Conjugation and Deconjugation of Ubiquitin Family
Volume 54
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Conjugation and Deconjugation of Ubiquitin Family
Volume 54
SUBCELLULAR BIOCHEMISTRY SERIES EDITOR !" #!$ !$ %!
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Conjugation and Deconjugation of Ubiquitin Family
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Marcus Groettrup
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Frederick C. Streich, Jr. and Arthur L. Haas
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Alexander Buchberger
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ACKNOWLEDGEMENT +2!# 2@ 22!2 + %^N__W 0
REFERENCES Z% 3! * !" & = !>2 2 2@H !# 3#
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REFERENCES Z . * NYL2WXP! # !H * !" ! +! (.! | ^H * . # = !>+ )!H )>|* \__[HZWN>\ZZ \ ! ! + ) ) 2! H " !.! @ @3 " * \__MK [HMZW>\W ^) 0 !. &+ H "!." ## ! !@ ! " !&# \__XKYHZ[MX>[X N2 22& % $!#2 !> +&\^! 2!# *(*NYL 8(ZJ+0NP = !> " 2!*\__ZKZ_XH[[X>XX M+|' ("! .. 0*%!. .! H &+ #2WX \__[KZM[HZ\>\Y [ * -! @! !Q!M ! 2!!# 8>!2WX> 2@ #%. !# %.#.! !"*ZWWYKW\H[_^>Z_ X)!.v )!%|*+Z^M = !!.!$ !.2WX>2NX> # %.#.! *\__NKZ[NHWX^>Y
79
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" 'ZWW[KZMHNYYN>WW ZM 2 ! # = ! !!.# !! *(*NYJ2WX = !!2 .!.*\__MKZ\_HX^>YN Z[! +* - & = !> " 2!*(*>NYJ2WX!3 ! !2*\__XKWH^XW>W_ ZX-! |' 8 ' | |*+ @2 +. # !! #!2!H"@# != 2 \__YKZXY\HXNN>Y ZY
* . B!2! H;. # &+ * NYJ2WX* 0#
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* 2# 2Z! Q\2 #* NY!""! = !>2!! 2!.!'2\__NKMHYZY>\N \[( .! '"! % !!!.#* NY2 = !> B!#!" 2! !""!! 2 ! 2 !v \__NK\ZHZ\X>^W \X !!>+ ! ) #!!-& Q\! Q^ # * NY " 2 !!!2 = !>2!!2 * \__NKZNHY\N>Y \Y Q! % % !! %& B(X ! 2 = !. !2 2WX! 8Z2 !"*\__YKZ^NHY_N>Z[ \W * & &+ 2WXJ|*+ ! @ !"
# #Z>2N ! 2NX . ! 2 !!. ! * \__NK \XWHNW[_W>Z[ ^_ 2# ! 8 ! !" !# 2 !. # # = !> "* NY
2!*\__[K\ZH\[Z>W ^Z )#( (Z2 2!= #.!## 2! #!2 H2.!(Z>32! 8' v \__NKNHX\Z>W ^\ - 2!vv (!(#Z! @* NY!# ! ! !' v \__[K\^HZ_M^>[N ^^ % 3 !.!!J !> # #|*+J2WX> !#Q!>^ .! '\__[K\MHZMNX>MY ^N+ !v v!. &#!"2WXJ" !> !!!.2!>! !.## .2XY!!2 > .!*\__[K\YZH^M^MW>[Y ^M % *!# 0 *@ '" ! ".! # " !> !!!. 2!J " ! 2! NY !!. ! ! !. !2 > .! 2! 8' \__WK\X[HZ\_Y>\_ ^[ ! 0 + |' 2 ! + ( ! . ! 2WX>2N> #Z !# *\__XK\Y\H\Z^[Z>W ^X(0 !. &*2 #2WXJ" !> !!!.2!" ! ! @!! ! \__^KZ_HYM[>[^
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2Q 2NX! #Z>2N'\__\K\ZHM[NM>M\ MX ) )" 3 - % (NJ+0N = #2 2 !.# = !'2! *\__ZKZ\HNZZN>\Y MYvv 22& & &+ * NYJ2WX! 2! ! 22! # '! \__ZKNZNH[M\>[ MW ' &Q * |3@!*+! !#'= 2 = !!! > &+ * NY*\__\KNHZ^N>W [_ !" ' - 8 - > &+ 2WXJ* NY2 2!= # !2 > 2!.!*\__\K\\H[\[>^N [Z ! @ 3- 2# = !> "*(*NYL 8(ZJ+0NP 2! L .. P!' (#0'Z! '\__\K\ZH[ZM>\Z [\8 B .* 8!.(* NY2 = # !2! ".! #%Z>*(-! 8Z2*\__^KZ[^H\Z>[ [^*- 3; & > &+ * NYJ2WX. 2! !# *\__^KZZMH^MM>[X [N | *!.' (23!&.2 # !>*3 *\__NKZNH[WW>XZZ [M|!.,+ *- 0 v* .!!! ..!. = !!# ""!
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REFERENCES Z- v v *!!.# ( @ ! !" = !>32! 2 * !ZWW^KZWMH^W^>^WW \! +@3 % = !! 3!H@ # &! *\___K Z_H^^M>^N\ ^032 ( (!. % @ 3- !"2! !2@ = ! 'ZWWYKZXH\\_Y>\\ZN N(!.B) ( ) ) !!! #*+W .! ! ! H! !. #"2!&! %!\___KZ[H\_\>\_^ M) (!.B)*+WH!@.! !""!.>. "2!!.!Q2 ! ! 2 +!*ZWW\KNHZM_X>ZMZY [*"$( ) !&&*+W 2Q !" !! . !""!. !#2!"2! @ *ZWW[KY[HZZM>Z\Z X) (!.B)* $!!2 !#!*+W 2Q ! "! . !! 2 #22.! !.2! + + ZWWYK[YH\^X>\NZ Y . -# 8- !"2! 2Q!""! .!! !2 !. \[ 2 ! 8 'ZWWYKZ\HN[W>NXY W
#!!-+ !# ! ###2 *+W> .! !8^ 2Q *!# \__MK[HXZ Z_-2 > ( .' ! 2! !> !! ! " #*+W .! \___K^__HZZ[W>ZZXY ZZ% 3! !( * Q 2Q#2 . 2 = # = !> !; . .! ! *+W> .! ! 8^ * ZWWYK WNH[ZM>[\^ Z\ ! ' ' #*+W .! 2Q .. # ! ! \__WKZXH^Z>N_ Z^(!!' 0" = + (|0& 2 *+W .! !#%\ 2 2. !!.! ("2!\__YKZ^MH\_Z^>\_\\ ZN0 * +@- !-*2WJ .! ! !+ N. ! 32!>2!!!>!2!! ! %! ("\__^KZXHZZ^_>ZZN_ ZM % ! +!@ 3 - 3 & = ! . " ! ((\ ! *
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" 75'!$ !2! !""! !#2M^ = !# = ! # L 8K # 2M^H = ! '((Y ! >ZP " 2 " 'Z "!. !$ L = ! ZK '((Y +++Z> ^K >Z 'Z> '\P Q2! # &+ & " 8 ! ! # 2 " '\ !; .!. !$ L# 2M^H = ! M J Z^K '((Y Z\K >Z WP 8! '\ ! ! @ 2 " '^ 8 . L# '^ !"" ! 2M^ ! & ZP $ "! !# 2 " 82M^8Q2 M ! M 2 "!"" ! \> = !! # 2M^ @ ! ! = !! 2 = !! L2 -NY>!3 = ! ! P # 2M^ 8 ! # 2M^ @ -NY>!3 = ! ! = 2 ! #2^__J*+ @ 'N = ! ! # ) ! = !! ## $! # 2M^ L! Q2 ! ! !P ! # 2M^ @ = ! ! ! 2M^ .! L-NY>!3 ! P
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! !! ! 2 !!! 2!2M^.! !
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CONCLUSION !2 @ 2M^ !# !! 2! ! ! = ! # ! # !2@ 2 ## 2M^ !""!. !#2M^" ! !# ; !. 2 " !#!! 2! " .
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ACKNOWLEDGEMENTS (B 8@# !#!!!*! L *P )3! # 22 ( 8 !. .! # ' 2! * ! L8+[ *P ! %!> 8 !! #
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ACKNOWLEDGEMENTS )!3 Q! $ '!!$!$!-!#!. #! 2&@3# @ 22 .! #( 8 !. .! # !!!' 2!@3#'Q ! *
REFERENCES Z & )#( '!2 .!H" 2!@#! !8 ' ZWWXKZZLZNPHZ\\X>Z\^^ \+2- )#( .2!! !H' ( !# 2! !! &!
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REFERENCES Z% ! + ! . !! "*("\__^KZWHM[M>YX \ 8 8 + #> 2!$ + ! +.! \__[K[_HMM^>[\ ^0 2 w{8 & &' 2 2 ZWWXK\NHZ\M>^Z N(@!- +3 = !>03+! + !+.! " \__WK XHNYM>WZ M ) )$ w{8&2 H+.# #> 2!$!.2 * ZWWYKW\H^[X>Y_ [0!% % & * 2 .! ! .2 @ .22 2 \__[KMWHZN_M>Z[ X+ > * ! )* $!#\_ 2 # ! 8!3 \___K^MH^[Y>XX Y& & . 0 2 & $!# 2 H\_ 2Q#
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^2!*\___K\_H\^[X>XX ^v'& %!.0 -!& = !>32! H!@@! !!@ %!\___K\NYHZ>ZN N%!>'!& . @- v!0! !! $!#('Z ! # 0+#*\__^K\XYH\YYW\>W__ M*!. @& 0+# !!"2!#2 ! 2 \__[KYH[[X>X[ [ v '& ! ! (> !H @ !. @ #/ 2 * \__WK \YNHY\\^>X X & > 2 2 H@ !&! *\__XKZXH^X_>[ Y ! - 3! | !. MM| 22 ! !" .! L >NP ##! " 3 ! 2! # ! @ 2 2 *\__NK\XWH\X\^^>Y W@ ( -' v'& 2!>W_ != >N2"! !! ! 2 * !\__MK^^XHMZX>\_ Z_ % " + 8 v8 ! 2 = #@!.>. !! 2 !! # 00 , &>'0 &'((8' !; . +!*\__^K ZMH\^_Y>ZW ZZ* & 3!! > !; .!.! > !; .!.!$ # 2 +!+ \__[KZN\H^ZY>^\ Z\* 3; 0 0'" !# .!!. ! 2 !! !" H 2 ! \__[K^WYHM\Z>W Z^ " ( 0 *( # '+\ 2 ! ! @ 2 !
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" 2.X2 +!> "!.!$# 2 ! !. ! 2.Z\2 % &'>Z[ % +! +>0*^*\__ZK\X[HZX_Z>[ ^_- &! !&&*>!.!#! 2.X2J*"\2 = #$! ! !# 'Z "!'Z>'\ 2Q#!*\__ZK\X[HWYN[>MN ^Z 0 ) | = !!H !! = ! "! ZWY^K\\HN^YY>WN ^\ 0 & ! # = ! "!.!$ 3! != ! *ZWY\K\MXHZ_^\W>^X ^^* !" ' *"!#!2 !# = !> "!.!$ *ZWY\K\MXH\M^X>N\ ^N+ 3* ! !!. = !! !! " \__ZKX_HM_^>^^ ^M v !# 2.H@ = !>3 "*\__ZK \H\ZZ>[ ^[ v - 3& &3& = !>3 2!2! \___K N_YHNYY>W\ ^X v $ &@ = !>3 !; .! !# 2. !*(" \__NKZMH\^Z>[ ^Y%!.0 0 !!.! $!# ! Z! \ 2!! # !!> "!.!$ 2Q8' 0ZWWWKNNYHZYM>W ^W( . $ -2%(! !#!$= # "!# = !>32! >Z*ZWWWK\XNHZ_[ZY>\N N_w!. % B w!.( 'Z(*Z ! = !> "!.!$ " @##! = !>3 2! * \__YKZ_NH\^\N>^N NZ*!! v B *8 &Z_J = !>32!> ! Q !2!2 ##! *\__[K\[HMZY_>W N\ v ! ) - . ! !$ " Z 2! ! !; .! # !#! L8P>! = !>3 %ZM2!'\__ZK\_H^[\>XZ
78=
c2W.%'!c2 '2 )c2W.%'!c2 c" .&`.!2 "' #^ c"),
N^ + % !3 *2!&& = !> "!.!$L'ZP.!#! 2 !+!ZWWXKZZH\Z^>\[ NN + | ( 2# # = !> "!.!$'Z#@! !# ! ! ! !" .! *ZWW\K\[XHZNXWW>Y_^ NM% B w8 w!.*!!.! $!#.!! !. !2 " = !
!; .!.!$'\wL '\wP2\__XK^NHZY^>Y N[8!) *) + ! !# "!!@ ! * .!.! ! 0 >8 !.! ZWW[KNNHWX>Z_^ NX0 v* +! w!.* *>! = !>32!L8 &Z_P ! !! "! 2! 32!2! (\+
ZWWWKW[HN^Z^>Y NY
3% % 2& = !>32!8 &Z_# "! !; . !! 22 *\__ZK\X[H^M^^N>N^ NW ! )!. 8 w* #!#!>! = !>32! %ZM *\__MK\Y_H\X^M[>[M M_
3 % % = !>3 2! @ !. ! !#!>.! ! # >2' !ZWWWK\WHN_^_>[ MZ '8 " ( *! !!! #!" # = !# Q2 !! ! > ' !ZWWXK\XH\NXZ>X M\0 3 3
* .++! 3! 8 &Z_ 2. !! ! #"! !! .!\__YK\XH[_[Y>XN M^0*% ! *!. 'Q2 !#8 &Z_.! . 2. !2 ! !. ! !!.! . ! ! .!\__^K\\H\MW\>[_^ MNB!.) )!.B 0 Bv ! #.!Q2 !!2 " ! # !! !#! !.) B v !.) ) 0B L !.P\__^K^MHZ_M^>[_ MM 3! .8 .>- !( %## ( @! !#8B+^ @ ! !#0% 0 ^ !@3# !. &> 0 !" \__[KY[HX\N>^X M[' !8 0!. ! !# !!
2! # ! !!. # = !>2 ! *\__Y MX ) !$ (# = !>32!8 &Z_!222 !! 2\__[KZXHWW[>Z__N MY* 8* %!&)* $!#! #!..@# > Z>! 22 ! 2!82 *\__^K\XYHMW\_>Y MW! -! 0!. 8 &Z_2 !. !# * \__[K\YZHZZNZ^>\Z [_8$+ 3( %&! 2! # ! %!\__\KZZH^\NW>M[ [Z 22 -" 8 &Z_ = !>!2!! .!#2 .! *\__MK\MH^NY^>WZ [\" .>% 8 8! 8Z_ !2.! 3#"2!2 ! .>2 # .! 'Q2+\__YKYNHZ_\>Z\ [^w!.() !.-& 0*%2M^!.". Q2 !#8 &Z_ .! 2. ! " ! ! .!\__[K\MH\^ZY>\X [N(3!" *!.( 8!.!>! .@ # ! !""
> "!# 2.@>! .! *! &\__\KZH\N>X [M 22 % 2'((Y >Z0! @ = !>32! 8 &Z_!
.!*\__NK\XWHZ[M_^>Z_ [[ -! & - - 8 3 >-! & &.!. # '((Y ! !; . # 2 .! Z*\__ZK\X[HN[[MM>[_ [X
3% -" ) '& ! # Z0= # !!. !#
.!# = !>38 &Z_*\__[K\YZH\__NM>MN [Y
* *!0 &!.!3+\^!3 ( 2 = !J2 2@ ZWWYK^WZHXZM>Y [W |! 8 !((" 2 = !! !22 # = !> ! \__MKZ\HX_Y>ZN X_
3% -" % 2(.!#8 &Z_ \[ 2 !2!!# = ! ! Z08' 0\__WKMY^HMWZ>N XZ-"
3% % 2& = !>38 &Z_! @ ( *[! $ .. !2 ! !*
\__YKZ\ZHN_XW>YY X\@ ' ! ! ( *[! = # 2. .!# ... !!.!*\__MK\Y_HN_\Y\>W\ X^+! w "v ( *[ ! .!!!2" ! !!3 @! 2.! + \__XKNNXHYMW>[^
2)]
A \_ 34 71 77 78 91>95 2 3L +0P 205 208 210 221 .. 243 244 !"!.!!.# L 0&P 165 192 !2 >2!. 2QL +*P 44 47 64 101 107>113 ! 2 ! ! ! L'( >(P 103 !" 2! 231 22 63 64 92 103 117 125 126 165 178 180 208 209 223 242 222Z 3>7 10>13 46 58 61 121 222ZJ ^ 3>7 10>13 46 121 ! QL &P 205 208>210 Q!>^ 20 25 27 80 81 >! ! 35
B > !! 48 62>64 71 177 + 137 138 141 * Z 47 79 81 108 189>192
C *! 49>52 64 70 71 88 105 110 111 116>118 120 123 124 127 129 165 177 179 180 189 191 192 198 203 210 221 236 242
* !. 137 138 * Q22 vL*+vP 139>141 * !.! 64 170 177 180 181 242 *(^> 25 142 143 Cdc48 17>27 136 139 141 142 * NY> #Z>2N 2Q 22 23 136 141 142 Cell cycle 17 22 48 58 60 79 108>111 117 126 165 170 178 179 181 198 242 *.@ 57 165 170 178 *2! 17 19 23 27 124 137 138 * + 23 119 124 143 *0 224 *+W .! L* P 46 48>52 57>59 61>65 81 95 Coupled monoubiquitination 31 34>38 * !M 49 51 58 61>64 81 83 Cul1 42 43 45>49 58>61 63 90 124 Cullin 41>44 46>52 57>64 81 89 95 127 * !>% = !. L*0P 41>52 57>60 62>64 *! L*v0(P 70 71 75 76 77 83 91>95 * !2 170 171 181 185 * ! !
! ! . L*8&8M_YP 143 78>
789
c2W.%'!c2 '2 )c2W.%'!c2 c" .&`.!2 "' #^ c"),
D ( #2 2L(2P 152>156 (.! 1 17 18 19 21>23 25>27 34 41 42 58>60 62>64 69 70 80 83 88 89 91 94 95 100>102 107>113 116 118>126 128 136>143 149>152 154>156 158 162 163 174 175 177>179 184 186 192 195 198 200 202>205 208>210 221 238 239 243 244 Den1 51 52 59 61 (! L(*P 221 242 Deneddylation 46 48>52 57>65 81 (2 2!L+ 2 !; .!P 149 154>156 (Z 140 141 (! 20 140 141 DeSUMOylation 162 173 175>179 202 204 ( = ! L( P 20 21 24>27 31 38 61>63 69>83 89 91>96 105 112 122 126 154 155 185 196 230 Deubiquitination 36 38 59 69 70 74 77 83 91 126 ("2! 17 32 48 51 57 73 83 89 94 120 129 152 156 165 170 177 179>181 219 242 Doa10 139 141>143 ( .. 83 156 219 224
E E1 2>5 7 8 10>14 23 31 32 42 58 61 89 100 121 152 159 160 162 165 171 185 195 196 215>218 221 228 230 238>243 E2 2>5 8 11>14 20 23 31 32 36>38 42>48 51 52 58>61 89 91>94 96 100 101 103 104 107>109 111>113 120 121 125 142 152 159 160 162 171 185 187 190 191 195 196 203 205 215>217 222>224 228 230 235 238>241 243 '\ !; .!.!$ W 159
E3 2 3 14 21>23 26 27 31 32 35>38 42 47 48 50>52 58 60 61 70 74 77 79>81 89>96 100 101 105 108 109 111>113 121 138 142 143 152 159>164 171 177 185 187 192 195 196 203 215>217 221>225 228 230 233 235 238 239 241 243 'N " 23 27 'N 18 27 '! 31 34 35 107 239 '!2 > 2! .!L' (P 19 20 22>27 123 136 137 139>143 '2 ZM 34>38 ' ( 22 23 25 137 139>142 ERdj5 137 ' .!> 2! "!.2! L'8+P 216 217 'Q 224
F 8! !!L8 P 191
G %!! 2! 170 175>177 181 % 137 gp78 20 23 141>143
H '*M 228 230>234 '*[ 224 230 ! [L ( *[P 243 0 >8 238 241 0 >8; !! 2Z_L8 &Z_P 72 73 238>244 %>* 140>142 0>Z0 101 103 + 101 103 . !!L P 188 191 192 204 Z 20 23 123 138>143 ZJ(^ 138>142 >(. 2Q 139 140 34>36 Z 137 141
2)]
I + 92 .. ! 142 . ! 142 ! 2! 2 71 89 143 228 235 241>243 !! 38 71 88 89 94 102 ! !$ 217 219>221 229 230 Innate immunity 217 220 !#!2 2L8 P 220 228 !#!L8P 2 71 215>221 223 224 228>233 235 241 242 !#!> .!ZML %ZMP 2>4 72 73 75 215>225 228>235 239 241 !#!> .!L %P 2 215 217 229 !#!>L8>P 241 242 22 49 59 61 78 126 164 165 170>177 179 180 196 200 209 215>217 220 222>224 22 ! 2 36 41 43 58 61 63 69 71>73 107 152 154 159 173 185 238 22 ". 160 164
W ZJ2!!!$ L P 49 57 61>63 69>73 80>83 2! 18 25 69>71 80 81
K -[^ = ! ! 90>95 100 103 105 108 125 187 -\ 137 138 141 -! 3 > L- P 103 178 179 217 218 220 229
L 0 ! 137 138 0! ! = ! 2Q L0 *P 23 90 93 101>105 0!2 = ! 71 100>105
78<
0! = ! ! 70 73 93 100 101 104 105 0!3. 2 44 73 75 77 81 83 101 112 0 $! 5 7 34 35 75 89 95 107 117>119 121 125>127 149 161>165 170>174 191 198 217 238 241>243 0 ! 12 23 31>33 35 37 38 41 43>47 58 70 89 107 108 117 120 121 127 129 152 155 159>162 173 184>188 196>198 202 203 209 216 222 223 233 235 238 239
M !! 137 Mdm2 50 60 61 70 117>123 125>129 178 2 49 57 58 62>64 71 72 80 *L; 2 2QP
2 143 241 109 110 111 113 163 164 178 Mnl1 137 141 ! Z 137 Monoubiquitination 31 33>38 42 89 100 107 121 125 2 &+ 150>152 155 156
N N 35>38 224 Y 2>8 10 11 13 41 43 44 46>52 57>63 72 73 78 81 83 95 116 117 119 121 125 127>129 171 223 230 239 240 243 '((Y Z!.L Z0P 243 244 ! 46>50 52 57>61 63 121 127 8> 23 71 73 77 88>96 100>105 198 242 8> ! L'P 23 89>91 93 94 102>105 163 >. ! 137
250
c2W.%'!c2 '2 )c2W.%'!c2 c" .&`.!2 "' #^ c"),
O 0'2@ 24 26 ! 2! 116>118 124 >W 138 141 Otu1 19 24>26 78 "! L& P! 18 69 70 71 73 77 78 92 94 95 111
2\X-2 60 61 63 64 p53 50 58 64 70 116>129 161 163 178 179 242 p97 17 18 20 109 112 113 141 +3! 37 38 143 +3!> !! 2> 3 2L+> 2P 143 +.! 32 57 64 65 156 158 180 +M 140 141 +3 = !>32!L+ 2P 149 151>156 +#!. ! !.!L+* P 35 36 186>188 198 199 203 + 32!L+0P 174 180 192 199 201 202 205 208>210 221 + 1 10 20>23 26 41 42 45 57>59 62>64 70 73 79 80 83 88>90 101 102 107>109 112 113 116 118 120>123 128 136 137 139 141>143 149>156 165 175 186 195 198 200 203 205 208>210 239 242>244 +
# L+# P 150>156 +!...! 19 80
R Rad23 23 109 113 120 141 Rad52 191 192 204 206 QZ 42 44 46>48 50 58 60>62 90 124
2# 190 191 ! ! 2QL&*P 140 141 #2ZJ#2\ 143 200 204 206 207 %L! !.!@.!P!. 20 23 42 92 140 143 195 203 205 207 Z 203 206 8N 193 200 201 205 207>210 Rpn10 109 Rub1 57
S \[ 2 1 10 21>23 26 41 58 59 62>64 90 107>109 112 120 128 143 239 243
ZJ \ 3>7 9>13
*8L -+Z> !>8>QP 42>49 89 95 119 123 124 127
[Z! ! 136 137 140
.. " 17 19
#> = !! 34
.3Z 142
. Z 203 204
.! 46 48 49 57 81 95
.!! ! 17 19 26 31 38 42 48 58 88 101 158 170 174
QM 193 200 203>207
QY 193 200 203>207 Smt3 170 172 173 178 196 197 203 206 Spt23 19 23 24 26
\ 187 188 198 203 204
& &>Z 217 242
4>7 9>13 19 20 32 42 45 47>49 58 69 72 75>83 90 92 101 107 137 138 149 158 159 188 190 192 195 199 201 202 208 223 235 239>241
>\ 35
! ! # 159
! !.#L P 159 160 162 164 165 186>189 192 195 199>203 207>209
2)]
L = !>P 2 3 7 10 72 73 116 117 119 121 124 125 128 129 158>165 170>181 184>189 191>193 195>210 223 230 232 239>241
2 2 L !2P 160 164 170>176 179>181 185 195 196 Senp1 164 165 171>180 200 210 Senp2 164 165 171>179 Senp3 164 165 171>177 Senp5 164 165 171>178 Senp6 164 171>177 180 181 200 209 Senp7 164 171>174 178 200
159>162 165 200
251
Z_ 101 107 109>113 Z0 3>5 12 215>221 228 230>234 239 Ube2S 107 109>113 = ! L P 10 18 20 22 25 32>35 75 77 92 101 120 141 243 244 Z 3>13 121 239>241 [ 238>243 = !> !!.!L (P 19 31>38 75 77 90 93 96 108 112 190 191 239 = ! !!.2!N^L +N^P 216>220 224 229 232 = ! ! 21>27 43>46 49 69 70 72>75 79>81 88>96 100>105 107>113 120 121 125 136 141 187 191 198 210 223 T Ubiquitin conjugation 2 3 17 50 &*> 25 142 88>90 94 96 100 239 241 &'N 141 143 = ! "2!L *+P &8 3>5 240 215 & 3 5 240 = !*>! L * P &!( . L&(%P 188 61 62 69>71 78>80 189 Ubiquitin deconjugation 88 93 94 &8> 93>95 100 102>104 241 242 = !>#!\L #\P 18 & 8[ 78 91>95 20>26 &! ! ! L&0 P 35 36 187 = !>#!^L #^P 20>22 191 24>26 & 149>151 156 = !>#!2@L 8( & 49 51 58 64 70 71 79 90 2@P 23 24 92>95 111 116>118 120 122>124 = !! !#L P 10 126 142 143 175 178 180 186 18 25 32>37 75 77 80 190 191 188 192 221 222 229 242 = !. 17>20 22 23 26 & 22 58 64 70 71 79 27 41 42 45 49>51 70 77 93 95 116 117 122 126 142 143 79 89 100>102 109 118 120 175 192 122>129 138 140>143 153 185 &2Z!#! 215 228 241 186 189>191 193 195 199 200 &2\!!! L(\P 17 203>208 210 215 216 18 21 143 = !>3L 0P 2 151 186 195 215 224 238>241 243 = !>32!L 0P 1 2 4 5 U 7>11 13 14 37 41 46 72 73 75 *Z\ 4 6 11>13 46 58 121 127 77 92 101 118 120 140 141 149 Y 12 14 216 217 220 228 152 195 196 198 215 223 228 230>234 229 232 233 239 241 243
252
c2W.%'!c2 '2 )c2W.%'!c2 c" .&`.!2 "' #^ c"),
= !>3 2 2 ZL 2ZP 164 165 170>174 178 185 196 200 204>206 = !>3 2 2 \L 2\P 164 165 170>173 178 185 196 200 202 204>206 = !>. BL BP2! 18>20 22 25 141 = ! 2 2 L +P 58 59 61 62 69>71 73>77 92 93 WM 124 126 217 USP15 58 59 62 92 95 USP18 75 217 Z 140 141
V |!# ! 216 223 | 124 217 219>221 224 228>230 235 242
] B&+>^ 138 141
Y v W 138 140 141 v>2 35 36
