSWHU HSLSK\WH (SRFK ,,,
6XEMHFW,QGH[ HURVLRQ VWUHDP HU\WKURPHWK\OGLVRDVSDUWDWH HVWHUV (VWKZDLWH:DWHU(QJODQG HWKDQRO (WKLRSLD HWK\OHQH DVSURGXFWRIDVVD\ see DFHW\OHQHUHGXFWLRQ DVVD\ VHQVRU HXHQGROLWK HXSKRWLFGHSWK]RQH HXWURSKLFDWLRQ&KDSWHU HYDSRULWH (YHUJODGHV H[IROLDWLRQ H[RELRORJ\ H[RSRO\PHU see H[WUDFHOOXODUSRO\VDFFKDULGH ([SRUWDGRUDGH6DO0H[LFR H[WUDFHOOXODUsee also SKRVSKDWDVH SRO\VDFFKDULGH '1$ SKRVSKDWDVHV SKRVSKRPRQRHVWHUDVH SRO\VDFFKDULGH &KDSWHU ([XPD&D\V%DKDPDV Plate 9
F )DDV)UHQFK3RO\QHVLD RIDFWRUV )DUTXKDU,QGLDQ2FHDQ IDUUHGOLJKW IDWW\DFLGV SRO\XQVDWXUDWHG IHOOILHOGVRLOV )HQWLQGHULYDWLYHV IHUPHQWDWLRQSDWKZD\ IHUQ VHHAzolla IHUUHGR[LQ IHUULFLURQ IHUWLOLVHU ILEULOVZDOO )LML ILODPHQW EHFRPLQJXQLFHOOXODU WDSHUHG WLSV
ILOWHUV Anopore, PHPEUDQH )LQODQG ILVFKHUHOOLQ )LVFKNDOWHUVHH%DYDULD ILVK SRQG DQGWR[LQV ILVKSRQGV IODJHOODWHV IODJHOOXP IODPLQJR IODYRGR[LQ IODYRQHJO\FRVLGH IODYRQRLG IORDWLQJ&\DQREDFWHULDOPDVV IORUDV )ORULGD(YHUJODGHV IORZFRXQWU\6FRWODQG IORZF\WRPHWU\ IOXRUHVFHLQ LVRWKLRF\DQDWH IOXRUHVFHQFH Plate 14 DXWRIOXRUHVFHQFH ODEHOOLQJ TXHQFKLQJ IOXRURFKURPH IRRG IRRGZHE IRUPDOGHK\GH IRUPDQLOLGH IRUPDWH IRVVLOV&KDSWHU )UDPQHV0RXQWDLQV$QWDUFWLF )UDQFH see %UHWDJQHFRDVW&DPDUJXH6DOLQVGH *LUDXG )UDQ]-RVHI/DQG IUHH]HIUDFWXUHHOHFWURQPLFURVFRS\ IUHH]LQJ )UHQFK3RO\QHVLD )UHVKZDWHU%LRORJLFDO$VVRFDWLRQ(QJODQG IUXFWRVH IUXFWRVHELVSKRVSKDWDVH IUXFWRVH ELVSKRVSKDWHDOGRODVH IXFRVH IXPDUDWH IXQJLFLGH IXQJXV see also P\FRELRQWSK\FRP\FHWH \HDVW
G JDODFWRVH ' JDODFWXURQLFDFLG JDVYDFXROH&KDSWHUHVS FROODSVHE\VKRFNZDYHV VWUXFWXUH JDVYHVLFOH &KDSWHU JDVSLQJ JDVWURSRG *DWXQ/DNH3DQDPD *DYLVKVDENKD *'+ VHHJOXWDPDWHGHK\GURJHQDVH JHHVH JHOEVWRII JHPPD *HQ%DQN JHQHH[FKDQJH JHQHWLFWROHUDQFH JHQRPH JHRVPLQ (trans- GLPHWK\OWUDQVGHFDORO JHRWDFWLFUHVSRQVH *HUPDQ\ VHHalso $O](LQHVEHUJHU =HQWUDOH JLDQWWRUWRLVH JLOOGDPDJH *MHOYLNVIMHOOD$QWDUFWLF JODQG LQ Gunnera, PXFXVVHFUHWLQJ JOLGLQJPRWLOLW\ UDWH JOREDOFKDQJH JORELQ JORHRFDSVLQ JOXFRVDPLQHSKRVSKDWH JOXFRVH JOXFRVHSKRVSKDWH JOXFRVHSKRVSKDWHGHK\GURJHQDVH 'JOXFLV\O JOXFRV\OJO\FHURO ' JOXFXURQLFDFLG 3JOXFXURQLGDVH JOXWDPDWH GHK\GURJHQDVH*'+ V\QWKDVH JOXWDPLQH V\QWKHWDVH*6
6XEMHFW,QGH[ JOXWDUDOGHK\GH JOXWDWKLRQH JO\FDQVKHDWK JO\FHURO JO\FLQH JO\FLQHEHWDLQH JO\FRJHQ JO\FRODWH H[FUHWLRQ JO\FRSURWHLQ *REL'HVHUW&KLQD *2*$7 JOXWDPLQHR[RJOXWDUDWH DPLQR WUDQVIHUDVH *ROGHQ*DWH+LJKODQGV1DWLRQDO3DUN 6RXWK $IULFD JROIFRXUVHV *UDPQHJDWLYHEDFWHULD *UDPSRVLWLYHEDFWHULD JUD]HUJUD]LQJ see also FRSHSRGF\FORSRLGRVWUDFRG *UHDW%DVLQ'HVHUW *UHDW6DOW/DNH8WDK *UHDW6LSSHZLVVHWW0DUVK0DVVDFKXVHWWV *UHHFH JUHHQDOJD see also LQGLYLGXDOJHQHUD JUHHQGHDWKFDS *UHHQ/DNH(WKLRSLD *UHHQ/DNH86$ JUHHQOLJKW JUHHQQRQVXOIXUEDFWHULD see also Chloroflexus *UHHQODQG6HD JURXQGZDWHU JURZWKUDWH *VFKQLW]EDFK5LYHU $XVWULD JXDQR JXDQRVLQH GLSKRVSKDWH GLSKRVSKDWH SS*SS JXDQRVLQHWHWUDSKRVSKDWH *XHUUHUR1HJUR0H[LFR Plate 8 *XOIRI$TDED *XOIRI0H[LFR *XOI6WUHDP J\SVXP Plate 17c
6XEMHFW,QGH[ H KDLU Azolla, 5LYXODULDFHDH KDOLWH +DPHL0D]RUVSULQJV KDSDOLQGROH +DUWEHHVSRRUW'DP6RXWK$IULFD +DUWLJDQ)LWFK6DQNRIIDOJRULWKP KDXVWRULD +DZDLL +DZNHVEXU\5LYHU1HZ6RXWK:DOHV KHDOWKIRRG KHDY\PHWDOV&KDSWHU Plate 23 +H/DFHOOV KHOLFDOPRUSKRORJ\ KHOLFDVH KHOLFLW\ KHOLRWKHUPDOKHDWLQJ KHOLRWURSLF KHPRSURWHLQ KHSDWRF\WH KHSDWRWR[LFRVLV KHSDWRWR[LQV KHSWDQXFOHRWLGH KHSWDSHSWLGH KHUELFLGH UHVLVWDQFH KHUELYRUHKHUELYRU\ KHWHURF\VW Plate 2d KHWHURWURSKVKHWHURWURSK\ EDFWHULD &\DQREDFWHULD KH[DGHFDQRLG KH[RVH VHHDOVRJOXFRVHIUXFWRVH +LOGDQ +,3 KLJKO\LWHUDWHGSDOLQGURPH VLWHV HYROXWLRQDU\FRQVLGHUDWLRQV SRVVLEOHUHODWLRQWRHQYLURQPHQW SRVVLEOHUROHLQ'1$UHSDLU KLVWLGLQH KLVWLGLQHSURWHLQ NLQDVH KRORW\SH KRPRDFHWDWH SDWKZD\ KRPRDQDWR[LQD KRPRODFWDWH
KRPRVHULQHODFWRQHV KRPRW\URVLQH KRUPRJRQLD KRUPRQH KRUQZRUW +RUVHVKRH0RXQWDLQ$QWDUFWLF +RW/DNH:DVKLQJWRQ +3/&KLJKSHUIRUPDQFHOLTXLGFKURPDWRJUDSK\ UHYHUVHGSKDVH UHYHUVHGSKDVHIDVWSURWHLQ KXPLF +XQDQ3URYLQFH&KLQD +XQWHU V+RW 6SULQJV 2UHJRQ Plate 5a, 5b, 5c +XURQLDQ6XSHUJURXS&DQDGD K\GURJHQSHUR[LGH K\GURJHQVXOSKLGH K\GURJHQDVHV K\GURODVHV K\GURPDJQHVLWH K\GURSKLOLF K\GURSKRELFSURWHLQ K\GURSRQLF K\GURVWDWLFSUHVVXUH K\GUR[DPDWH K\GUR[\EXW\UDWH QK\GUR[\JXDQLGLQHPHWK\OSKRVSKDWH K\GUR[\WHWUDORQHQH K\SHUEUDG\WHO\ K\SHUVDOLQLW\ &KDSWHU K\SHUVDOLYDWLRQ K\SHUHXWURSKLFK\SHUWURSK\ K\SKD K\SROLWKLF K\SRYRODHPLFVKRFN
I ,FHODQG ,FHODQGVSDU LGHQWLILFDWLRQ LPPXQRDVVD\ LPPXQRFKHPLVWU\ LPPXQRJROGODEHOOLQJ LPSHUYLRLXVURFN ,QGLD VHHalso $OO,QGLD &RRUGLQDWHG5LFH,PSURYHPHQW3URMHFW /RQDU /DNH0DGXUDL1DJDODQG 3RRQD ,QGLDQLQN
6XEMHFWOQGH[
,QGLDQ2FHDQ see also $OGDEUD$VWRYH)DUTXKDU 6W3LHUUH SRVVLEOHRFFXUUHQFHRINostoc DWGHSWK LQGROHDFHWLFDFLG LQGROHSKRVSKDWHK\GURODVH ,QGRQHVLD LQGXVLXP LQIHFWLRQSURFHVV LQIHFWLYHVWUDLQVRI&\DQREDFWHULD LQIHFWLYLW\RIF\DQRSKDJH ,QQHU0RQJRQLD LQVHFW VHHDOVR'LSWHUD PLGJH LQVHFWLFLGH WROHUDQFH ,QVWLWXWH)UDQoDLVHGX3pWUROHXP ,QWHUQDWLRQDO%RWDQLFDO&RQJUHVV ,QWHUQDWLRQDO&RGHVRI1RPHQFODWXUH ,QWHUQDWLRQDO&RPPLWWHHRQ7D[RQRP\RI9LUXVHV ,QWHUQDWLRQDO5LFH5HVHDUFK,QVWLWXWH LQWHUWLGDO&KDSWHU LQWUDFHOOXODU&\DQREDFWHULD LQWURQ LRQLVLQJUDGLDWLRQ ,UDT ,UDTLPDUVKHV ,UHODQG see %XUUHQ&DKHU5LYHU /RXJK1HDJK 1,UHODQG LURQ OLPLWDWLRQVWUHVV DQGQLWURJHQDVH RFHDQLFVXSSO\ R[LGDWLRQ R[LGH VXOILGH Plate 8d ,55,,QWHUQDWLRQDO5LFH5HVHDUFK,QVWLWXWH ,VDU5LYHU $XVWULD LVLGLD LVRDONDQHV LVRFLWUDWHGHK\GURJHQDVH LVRPHUDVHV ,VUDHO see also 'DQDUHDVHZDJH WUHDWPHQWSODQW'DQ:DVWHZDWHU5HFODPDWLRQ 3URMHFW/DNH.LQQHUHW0DNKWHVK5DPRQ 1HJHY6RODU/DNH ,WDO\ see also &DVD$XUHXP/DNH0DJJLRUH
J -DPDLFD
-DSDQ see also -LQWVXULYHU /DNH%LZD0LQLPDWD%D\ -DYD -HEHO0DUUD 6XGDQ -HW3URSXOVLRQ/DERUDWRULHV -LQ'\QDVW\ -LQWVXULYHU-DSDQ
K NDQDPLFLQ NDWDEDWLFZLQGV .DWPDL$ODVND NHURJHQ NHWRGHR[\SKRVSKRJOXFRQDWHDOGRODVH NHWRJOXWDUDWH NHWRQHV NLGQH\GDPDJH NLQDVHV .LUNWRQ/LPHVWRQH .ODPDWK/DNH2UHJRQ .RQJVIORUGHQ6YDOEDG1RUZD\ .UDNDWDX .XZDLW
L /D*RUFH0RXQWDLQV$QWDUFWLF ODFWDPDQWLELRWLFV ODFWDWH GHK\GURJHQDVH /DJXQD*XHUUHUR1HJUR0H[LFR /DKRUH3DNLVWDQ /DNH$OH[DQGULQH$XVWUDOLDPlate 11e /DNH$SRSND)ORULGD /DNH$UDQJXDGL(WKLRSLD /DNH$\GDW)UDQFH /DNH%DLNDO5XVVLD /DNH%DODWRQ+XQJDU\ /DNH%DQJZHRORX=DPELD /DNH%HULD6UL/DQND /DNH%LZD-DSDQ /DNH%RGRX&KDG /DNH%RJRULD .HQ\D /DNH%RQQH\$QWDUFWLFD /DNH&KDG$QWDUFWLF /DNH&KDGFHQWUDO$IULFD /DNH&KLWX(WKLRSLD /DNH&KRLFH$QWDUFWLF /DNH&RQVWDQFH*HUPDQ\ /DNH'DULED6XGDQ /DNH'DUWPRXWK$XVWUDOLD
6XEMHFW,QGH[ /DNH(OPHQWHLWD.HQ\D /DNH(UNHQ6ZHGHQ /DNH)U\[HOO$QWDUFWLFD /DNH*HRUJH8JDQGD /DNH*MHUVM¡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late 3B ODPLQDWLRQPlate 1e: see also VWURPDWROLWH /DQJMDQ1DWXUH5HVHUYH1RUWKHUQ3URYLQFH6RXWK $IULFD /DQJPXLUFLUFXODWLRQ /DUVHPDQQ+LOOV$QWDUFWLFD ODUYDH ELYDOYH
FKLURQRPLG JDVWURSRG PLGJH PRVTXLWR /DRV ODWHUDOJHQHWUDQVIHU OD[DSK\FLQV OHDG OHDG]LQFPLQLQJPlate 23 OHDI Azolla, PRVV OHFWLQ OHJXPH OHXFLQH /LE\D OLFKHQ OLJDVHV OLJKW FDSWXUH IDUUHG JUHHQ UHG VSHFWUXP XQGHUZDWHU OLPHVWRQH&KDSWHU &DUERQLIHURXV KLJKPDJQHVLDQ PHWDPRUSKRVHG QXWULHQWFRQWHQW WURSLFDOWHUUHVWULDO /LPIMRUGHQ'HQPDUN OLPLQJ OLQGDQH /LQQDHXV7HUUDFH$QWDUFWLF DFLG OLSLGV ELRPDUNHU OLSRROLJRVDFFKDULGH OLSRSRO\VDFFKDULGH OLWKRVRO /LWKXDQLD /LWWOH&RQHVWRJD&UHHN86$ OLYHU FDUFLQRJHQ IDLOXUH KDHPRUUKDJH QHFURVLV OLYHUZRUW OLYHVWRFN
ORDP /RFK%RUUDOOLH6FRWODQG /RFK,QVK6FRWODQG /RQDU/DNH,QGLD ORQJWHUPYLDELOLW\ /RV$QJHOHV&DOLIRUQLD /RXJK1HDJK1,UHODQG /RZHU*H\VHU%DVLQ
M 0DF4XDULH,VODQG PDFUROLGH 0DGXUDL,QGLD PDJQHVLXP KLJKFRQWHQWLQOLPHVWRQH LRQV ORZFRQWHQWLQOLPHVWRQH PDJQHWRWDFWLFUHVSRQVH PDL]H 0DNKWHVK5DPRQ,VUDHO PDODWH PDODWKLRQ 0DOKDP(QJODQG 0DOL PDOWRVH PDPPDOV 0DPPRWK+RW6SULQJV7HUUDFHV 0DQJDOLD8NUDLQH PDQJDQHVH LRQV R[LGH PDQJURYH Plate 25a 0DQLNJDQM%DQJODGHVKPlate 25d PDQQLWRO PDQQRVH 'PDQQXURQLFDFLG 0DU6ORXJK&DOLIRUQLD PDUEOH 0DUH V(JJ6SULQJ2UHJRQPlate 24a 0DULH%\UG/DQG$QWDUFWLF PDUO
6XEMHFW,QGH[ 0DUV PDVVFXOWLYDWLRQ PDVVVSHFWURPHWU\ PDWUL[DVVLVWHG ODVHUGHVRUSWLRQLRQLVDWLRQ 0$/', PDWV&KDSWHU&KDSWHU &KDSWHU EOLVWHU FROXPQDUOLIWRII GHHSIODW PRDW SLQQDFOH SRO\JRQDO Plate 19b SURVWUDWH SXVWXODU Plate 8c, 8e, 8g VKDOORZIODW VPRRWK Plate Sa, 8b, 8d, 8f PDWWRFNLQJ 0DXGH:HLU0XUUXPELGJHH5LYHU 0DZVRQ5RFN$QWDUFWLF 0F0XUGR 'U\9DOOH\V &KDSWHU ,FH6KHOI 6RXQG PHJDVSRURFDUS 0HOOXPLVODQG *HUPDQ\ PHOWZDWHU PHUFXULFR[LGH PHUFXU\ UHGXFWDVH PHWDOLPQLFPHWDOLPQLRQ PHWDOORWKLRQHLQ PHWHRJHQH 0HWHRUSDSLHU PHWKDQH PHWKDQRJHQ /PHWKLRQLQH'/ VXOIR[LPLQH06; PHWKR[\SKHQDQWKUHQH PHWK\ODWHGDPLQHV PHWK\OHQHEOXH PHWK\OIRUPDPLGH PHWK\OQDSKWKDOHQHV PHWK\O SHQWDD]DELF\FOR>@QRQDQ WULHQRQH 0H[LFR see also ([SRUWDGRUD GH6DO *XHUUHUR1HJUR/DNH7H[FRFR 7HKXDFDQ PLFULWH PLFURF\VWLQV&KDSWHU VWUXFWXUH PLFURHOHFWURGH
6XEMHFW,QGH[ PLFURIRVVLOV PLGJH 0LHVFDWWHU PLQHVSRLOZDVWHPlate 23 0LQLPDWD%D\-DSDQ 0LRFHQH0RQWHUH\)RUPDWLRQ 0LVVVRXUL%RWDQLF*DUGHQ PLWRP\FLQ& PL[LQJ DUWLILFLDO WXUEXOHQW 0RDE8WDK PROHFXODUFORFN PROOXVF PRO\EGHQXP 0RQJROLD PRQRFORQDODQWLERG\ PRQRJDODFWRV\OGLDF\OJO\FHURO 0RQWHYLGHR8UXJXD\ PRQXPHQWV 0RVHVFDYH6RXWK$IULFD PRVTXLWRHV PRVV PRWLOLW\ JOLGLQJ SKRWRUHVSRQVHV see also SKRWRSKRWRSKRELFSKRWRWD[LV VLJQDOOLQJ VZLPPLQJ YHUWLFDO PRXVHDVVD\V 0RXQW)OHPLQJ$QWDUFWLF 06; see /PHWKLRQLQH'/ VXOIR[LPLQH 0W$XFDQTXLOFKH&KLOH PXFLODJH 0XOX3DUN,QGRQHVLD 0XUUD\-DPHV 0XUUXPELGJHH5LYHU$XVWUDOLD PXVFOHIDVFLFXODWLRQV 0XVKURRP6SULQJ
N 1$'+GHK\GURJHQDVH
1$'3+ 1DJDODQG,QGLD 1DPLELD QDQDIODJHOODWHV QDSWKDOHQH QDSWKHQH QDSWKRO 1$6$ QDWURQODNHV QHFULGLXP QHHP 1HJHYGHVHUW Plate 22a, 22b, 22c, 22d, 22e QHPDWRGHV 1HW3HQ/LYHU'LVHDVH 1HWKHUODQGV QHXURWR[LQ 1HXVH5LYHUHVWXDU\ Plate 10d QHXWUDOUHG 1HYDGD 1HZ0H[LFR 1HZ6LEHULDQ,VODQGV 1HZ=HDODQG QLFNHO 1LQJ[LD3URYLQFH&KLQD QLWUDWH DVVLPLODWLRQ UHGXFWDVH WUDQVSRUWWUDQVSRUWHU QLWULI\LQJEDFWHULD QLWULWH DVVLPLODWLRQ UHGXFWDVH WUDQVSRUW WUDQVSRUWHU QLWURJHQ DVVLPLODWLRQ H[WUDFHOOXODU IL[DWLRQ &KDSWHU JOREDOUHJXODWLRQ OLPLWDWLRQ RUJDQLF VHHDOVRXUHD SKRVSKRUXVUDWLR VWRUDJH QLWURJHQDVH SQLWURSKHQ\O SKRVSKDWH 1L]]DQD1HJHY 105QXFOHDUPDJQHWLFUHVRQDQFH
QRGXODULQ 1RPDUVNLLQWHUIHUHQFHPLFURVFRS\Plate 24e QRQDNHWLGH 1RUGHQVNLROG$GROI(ULN 1RUULV%DVLQ
O 2DNODQG&DOLIRUQLD RFWDGHFDQRLFDFLG 2FWRSXV 6SULQJ
6XEMHFW,QGH[ VHQVLWLYLW\ VXSHUVDWXUDWLRQ R[\JHQDVH 2]RQH+ROH R]RQRO\VLV
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phagnum FHOOV LQ Synechococcus PDW SKDHRVRPH SKDJRF\WRVLV
6XEMHFW,QGH[ 3KDQHUR]RLF SKDUPDFHXWLFDO SKHQDQWKUHQH SKHQDQWKUROV SKHQDQWKUROLQH SKHQROLFFRPSRXQGV SKHQROV SKHQ\ODODQ\OW51$V\QWKHWDVH SKHQ\OSURSDQRLG 3KLOLSSLQHV see ,QWHUQDWLRQDO5LFH5HVHDUFK ,QVWLWXWH/DNH/DQDR SKORHP SKRUEROHVWHU SKRVSKRULWH SKRVSKDWDVH H[WUDFHOOXODU SKRVSKDWHSKRVSKRUXV DLUERUQH IHUWLOL]DWLRQRIPDW IHUWLOL]DWLRQRIULFHILHOGV IUHVKZDWHUSODQNWRQ DQG Gloeotrichia, LQKLELWLRQRIFDOFLWHQXFOHDWLRQ OLPLWDWLRQVWUHVV SKRVSKRUXV ORZDPELHQWOHYHO LQPDWV DQG Microcystis, RFHDQLF RUJDQLF RYHUSOXV UHGXFWLRQUHPRYDO LQURFN SUHFLSLWDWLRQ SXOVHHIIHFW UHPRYDO DQGVDOLQLW\ VHDVRQDOFKDQJHV LQVRLOV VWLPXODWLRQRIQLWURJHQDVHDFWLYLW\ DQGWR[LQV ZKROHODNHHQULFKPHQW DQG]LQFWR[LFLW\ SKRVSKDWLG\OFKROLQH SKRVSKDWLG\OJO\FHURO SKRVSKRGLHVWHUDVH SKRVSKRIUXFWRNLQDVH SKRVSKRJOXFRQDWHGHK\GUDVH SKRVSKRJO\FRODWH SKRVSKROLSLGV SKRVKRPRQRHVWHUV SKRVSKRPRQRHVWHUDVH SKRVSKRUHOD\SDWKZD\
SKRVSKRULEXORVH NLQDVH SKRVSKRULERV\O DPLQRLPLGD]ROH FDUER[\ODVH SKRVSKRVHULQH SKRVSKRWKUHRQLQH SKRVSKRW\URVLQH SKRWR DGDSWDWLRQ SKRWREOHDFK SKRWRGDPDJH SKRWRH[FUHWLRQ SKRWRKHWHURWURSK\ SKRWRLQKLELWLRQ SKRWRNLQHVLV SKRWRR[LGDWLRQ SKRWRSKRELF see also PRWLOLW\ SKRWRUHDFWLYDWLRQ SKRWRUHFHSWRU SKRWRUHVSLUDWLRQ SKRWRV\PELRGHPH SKRWRV\QWKHVLV LQKLELWLRQ SKRWRV\VWHP, SKRWRV\VWHP,, SKRWRWD[LV VHHalso PRWLOLW\ SK\FRELOLQVSK\FRELOLSURWHLQV see also SK\FRF\DQLQ SK\FRHU\WKULQ SK\FRELOLVRPH SK\FRF\DQLQ SK\FRHU\WKULQ SK\FRHU\WKURELOLQ SK\FRP\FHWH SK\FRXURELOLQ SK\ORJHQHWLFUHODWLRQVKLSWUHH SK\WLFDFLG SK\WRFKURPH SK\WRSDWKRJHQ 3,FXUYH SLFRF\DQREDFWHULD &KDSWHU SLFRHXNDU\RWH SLQQDFOHV SODVPDGHVPDWD SODVPDOHPPD SODVPLG SODVWLG SODVWRF\DQLQ SODVWRTXLQRO
SODVWRTXLQRQH34 SODWLQ SODWLQJWHFKQLTXH SORXJKLQJ SQHXPDWRSKRUH Plate 25a SRLVRQLQJVHHWR[LQV SROOXWLRQ SRO\FORQDODQWLERG\ SRO\JOXFDQJUDQXOHVSRO\JOXFRVH SRO\KDOLQH SRO\KHGUDOERG\ see FDUER[\VRPH SRO\NHWLGH 3RO\QHVLD SRO\RO SRO\SHSWLGH SRO\SKRVSKDWDVH SRO\SKRVSKDWH ERG\JUDQXOH NLQDVH SRO\SORLG\ SRO\VDFFKDULGH &KDSWHU VXOIDWHG SRO\VXOILGH SRO\XUHWKDQH SRQG PDQDJHPHQW RXWGRRUFXOWXUHPlate 27a, 27b ZDVWHVWDELOL]DWLRQ 3RRQD,QGLD SRUHPHPEUDQH LQ Geosiphon, ZDWHU SRUSK\UD 3RUW$UDQVDV7H[DV SRWDVVLXPLRQV SRWDVVLXPSKRVSKDWH 3UHFDPEULDQ SUHFLSLWDWLRQ DWPRVSKHULF PLQHUDO see also FDOFLWH SUHGDWLRQsee also JUD]LQJ SUHVHUYDWLRQRISLFRSODQNWRQ 3ULHVWOH\*ODFLHU$QWDUFWLF SULPDVHV 3ULQFHVV(OL]DEHWK/DQG$QWDUFWLF 3ULQFLSDO&RPSRQHQW$QDO\VLV SURFKORURSK\WHVsee also Prochlorococcus, Prochloron, Prochlorothrix SURGLJLRVLQ
6XEMHFW,QGH[ SURKHWHURF\VW SUROLQH SURPRWHUELQGLQJIDFWRUV SURSLRQDWH SURWHDVH SURWHLQ GHJUDGDWLRQ see also SURWHDVHV NLQDVH SKRVSKDWDVHV SKRVSKDWDVHLQKLELWLRQDVVD\ SKRVSKRU\ODWLRQ 89VKRFN 3URWHUR]RLF SURWRQHPD Plate 23c, 23f SURWRSODVW SURWR]RD see also DPRHED FLOLDWH SVHXGRILODPHQW SV\FKURSKLOLF SV\FKURWROHUDQW SWHULGRSK\WH see also Azolla 3XHUWR5LFR SXULQHELRV\QWKHVLV SXUSOHVXOIXU EDFWHULD 3\UDPLG/DNH86$ S\ULGLQHV QXFOHRWLGH S\ULPLGLQHV S\ULWH S\UURORS\ULPLGLQH
Q TXDUW] 4XHEHFODNHV
R UDLQZDWHU 5DPDQVSHFWURVFRS\ 5DQJLURD$WROO 5DQJRRQ0\DQPDU UDVKVNLQ 5D\OHLJKVFDWWHU UHFRJQLWLRQSURFHVVHV UHFRPELQDWLRQ UHGOLJKW see also FKURPDWLFDGDSWDWLRQ 5HG6HD see also 6RODU/DNH 5HGILHOGUDWLR UHGR[ FRQWURO SURWHLQ
6XEMHFW,QGH[ VHQVLQJ VWDWH UHHIV UHJXODWHGULYHU UHK\GUDWLRQ 5HVROXWH%D\ UHVSLUDWLRQ 5)/3 UKDPQRVH UKL]RELD UKL]RPH UKL]RSRG ULERVH ULERVRPHV ULEXORVHELVSKRVSKDWH FDUER[\ODVH ULEXORVHSKRVSKDWH ULEXORVHSKRVSKDWHNLQDVH ULFH ILHOG&KDSWHUHVS JHUPLQDWLRQ JUDLQ\LHOG VWUDZ 5LIW9DOOH\V(DVW$IULFD 51$ KHOLFDVH P51$ SRO\PHUDVH 6U51$ W51$V\QWKHWDVH 5REHUWVNROOHQ$QWDUFWLF 5RPDQLD URRW DSRJHRWURSLF FRUDOORLG H[XGDWHV ODWHUDO QRGXOH 5RVV'HVHUW ,VODQG 5RWK$OEUHFKW URWLIHU 5XELV&2 5XWODQG:DWHU(QJODQG 5XVVLD
VDEKND &KDSWHU VDFULILFLDOFHOO 6DKDUD 6DOGD/DNH7XUNH\
VDOLQHVDOLQLW\&KDSWHU 6DOLQVGH*LUDXG)UDQFH VDOLWH VDOPRQ VDOWFU\VWDOL]DWLRQ VDOWHUQSRQGV 6DQ'LHJR&DOLIRUQLD 6DQ6DOYDGRU,VODQG%DKDPDV VDQG VDQGGXQH 6DQWD%DUEDUD&DOLIRUQLD 6DUJDVVR6HD VDWHOOLWHVHQVLQJ 6DWRQGD&UDWHU/DNH,QGRQHVLD 6DXGL$UDELDPlate 1h VD[LWR[LQ VFDQQLQJHOHFWURQPLFURVFRS\ VFDULGILVK 6FKLUPDFKHU2DVLV$QWDUFWLF 6FRWODQG see also IORZFRXQWU\ /RFK%RUUDOOLH/RFK,QVK:HVW/RWKLDQ 6FRWW5REHUW)DOFRQ VF\WRQHPLQ Plate 24e, 26f, 31, 32 VF\WRSK\FLQ VHDJUDVV VHDXUFKLQ VHFRQGDU\PHWDEROLWH&KDSWHU VHGLPHQWDWLRQ VHGRKHSWXORVHELVSKRVSKDWDVH 6(0VFDQQLQJHOHFWURQPLFURVFRS\ 6HQHJDO FRUDOORLGURRWV Azolla OHDI VHSWD VHULDWHPRUSKRORJ\RINostoc), 6HUHQGLSLW\0HDGRZV
VKHDWK SLJPHQWDWLRQ see JORHRFDSVLQVF\WRQHPLQ VKHHS VKHOOILVK VKLQRULQ VKRFN SURWHLQ KHDW R[LGDWLYH VWUHVV 89 VKULPS 6LEHULD VLGHURSKRUHV VLJQDOV 6LJQ\ ,VODQG VLOLFHRXV KRWVSULQJ VLOLFD VLOLFDWH VLOLFRIODJHOODWHV VLOLFRQ VLON 6LQDL see also 6RODU /DNH VLQJOHW 2 VLQNLQJ VNLQUDVK 6ORYDNLD VQRZ VRGLXP LRQV VRGLXP GLWKLRFDUEDPDWH VRLO &KDSWHU FDWHJRULHV IRU DOJDH FOD\ HQULFKPHQWFXOWXUH ORDP UHFRYHU\ VKHDU VWUHQJWK YROFDQLF 6RODU /DNH VRODU SRQG IRUPHUO\,VUDHOQRZ (J\SW VRODUSRQGVVDOWHUQV 6RPDOLODQG 6RQDUDQ 'HVHUW 6RQDUJDRQ %DQJODGHVK VRUELWRO 6RVD7H[FRFRFRPSDQ\ 5 VRXS Plate l i 6RXWK$IULFD see also &ODUHQV6DQGVWRQH *ROGHQ *DWH +LJKODQG 1DWLRQDO 3DUN +DUWHEHHVSRRUW 'DP /DQJMDQ1DWXUH5HVHUYH 0RVHV FDYH 6RXWKHUQ K\EULGL]DWLRQ
6XEMHFW ,QGH[ 6RXWKHUQ9LFWRULD/DQG 6SDLQ see also $OKDPEUD $OLFDQWH /DNH6DQWD2ODOOD9DOHQFLD VSHFWUDOFRPSRVLWLRQ VSHOHRWKHP 6SHQFHU*XOI$XVWUDOLD VSLUDOPRUSKRORJ\ 6SLUXOLQD0H[LFDQDFRPSDQ\ 5 6SLWVEHUJHQ VSRLOKHDS VSRQJH VSRURFDUS VSRUWV WXUI VSRUXODWLRQ see DNLQHWHEDHRF\WH EDFWHULDO 6UL /DQND 6W3LHUUHLVODQG ,QGLDQ 2FHDQ VWHDG\VWDWH 6WHFKOLQVHH*HUPDQ\ VWHPJODQG VWHQRKDOLQH 6WRNHVHTXDWLRQ VWUDWLILFDWLRQ VWUDWLJUDSK\ VWUDZ EDUOH\ ULFH VWUHDPV &KDSWHU &KDSWHU $UFWLF VWURPDWROLWH &KDSWHU GHILQLWLRQ VWURQWLXP VW\URIRDP VXEOLWKLF VXEOLWWRUDO&KDSWHU VXFFHVVLRQ IUHVKZDWHUURFN WHUUHVWULDO VXFFLQDWH VXFURVH VXFURVHSKRVSKDWHV\QWKDVH 6XGDQ 5 6XGDQ6DKHO]RQH5 , 6XIIRON:DWHU%RDUG (QJODQG VXJDUEHHW VXJDUFDQHZDVWH VXOIDWH 5 ELQGLQJSURWHLQ UHGXFLQJEDFWHULD Plate 17c UHGXFWLRQ XSWDNH
6XEMHFW ,QGH[ VXOILGH VXOSKLGH 40, 41,66,68,71,77,81,83,91,
100-102,105-108,265,286,287, 292,294,382, 444 WROHUDQFH 81,83 VXOILWH 82,427 VXOIRTXLQRYRV\OGLDF\OJO\FHUROV 315 VXOIRWUDQVIHUDVH SDWKZD\ 427 VXOIXU 88,244,308,309,386 F\FOH 106 HOHFWURQ DFFHSWRU 88 GHILFLHQF\ OLPLWDWLRQ 386,427-428 6XQELJJLQ (QJODQG Plate 26c VXSHUR[LGH DQLRQ 604 GLVPXWDVH 52,548,604 UDGLFDO 548 VXSHUSKRVSKDWH IHUWLOL]HU 245 VXSUD OLWWRUDO 598 VXUIDFHWRYROXPH UDWLR 21 6XUWVH\,FHODQG242 6YDOEDUG 335 6ZDOH 5LYHU (QJODQG Plate l g 6ZHGHQ 199,200, 201,203,207,219,223,537 VZLPPHU V LWFK 626 VZLPPLQJ PRWLOLW\ &\DQREDFWHULD 208,382,383 6ZLW]HUODQG 203 V\PELRQW V\PELRVLV 8,125,132,142,246-247, 474,&KDSWHU 19, V\PELRVRPH 545 V\QHUJLVWLF HIIHFW ZLWK WR[LQV 623
T WDLO F\DQRSKDJH 565,581 7DPDQUDVVHW $OJHULD 512 WDSLRFD IDFWRU\ 512 WDU VDQGV 243 WDUGLJUDGH 337 WDVWH 615 WDXWRP\FLQ 625 WD[RQRP\ F\DQREDFWHULDO 330,497 F\DQRSKDJH 565 7D\ORU *ULIILWK 322 7D\ORU 9DOOH\ 346 7HKXDFDQ 0H[LFR 242 WHPSHUDWXUH JUDGLHQW SODWH 247 7HPSOH RI $SROOR 'LGLP 7XUNH\ 274 7HUUD 1RYD %D\ $QWDUFWLF 328 WHWUHS\UUROH 405,407, 593 7H[HO 7KH 1HWKHUODQGV 103 7KDLODQG 236,51 WKDODVVRKDOLQH 283,285
667 WKHUPRFOLQH 167 WKHUPRJHQH 269 WKHUPRSKRELF 54 WKHUPRWROHUDQW 42 WKLRF\DQDWH 427 WKLRUHGR[LQ 376,377,386 WKLRVXOIDWH 82,300,427,428 WK\ODNRLG 315,381,387,388,399,400,411,475,
507,508,536
7LPEHU 3HDN $QWDUFWLFD 350,355 WLQWHQVWULFKH 260-261,270,272,273,275,352, 468,602,Plate 16a, 30b 7ROHUDEOH 'DLO\ ,QWDNH 7', 624 WRPDWR 528 WR[LQV 8,150,&KDSWHU 22 LQIOXHQFH RI SKRVSKDWH 617 SURGXFWLRQ 570 ]LQF UHTXLUHPHQW IRU SURGXFWLRQ 449
trans-9,10-dihydroxy-9,10-dihydrophenanthrene, 314 WUDQVFULSWLRQDO DFWLYDWRUV 386 LQLWLDWLRQ 423 trans- , GLPHWK\OWUDQVGHFDOJHRVPLQ 496 WUDQVIHUDVHV 444 WUDQVSRUWHUV 422,425,448 WUDQVSRVRQ 485,539,546 WUDSSLQJ SDUWLFOHV 265,267 WUDYHUWLQH &KDSWHU 9 WUHHV 242 WUHKDORVH 85,86,282,296,297,351,355,356 WULFKORUIRQ 472 WULIOXRSHUD]LQH 371 WULPHWK\ODPLQH 85 WULRVH SKRVSKDWH 422 WULSDUWLWH OLFKHQ 547,549,550,551 WULSKHQ\O WHWUD]ROLXP FKORULGH 77& 135,136,488 WURXW 626 WU\SVLQ 508 WU\SWRSKDQ 333,418 7VKLSLVH 6DQGVWRQH 358 77& WULSKHQ\O WHWUD]ROLXP FKORULGH 136,488 WXEHUFLGLQ 618 WXPRXU SURPRWLRQ 622,625 VXSSUHVVRU SURWHLQ 625 WXQGUD VWUHDPV 327 WXQLF FHOOV 532,535 WXQLFDWH 337 WXUEXOHQFH 129,131,140-142,&KDSWHU 6 HVS 158-160-162,173),595 WXUJRU SUHVVXUH 154-159 7XUNH\ 262,268,274
6XEMHFW ,QGH[
7XVFDQ\ ,WDO\ 7ZLQ %XWWH 9LVWD 6SULQJV
U XELTXLWLQ 8NUDLQH XOWUDROLJRWURSKLF XOWUDVRXQG XOWUDVWUXFWXUH XOWUDYLROHW UDGLDWLRQ &KDSWHU 8QLYHUVLW\ 9DOOH\ %HDFRQ +HLJKWV $QWDUFWLF 8SSHU 7HHVGDOH (QJODQG XSWDNH K\GURJHQDVH XUEDQ UXQRII XUHD XUHDVH XULG\O\O WUDQVIHUDVH 8QLWHG .LQJGRP XURQLF DFLGV 8UXJXD\ 86$ VHH also %HDXIRUW %LUG 6KRDO &DOLIRUQLD &DSH &RG &RORUDGR 3ODWHDX &URRNHG /DNH (YHUJODGHV +XQWHU V +RW 6SULQJV -HW 3URSXOVLRQ /DERUDWRULHV .ODPDWK /DNH /DNH +XURQ /DNH 0HQGRWD /DNH 0LFKLJDQ /DNH :DVKLQJWRQ /LWWOH &RQHVWRJD &UHHN 0DUH V (JJ 6SULQJ 1$6$ 1HXVH 5LYHU HVWXDU\ 3HQQV\OYDQLD ODNHV 3RUW $UDQVDV 8WDK :DOGR /DNH :RRGV +ROH +DUERU
V YDFXROH JDV &KDSWHU HVS OLTXLG 9DOHQFLD 6SDLQ YDQDGLXP YDQ /HHXZHQKRHN
YDUQLVK GHVHUW YDVFXODU SODQWV 9DXFKHU -HDQ 3LHUUH (WLHQQH YHKLFOH XVH 9HVWIROG +LOOV $QWDUFWLF YLDELOLW\ WHVWV 9LFWRULD /DQG $QWDUFWLF 9LNLQJ 3URJUDPPH YLRODFHLQ YLWDPLQ YLWDPLQ % YROFDQLF 9ROOHQZHLGHU2(&' PRGHO YRPLWLQJ
W :DGL 1DWUXQ (J\SW :DOGR /DNH 2UHJRQ ZDVWH VWDELOL]DWLRQ SRQG ZDWHUEORRP VHH EORRP :DWHUIDOO %HFN 0DOKDP (QJODQG ZDWHUORJJLQJ ZDWHU SRWHQWLDO VWUHVV ZHDWKHULQJ :HVW /RWKLDQ 6FRWODQG ZKHDW :KLWH 0W 6XPPLW &DOLIRUQLD ZLQG DFWLRQ VHH DOVR NDWDEDWLF ZLQGV ZLQG VXUIHU :RRGV +ROH +DUERU :RUOG +HDOWK 2UJDQL]DWLRQ
X [DQWKLQHV [DQWKRSK\OO [DQWKRSK\WH [\ORVH
Y \HDVW
6XEMHFW ,QGH[
Z =DLUH 1 =DPELD ]HD[DQWKLQ ]LQF &KDSWHUPlate 23 GHILFLHQF\ LQ ULFH ILQJHUSURWHLQ KRPRHRVWDVLV RFHDQLF DQG WR[LQ SURGXFWLRQ WULDG ]RRSODQNWRQ ]\PRJUDP