Horticultural Reviews, Volume 29: Wild Apple and Fruit Trees of Central Asia

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HORTICULTURAL REVIEWS Volume 29 WILD APPLE AND FRUIT TREES OF CENTRAL ASIA

edited by

Jules Janick Purdue University

John Wiley & Sons, Inc.

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HORTICULTURAL REVIEWS Volume 29 WILD APPLE AND FRUIT TREES OF CENTRAL ASIA

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Horticultural Reviews is sponsored by: American Society for Horticultural Science

Editorial Board, Volume 29 Philip L. Forsline Elizabeth E. Dickson Maxine Thompson Roger D. Way

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HORTICULTURAL REVIEWS Volume 29 WILD APPLE AND FRUIT TREES OF CENTRAL ASIA

edited by

Jules Janick Purdue University

John Wiley & Sons, Inc.

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This book is printed on acid-free paper. Copyright © 2003 by John Wiley & Sons, New York. All rights reserved. Published by John Wiley & Sons, Inc., Hoboken, New Jersey Published simultaneously in Canada No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted under Section 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of the Publisher, or authorization through payment of the appropriate per-copy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, (978) 750-8400, fax (978) 750-4470, or on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, (201) 748-6011, fax (201) 748-6008, e-mail: [email protected]. Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives or written sales materials. The advice and strategies contained herein may not be suitable for your situation. You should consult with a professional where appropriate. Neither the publisher nor author shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. For general information on our other products and services or for technical support, please contact our Customer Care Department within the United States at (800) 762-2974, outside the United States at (317) 572-3993 or fax (317) 572-4002.

Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Library of Congress Cataloging-in-Publication Data: ISBN: 0-471-21968-1 ISSN: 0163-7851 Printed in the United States of America 10

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Contents Contributors Dedication: Calvin R. Sperling

vii ix

Philip L. Forsline

1. Collection, Maintenance, Characterization, and Utilization of Wild Apples of Central Asia

1

Philip L. Forsline, Herb S. Aldwinckle, Elizabeth E. Dickson, James J. Luby, and Stan C. Hokanson I. II. III. IV. V. VI. VII. VIII.

Introduction Germplasm Acquisition Central Asian Collections Maintenance Distribution Characterization and Evalution Utilization Conclusion Literature Cited

2. The Wild Apple Tree of Kazakhstan

2 12 24 37 40 45 56 57 58

63

A. D. Dzhangaliev I. II. III. IV. V. VI.

Introduction Historical Review The Role of Wild Apple in the Vegetative Cover of Mountain Regions in Kazakhstan The Influence of Wild Apples on the Structure of the Environment Characteristics of Wild Apple Growth and Development Intraspecific Polymorphism of Wild Apple

65 69 83 121 156 207

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CONTENTS

VII. VIII. IX.

Utility and Biochemical Characterization of Wild Apple Fruit Preservation of Wild Apples Conclusion Literature Cited

3. The Wild Fruit and Nut Plants of Kazakhstan

246 272 280 285

305

A. D. Dzhangaliev, T. N. Salova, and P. M. Turekhanova I. II. III. IV. V. VI.

Introduction Pome Fruits Stone Fruits Small and Vine Fruits Other Fruits Nuts Literature Cited

308 311 324 332 349 362 370

Subject Index

373

Cumulative Subject Index

375

Cumulative Contributor Index

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Contributors Herb S. Aldwinckle, Department of Plant Pathology, Cornell University, Geneva, NY, 14456-0462 Elizabeth E. Dickson, Faculty of Environmental Design, University of Calgary, Calgary, AB, Canada, T2N 1N4 A. D. Dzhangaliev, Kazakhstan Academy of Science, Interbranch Laboratory for the Protection of Germplasm, Main Botanical Garden, Almaty, Republic of Kazakhstan Philip L. Forsline, U.S. Department of Agriculture, Agricultural Research Service, Plant Genetic Resources Unit, Cornell University, Geneva, NY, 144560462 Stan C. Hokanson, University of Minnesota, Department of Horticultural Science, St. Paul, MN, 55108 James J. Luby, University of Minnesota, Department of Horticultural Science, St. Paul, MN, 55108 T. N. Salova, Kazakhstan Academy of Science, Interbranch Laboratory for the Protection of Germplasm, Main Botanical Garden, Almaty, Republic of Kazakhstan P. M. Turekhanova, Kazakhstan Academy of Science, Interbranch Laboratory for the Protection of Germplasm, Main Botanical Garden, Almaty, Republic of Kazakhstan

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Calvin R. Sperling

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Dedication: Calvin R. Sperling This volume of Horticultural Reviews is dedicated to the memory of Dr. Calvin Sperling for his tireless efforts to contribute to the world’s preservation of apple and tree fruit germplasm in the Central Asian habitats. Dr. Sperling died of cancer on May 20, 1995, at the untimely age of 38, a deep loss felt by all scientists in the international germplasm community. Dr. Sperling was born February 7, 1957, in Farwell, Minnesota, the son of Clarence and Doris Sperling. His boyhood was spent on a dairy farm in Alexandria, Minnesota. As a child, he accompanied his father, the county weed and agriculture inspector, on trips to identify and eradicate harmful weeds. Calvin graduated from North Dakota State University in 1979 with a B.S. in botany, followed by an M.S. in 1981 and a Ph.D. in biology from Harvard University in 1987. He married Debra Gilmore, and their son, Carl, was born in 1992. Calvin’s involvement in botany and plant collecting began as an undergraduate student. He undertook a five-year floristic survey and plant collection (over 5000 specimens) of Minnesota flora. While he pursued advanced degrees, he made four trips of two to five months duration to South America (Colombia, Ecuador, Peru, Bolivia, and Brazil) to collect plant germplasm, to study hummingbird behavior, hummingbird pollinated plants, ethnobotany, floristic variation, cultivation, and the usage of the tuber crop, Ullucus tuberosum. From 1984 to 1986, while still a graduate student, Calvin began his career with the U.S. Department of Agriculture-ARS as a consultant to conduct research on the ecogeographic distribution of wheat and wild relatives in Israel and Turkey and to conduct herbarium studies in England and Scotland. After receiving his Ph.D., he accepted a position as botanist with the National Plant Germplasm Resources laboratory, USDA-ARS, Beltsville, Maryland, and was named Plant Exploration Officer. He coordinated, and at times led, the agency’s foreign and domestic trips. Using a genepool concept, Dr. Sperling developed a rational means for establishing priorities for the National Plant Germplasm System (NPGS), which funded his explorations. He took into account plant rarity, plant utility in breeding, and input from plant breeders and the NPGS germplasm committees, including over 40 specific crop germplasm committees. These priorities resulted in explorations to acquire genetic ix

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DEDICATION: CALVIN R. SPERLING

resources to fill in the gaps identified in the existing NPGS collections. His responsibilities included implementation and coordination of exploration for the NPGS, research on wild relatives and progenitors of cultivated plants, and research on in situ preservation of crop genetic resources. Dr. Sperling oversaw the management of nearly 100 successful explorations to 40 different countries in the years 1987 to 1995. These trips resulted in the addition of thousands of new accessions to the germplasm collections. Among these were exploration trips to Central Asia (Kazakhstan, Uzbekistan, and Tajikistan) where he collected progenitors of food legumes as well as cultivated and wild fruit crops. Dr. Sperling led the first expedition to collect Malus in Central Asia in September 1989. While in Kazakhstan, Dr. Sperling and the team of American scientists met Professor Aimak D. Dzhangaliev, who introduced them to the wild apple germplasm in the area near Alma Ata. He helped to arrange an October 1992 visit of Professor Dzhangaliev to the United States, a visit that provided the catalyst for three successful expeditions in 1993, 1995, and 1996. This volume of Horticultural Reviews in many ways is a culmination of his efforts to organize research on wild apple germplasm. Dr. Sperling’s efforts to incorporate botanical science and ethnobotany in the NPGS agricultural explorations have been widely recognized. He was selected by Fortune Magazine as one of its “25 most fascinating Business People of 1989.” In 1995, the U.S. Department of Agriculture presented him with the Frank Meyer Medal for Plant Genetic Resources. The Healing Forest Conservancy, in cooperation with Conservation International, awarded him the first Richard Evans Schultes Award for Contributions to Ethnobotany for his role in coordinating global plant explorations of the National Plant Germplasm System. He was also awarded the N. I. Vavilov Medal for Botany from the Vavilov Plant Breeding Institute in St. Petersburg, Russia. Calvin was an extremely dedicated scientist. His warm personality and modest demeanor endeared him to all who knew him. In his short life, he became one of the nation’s foremost ethnobotanists known for excellence in field research and his work to conserve biodiversity and improve crop plants worldwide. The legacy of Calvin Sperling will live on as the unique genetic material from his many projects is used by plant breeders to improve their crops. Philip L. Forsline U.S. Department of Agriculture-ARS Cornell University Geneva, New York

1 Collection, Maintenance, Characterization, and Utilization of Wild Apples of Central Asia Philip L. Forsline U.S. Department of Agriculture, Agricultural Research Service, Plant Genetic Resources Unit, Cornell University, Geneva, NY 14456-0462 Herb S. Aldwinckle Department of Plant Pathology, Cornell University, Geneva, NY 14456-0462 Elizabeth E. Dickson Faculty of Environmental Design, University of Calgary, Calgary, AB, Canada, T2N 1N4 James J. Luby University of Minnesota, Department of Horticultural Science, St. Paul, MN 55108 Stan C. Hokanson University of Minnesota, Department of Horticultural Science, St. Paul, MN 55108

I. INTRODUCTION II. GERMPLASM ACQUISITION III. CENTRAL ASIAN COLLECTIONS A. 1989 Collection B. 1993 Collection C. 1995 Collection 1. Geographic Sites in 1995 D. 1996 Collection 1. Geographic Sites in 1996 Horticultural Reviews, Volume 29, Edited by Jules Janick ISBN 0-471-21968-1 © 2003 John Wiley & Sons, Inc. 1

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IV. MAINTENANCE V. DISTRIBUTION VI. CHARACTERIZATION AND EVALUATION A. Core Collection and Main Collection B. Central Asian Collections 1. Disease and Pest Resistance 2. Environmental Stress Tolerance 3. Plant Stature 4. Molecular Genetic Diversity VII. UTILIZATION VIII. CONCLUSION LITERATURE CITED

I. INTRODUCTION Apple is the most ubiquitous and well-adapted species of temperate fruit crops. It is grown in high latitude regions of the world where temperatures may reach –40°C to high elevations in the tropics where two crops may be grown in a single year (Janick 1974). Apples are the fourth most important world fruit crop following all citrus types, grapes, and bananas. The apple, along with many of the important temperate fruit crops, belongs to the Rosaceae or rose family. Apple, pear, quince, medlar and a few other species have been classified into the subfamily, Pomoideae, the pome fruits, having fruits with two to five carpels enclosed in a fleshy covering. The genus Malus consists of about 27 wild species (Table 1.1). Most of the species intercross and, since self-incompatibility is common, seed obtained from a botanic garden are mostly interspecific or intercultivar hybrids. It is therefore difficult to be certain of the authenticity of some species names. Some taxon formerly listed as species (Way et al. 1990) are now classified as cultivated species (Table 1.1) because there is no record of their having wild origins (Li 1989; Li 1996). The cultivated apple is likely the result of interspecific hybridization and at present, the binomial Malus ×domestica Borkh. has generally been accepted as the appropriate scientific name, replacing the previously common usage of M. pumila (Korban and Skirvin 1984). Malus sieversii Lebed., a wild apple species native to Central Asia, has been recognized as a major progenitor of the domesticated apple, M. ×domestica (Way et al. 1990; Ponomarenko 1987 and 1992; Morgan and Richards 1993; Juniper et al. 1999). In ancient times, apple seeds and trees were

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Baccatae (Rehd.) Rehd.

Baccatus Jiang

Sorbomalus Zabel.

Sieversinae Langenf.

Malus Langenf.

Kansuenses (Rehd.) Rehd.

Sieboldiane (Rehd.)

Sikkimenses Jing

Hupehenses Langenf.

Series

Malus sections, series and primary species.

Sections

Table 1.1.

M. sieboldii (Regel) Rehd. var. sargenti (Rehd.) M. kansuensis (Batal.) Schneid. M. transitoria (Batal.) Schneid. M. toingoides (Rehd.) Hughes M. komarovii (Sarg.) Rehd. M. xiaojinensis Cheng et Jiang M. fusca (Raf.) Schneid.

M. baccata (L.) Borkh. var. mandshurica (Komorov.) Likh var. sachalinensis (Juz.) Ponom. var. himalaica (M.) Vass. M. hupehensis (Pampan.) Rehd. M. halliana (Anon.) Koehne M. sikkimensis (Wenzig) Koehne

M. sieversii (Lodeb.) Roem. subsp. kirghisorum (Al.) Ponom. form. niedzwetzkyana (Dieck) Langenf. M. orientalis subsp. montana (Uglitz) Likh. subsp. turkmenorum (Juz.) Langenf. M. sylvestris (L.) Mill. var. praecox (Pall.) Ponom.

Primary Species

(continued)

4 M. yunnanensis (French) Schneid. M. prattii (Hemsl.) Schneid. M. honanensis Rehd. M. ombrophilla Hand.-Mazz. M. florentina (Zuccagni) Schneid.

Yunnanenses Rehd.

M. trilobata (Poiret) Schneid.

Eriolobus (D.C.) Schneid.

Adapted from Way et al. (1990); Langenfelds (1991); Ponomarenko (1992); Li (1996); Li (pers. com.).

Note: Cultivated Malus species and Malus species hybrids (secondary species) include: M. ×arnoldiana (Rehd.) Sarg. (baccata × floribunda) M. asiatica Nakai M. ×atrosanguinea ((Spaeth) Schneid. (halliana × sieboldii) M. ×dawsoniana Rehd. (fusca × domestica) M. floribunda Siebold M. ×domestica Borkh. M. ×hartwigii Koehne (halliana × baccata) M. ×magdeburgensis Schoch. (spectabilis × domestica) M. ×micromalus Mak. (baccata × spectabilis) M. ×platycarpa Rehd. (cornonaria × domestica) M. prunifolia (Willd.) Borkh. M. pumila Miller M. ×purpurea (Barbier) Rehd. (neidzwetzkyana × atrosanguinea) M. ×robusta (Carr.) Rehd. (baccata × prunifolia) M. ×soulardii (Bailey) Brit. (ioensis × domestica) M. spectabilis (Ait.) Borkh. M. ×sublobata (Dipp.) Rehd. (prunifolia × sieboldii) M. ×zumi (Mats.) Rehd. (mandshurica × sieboldii)

M. doumeri (Bois.) Chev. M. melliana (Hand.-Mazz.) Rehd. M. tschonoskii (Maxim.) Schneid. M. laosensis Chev.

Docyniopsis Schneid.

Florentinae Rehd.

Primary Species

Series

M. ioensis (Wood.) Brit. M. coronaria (L.) Mill. M. angustifolia (Ait.) Michx.

(continued)

Chloromeles (Decne.) Rehd.

Sections

Table 1.1.

1. COLLECTION AND UTILIZATION OF WILD APPLES OF CENTRAL ASIA

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likely dispersed from Central Asia, east to China and west to Europe, via trade caravan routes popularly referred to as the “Silk Road” (Juniper et al. 1999). This flow of apple germplasm declined over the last few centuries as overland trade through the region decreased and ceased in the twentieth century as Central Asia was isolated for political reasons. In the 1920s, Vavilov (1930) traveled through Central Asia and reported that large wild stands of M. sieversii existed in specific localities and suggested the region as a center of origin for the domesticated apple. Dzhangaliev (1977), while confirming the contemporary existence of the wild apple forests, also noted that they were under pressure in some areas due to urbanization, agriculture, grazing, and wood harvesting. In the 1980s, the U.S. Department of Agriculture (USDA) National Plant Germplasm System recognized that M. sieversii was a critical species that lacked representation in its Malus collection at the Plant Genetic Resources Unit (PGRU) in Geneva, New York. The material was critical because present cultivars of the commercial apple had a narrow genetic base and most commercial production was based on very few cultivars (Kresovich et al. 1988; Morgan and Richards 1993; Noiton and Alspach 1996; Hokanson et al. 1998). Malus sieversii could be a valuable genetic resource for the domesticated apple potentially containing more genetic diversity for important horticultural and environmentally adapted traits (Korban 1986; Way et al. 1990; Janick et al. 1996). M. sieversii is diverse with the wild trees bearing a full range of forms, colors, and tastes. Recent collection trips to Central Asia (Section III) have verified that M. sieversii is very diverse and has all the qualities present in M. ×domestica (Forsline et al. 1994; Forsline 1995). The east/west trade routes that eventually became the “Silk Road” passed through this region on the way to China to the east and to the Middle East, past the Black Sea, to the west. Travelers on foot, camels, and horses likely began dispersing this germplasm as long ago as Neolithic times with routes being well established by the Bronze Age (Juniper et al. 1999). Ruminants such as deer native to the area and donkeys, mules, and horses used by humans along with humans themselves avidly ate these apples. No doubt the best were selected and this narrowed the genepool as it was dispersed. The seeds pass undamaged through alimentary canals. Thus seedlings would have been randomly established along the length of the trade routes and hybridization between previously isolated species then became possible. A number of species likely contributed to the genetic makeup of the domestic apple. Malus orientalis found in western sections of the trade

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routes in the Russian Caucasus as well as in Turkey does not have the diversity of fruit quality, but may have contributed other valuable traits such as later blooming, adaptation to a wider array of habitats, and capacity for longer storage of the apples. Others that may have been contributing parents include, Malus sylvestris the European crabapple bearing small astringent, greenish-yellow fruits, native to an area from Britain across Europe to the Balkans, and Malus baccata and some of its subspecies or natural hybrids (M. mandshurica, M. prunifolia, and M. asiatica) on the eastern side of the trade routes. Recent evidence has shown M. sylvestris is an unlikely contributor to the genetic makeup of the commercial apple (Wagner and Weeden 2000). However, it may have been involved in the background of cider-type apples selected in Spain, France, and Britain. Selected cultivars likely arose from random hybridizations and they were maintained by vegetative propagation, especially grafting which is a very ancient horticultural technology. B. E. Juniper (pers. comm.) reported that Oxford University scholars have found Babylonian cuneiform tablets dating to 2000 BCE depicting graftage. The Greeks knew about grafting and it was discussed in the writings of Theophrastus (487–287 BCE) as well as Roman agricultural writers such as Cato (234–149 BCE), Varro (116–27 BCE), Virgil (70–19 BCE), Pliny (12 BCE–70 CE), and Columella (first century CE). By the nineteenth century, England claimed over 2500 cultivars and many more were known at that time in Russian territories (Morgan and Richards 1993). In North America, settlers relied on apple trees originating from seeds of apples collected from early plantings established in the Tidewater region of the East Coast (Calhoun 1995). Seedling orchard establishment continued in North America well into the nineteenth century. As a result, higher levels of genetic diversity accumulated in North America than in Europe at the time when selecting and grafting existing cultivars was the norm. The potential for hybridization was almost infinite and North America became a “vast experimental station,” where selecting promising seedling apple cultivars was practiced on a large scale. From this vast grow-out, many of the world’s important cultivars of unknown parentage arose such as ‘Delicious’ (‘Hawkeye’), ‘Golden Delicious’, ‘McIntosh’, ‘Jonathan’, ‘Rome Beauty’, and ‘Northern Spy’. Many of these cultivars have been used to breed new apple cultivars such as ‘Cortland’, ‘Empire’, ‘Jonagold’, ‘Fuji’, ‘Gala’, and ‘Pink Lady’. The introduction of resistance to apple scab (pathogen, Venturia inaequalis (Cke.) Wint.) with the incorporation of the Vf gene from M. floribunda 821 indicated the potential of interspecific crosses to introduce

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new genes into commercial cultivars (Crosby et al. 1992). In addition, gene transformation is now able to insert genes from any source into existing high-quality cultivars. Some genetic transformants are now in the testing stage (Norelli and Aldwinckle 2000). It is from this history that we recognize the importance of establishing genebanks to preserve valuable germplasm that has been accumulated over many millennia by human activities. The Plant Genetic Resources Unit (PGRU) in Geneva, New York is part of the National Plant Germplasm System (NPGS) administered by the U.S. Department of Agriculture (USDA), Agricultural Research Service (ARS). The activities of the NPGS have been reviewed in a special volume of Plant Breeding Reviews (Janick 1989). Repositories for clonally propagated crops were not introduced to the system until the 1980s. An early assessment of the germplasm reserves of pome fruits (Lamb 1974) publicized the need for establishing clonal germplasm repositories. The PGRU is devoted to curating Malus, Prunus, and Vitis as well as many seed propagated vegetable crops. The Malus collection is the most extensive collection consisting of nearly 4000 accessions. Previous reports on the activities of PGRU have been published (Forsline 1987, 1988, 1992, 2000; Forsline and Way 1993) as well as the status of apple germplasm collections in the world (Way et al. 1990). The apple collection at PGRU was established in 1984 according to the clonal repository plan of the NPGS (Barton 1975; Brooks and Barton 1977). The majority of these accessions (2438) are clonally propagated and stored as duplicate orchard trees. Dormant buds of 2000 accessions are stored in a back-up collection in liquid nitrogen at the National Seed Storage Laboratory (NSSL) in Fort Collins, Colorado (Forsline et al. 1999). A core subset has been established including 206 clones (Forsline 1996) that is a test-array of the most genetically diverse accessions available for evaluation of specific genetic traits. Approximately 3000 accessions are distributed annually. Accession history, characterization, and evaluation are documented in the Germplasm Resources Information Network (GRIN) (USDA 2000). The core subset and 70 percent of the remainder of the collection has been characterized with 25 morphological descriptors. In addition to the clonal collection, since 1988, approximately 1500 accessions of wild Malus spp. from centers of origin throughout the world are preserved at PGRU as seed lots. Accessions of Malus sieversii have been collected in Central Asia (mostly from Kazakhstan) from 12 distinct habitats and 894 tree sources (distinct accessions or seed lots). Collaborative evaluation for disease resistance and horticultural and molecular characterization is being conducted on 25,000 of these seedlings in 24 worldwide laboratories. Information on the Malus

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collection and other commodities in the NPGS can be accessed on the Web at: http://www.ars-grin.gov/npgs/. The procedures for collection, conservation, evaluation, and documentation of Malus germplasm have been described (Forsline 2000). Until 1989, wild Malus germplasm from the Asian center of origin was unavailable (Dickson and Forsline 1994; Forsline 1995; Hokanson et al. 1997b). In 1989, policy changes in the former Soviet Union permitted U.S. scientists to establish collaborative efforts with Central Asian counterparts to conserve this germplasm. Subsequently, with funding through the USDA National Germplasm Resources Laboratory (NGRL) and effort from personnel at the NGRL, the PGRU, and cooperating scientists from other institutions, contacts with scientists and government officials in Central Asian countries were initiated and collaborative collection expeditions were undertaken in 1989, 1993, 1995, and 1996. The four expeditions were conducted with equal funding from two programs in the USDA/ARS including the NPGS and the International Research Programs (IRP). Personnel participating in each expedition are listed in Table 1.2. The late Dr. Calvin Sperling led the initial trip with Herb Aldwinckle and Elizabeth Dickson in 1989 to develop contacts and determine the availability of Malus germplasm. Philip Forsline led the first follow-up trip in 1993 after detailed planning in 1992 with our main host, Professor Aimak Dzhangaliev. In 1994, a Specific Cooperative Agreement (SCA) was developed through USDA/ARS/IRP as a four-year project for research on Wild Apple Germplasm, funding the cooperative research activities of our main cooperator and expedition host in Kazakhstan, Professor Aimak Dzhangaliev. This agreement is one of many that IRP has funded for research in Central Asia. Following the development of the SCA, a third trip in 1995 included expeditions to sites that had not previously been accessed. In the fourth and final expedition in 1996, some of the objectives of the SCA were fulfilled with the cooperator, Professor Dzhangaliev. Dr. Stan Hokanson implemented expansion of our cooperative work in Kazakhstan and participated in this final expedition. The expeditions focused mainly on Kazakhstan where the primary collaborator was Professor Aimak Dzhangaliev of the Academy of Sciences of the Republic of Kazakhstan whose laboratory had researched the variation in M. sieversii in Kazakhstan over several decades (Dzhangaliev 1977). These expeditions are documented in several publications (Dickson and Forsline 1994; Forsline et al. 1994; Forsline 1995, 2000; and Hokanson et al. 1997b) and in GRIN (USDA 2000) on the Web at: http://www.ars-grin.gov/ars/NoAtlantic/Geneva/kaz_trip .html.

Main host for all collections, Professor, Head of Interbranch Lab for Protection of Germplasm, Main Botanical Garden, Kazakhstan Natl. Acad. of Sci. Interpreter for expedition, student, Almaty Univ. Leader of 3 expeditions from USA, Curator, PGRU, USDA-ARS, Cornell Univ Interpreter for expedition, junior scientist, Kazakhstan Natl. Acad. Sci. Post-Doc, PGRU, USDA-ARS, Cornell Univ.; presently, Professor, Univ. of Minnesota Fruit Breeder, INFRUTEC Host in Karatau area, forestry officer of Boraldy forest Interpreter for expedition, Professor of Philology, Kazakh State Agrarian Univ.

Dzhangaliev, A.

Isetbaev, R. Issayev, M. K.

Human, T.

Hokanson, S.

Gowhar, M.

Dusembina, S Forsline, P. L.

Stellenbosch, South Africa Chimkent, Kazakhstan Almaty, Kazakhstan

St.Paul, MN, USA

Almaty, Kazakhstan

Almaty, Kazakhstan Geneva, NY, USA

Almaty, Kazakhstan

Urdzhar, Kazakhstan Bellville, South Africa Almaty, Kazakhstan Almaty, Kazakhstan Calgary, Al., Canada

Almaty, Kazakhstan Almaty, Kazakhstan Almaty, Kazakhstan Geneva, NY, USA Almaty, Kazakhstan

City/Country

(continued)

1995 1993, 1995 1996

1996

1989, 1993, 1995, 1996 1993 1993, 1995, 1996 1995

1996 1995, 1996 1996 1989 1993, 1995, 1996 1995, 1996 1995 1993 1995, 1996 1989, 1993, 1995, 1996

Expedition Dates

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Bekbaeva, B. Britz, G. Butenok, L. Chekalin, S. V. Dickson, E.

Minister for Kazakhstan Ministry of Agriculture Director, M.A. Aitkhozhin Inst., Kazakhstan Natl. Acad. of Sci. Professor, Dept. of Plant Pathology, Kazakhstan Natl. Acad. of Sci. Professor, Dept. of Plant Pathology, Cornell University Deputy Minister, Ministry of Ecology and Bioresources, Chair, Committee of Forestry Host, Semipalatinsk Region of the Tarbagatai Area Pome Fruit Development, UNIFRUCO LTD Botanical assistant of Professor Dzhangaliev Vice Director, Main Botanical Garden, Kazakhstan Natl. Acad. of Sci. L. H. Bailey Hortorium, Cornell University, presently U. of Calgary

Affiliation/Address

Participating personnel in four expeditions to Kazakhstan.

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Ahymbekov, S. S. Aitkhozhina, Nagima Aitkhozhina, Nazira Aldwinckle, H. S. Amanbayev, A. K.

Expedition Personnel

Table 1.2.

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10 Plant Pathologist, Washington State University Apple Breeder, Hort. Res. Inst. Senior officer of Sarkand Forest, main host in Djungarsky area Collector of ornamental species, Professor of Horticulture, Univ. of Minnesota Director, Main Botanical Garden, Kazakhstan Natl. Acad. of Sci. Wife of Professor Dzhangaliev, Pomologist (wild apricots), Main Botanical Garden Senior forest officer of the Boraldy forest

Mink, G. Noiton, D.

Ovchinnikov, I. D.

Pellet, H.

Serikbaev, S.

Rachimbaev, I. Salova, T.

Kuzubaev, N. H.

Kolbintsev, V.

Kunakbayev, Y.

Botanist for collection in the Karatau area, Main Botanical Garden Professor of Pomology and Plant Breeding, Univ. of Minnesota Forestry officer in the Almaty region, host for collections in Ketmen area Occasional interpreter, Science and Economic Assistant, Embassy, USA Assistant in Karatau area, Senior Science worker of Aksu-Jabagli Nature Preserve Senior officer of Sarkand Forest, Host in Djungarsky area

Affiliation/Address

Chimkent, Kazakhstan

Almaty, Kazakhstan Almaty, Kazakhstan

St. Paul, MN

Prosser, WA, USA Havelock North, New Zealand Sarkand, Kazakhstan

Sarkand, Kazakhstan

Chimkent, Kazakhstan

Almaty, Kazakhstan

Almaty, Kazakhstan St. Paul, MN Almaty, Kazakhstan

City/Country

1993 1989, 1993, 1995, 1996 1993, 1995, 1996

1993,1996 1993, 1995, 1996 1996

1993, 1995, 1996 1993

1993

1996

1993 1995 1996

Expedition Dates

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Ivashenko, A. A. Luby, J. J. Kashaganov, B.

(continued)

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Expedition Personnel

Table 1.2.

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Venglovsky, B.

Unruh, T. Urazaev, F. R. Ustemirov, K. J.

Host for all Karatau and Talasky areas, General Director, Chimkent region Research Entomologist, USDA, ARS, Yakima Agric. Res. Lab. Main guide in Tarbagatai area, forestry officer, Semipalatinsk region Travel arrangements to Chimkent region, Foreign Relations, Com. of Forestry Host for expedition in Kyrgyzstan, Professor, Inst. of Forestry and Walnut Breeding

Bishkek, Kyrgyzstan

Yakima, WA, USA Urdzhar, Kazakhstan Almaty, Kazakhstan

Chimkent, Kazakhstan

Almaty, Kazakhstan

Chimkent, Kazakhstan

Almaty, Kazakhstan Beltsville, MD, USA

Almaty, Kazakhstan

1989, 1993, 1995, 1996 1993, 1995, 1996 1996 1995, 1996 1993, 1995, 1996 1993

1996

1995, 1996 1989

1993

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Turganov, G.

Turehanova, R.

Cohost of 1993 expedition, Professor, Res. Inst. of Fruit and Viticulture Botanical assistant of Professor Dzhangaliev, Almaty, Kazakhstan Leader of 1989 expedition, Plant Exploration officer, USDA-ARS (deceased, May, 1995) Host for Karatau and Talasky areas, forestry officer for Chimkent region Botanist, assistant for all expeditions, Main Botanical Garden

8/8/02

Tulemisov, E.

Sotnikov, V. Sperling, C.

Smuryghin, A.

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P. FORSLINE, H. ALDWINCKLE, E. DICKSON, J. LUBY, AND S. HOKANSON

The expeditions successfully introduced large quantities of seeds as well as a limited number of clonal accessions to the ex situ collection at the PGRU. The collections aroused much interest among researchers in the United States, Canada, and several other countries. Upon request to the Germplasm Curator at the PGRU, scientists were provided with samples of the collections that they are currently evaluating for various traits at their respective home locales. Extensive evaluation is also being conducted at the PGRU in collaboration with Cornell University scientists. Thus, in less than a decade since the germplasm became available, a substantial international evaluation effort was mobilized rapidly and spontaneously. In this volume, we summarize progress in evaluation of the collections and prospects and plans for their utilization based on reports that were graciously provided to us by the cooperating researchers. II. GERMPLASM ACQUISITION Since 1984, 2438 Malus clones have been acquired at the PGRU. The majority were obtained early in the life of the apple repository from breeders’ collections throughout the United States. Most were cultivars of Malus ×domestica such as those of the National U.K. Repository described by Smith (1971). The U.S. apple repository was established at the New York Agricultural Experiment Station at Geneva, in part because of the large collection of Malus (Way 1976) that Cornell apple breeders at the station had collected during the previous century. Nearly 50 percent of the 2438 clones were originally acquired by breeders at Cornell University (Lamb 1974; Way 1976), although other Malus species were collected from botanical gardens as well as breeders’ collections. Significantly less than five representatives were available for each of 30 Malus species (Table 1.1). These species matched the published morphological descriptions (Yü 1979), but critical passport information including the country of origin, and information about the collectors and habitat of origin were generally lacking. The Malus in germplasm collections were considered genetically vulnerable (Way et al. 1990) because of the large gaps that existed in regard to wild apple germplasm. The first initiative to increase genetic diversity for wild germplasm was begun in 1987 with collection expeditions for four native North American Malus species (Dickson et al. 1991). Although North American germplasm has not thus far contributed to the cultivated genome of M. ×domestica, it will almost certainly have potential for future utilization. Until 1989, wild Malus germplasm from the Asian center of origin was unavailable (Dickson and Forsline 1994; Forsline 1995; Hokanson et al.

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1. COLLECTION AND UTILIZATION OF WILD APPLES OF CENTRAL ASIA

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1997b; Hokanson et al. 1999). The material was critical because present cultivars of the commercial apple have a very narrow genetic base (Morgan and Richards, 1993). This has subsequently been confirmed by genetic, biochemical, and molecular methods (Kresovich et al. 1988; Noiton and Alspach 1996; Hokanson et al. 1998) and assessment using biochemical and molecular tools confirmed this (Kresovich et al. 1988, 1990). The existence of wild stands of M. sieversii, the progenitor of cultivated apple, had been confirmed by Vavilov (1930) and Dzhangaliev (1977) in specific localities in Central Asia. These populations could be expected to contain genetic diversity for many important traits, such as resistance to biotic and abiotic stresses, fruit quality, tree attributes, and other genes that may solve as yet unforeseen problems (Korban 1986; Way et al. 1990; Janick et al. 1996). The 1500 accessions of wild apple collected since 1987 include mostly seed populations from the following regions: (1) 200 accessions of four North American species across a broad geographic range; (2) 102 accessions of seven Chinese species from Sichuan; (3) 86 accessions of M. orientalis from the Russian Caucasus and the mountains of Turkey south of the Black Sea; (4) approximately 100 accessions of other Malus species obtained from direct exchange with collaborators around the world; and (5) 892 accessions of M. sieversii collected in four different years from 12 areas (Fig. 1.1) in Central Asia. The Central Asian M. sieversii specimens are from Kyrgyzstan, Uzbekistan, Tajikistan, and Kazakhstan whose regions are characterized by mountain ranges (Zailisky, Djungarsky, Karatau, Ketmen, and Talasky) as depicted in Fig. 1.1 and described in Tables 1.4 and 1.5. Highlights of some of these collection activities (Dickson and Forsline 1994; Forsline 1995; Hokanson et al. 1997b) can be found on the Web at: http://www.ars-grin.gov/ars/NoAtlantic/Geneva/kaztrip.html. The Central Asian M. sieversii collections are primarily from Kazakhstan with additional samples from Kyrgyzstan, Uzbekistan, and Tajikistan. The species is a dominant overstory component in montane forests in these countries as depicted in Fig. 1.1 and described in Table 1.5. Most collections were made as seeds, and altogether over 120,000 seeds were collected from 894 trees (Tables 1.6 and 1.7). Many of the seeds were selected at random from trees at the collection sites, but a concentrated effort was made in the 1995 and 1996 expeditions to identify and collect from trees that appeared to possess horticulturally desirable characters. These trees have been termed “elites” (Plate 1). Fruit size of M. sieversii from the different regions in Kazakhstan was diverse, but some accessions had fruit with an average diameter greater than 60 mm, approaching the size of many commercial cultivars (Table 1.8). Areas 5, 9, 11, and 12 (Fig. 1.1) had the largest fruit that also closely resembled

Fig. 1.1.

Map of regions in Central Asia where Malus sieversii (Lebed.) was collected in 1989, 1993, 1995, and 1996.

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15

Descriptors used to characterize apple collections in Kazakhstan.

Category/ Descriptor No.a

Descriptor

Descriptor Defined

Code/Defined

Morphology/4

Calyx Basin (CALYXBASIN)

Appearance of Calyx Basin

Morphology/21

Fruit Bloom (FRUITBLOOM)

Rating of Natural Bloom (Wax) on Fruit at Maturity

Morphology/22

Fruit Flesh Color (FRTFLSHCOL)

Fruit Flesh Color

Morphology/23

Fruit Flesh Firmness (FRTFLSHFRM)

Fruit Flesh Firmness at Maturity

Morphology/24

Fruit Flesh Flavor (FRTFLSHFLA)

Fruit Flesh Flavor

Morphology/26

Fruit Ground Color (FRTGRNDCOL)

Fruit Ground Color at Maturity

1 2 3 4 5 6 7 1 2 3 4 1+1 2+2 2+3 2+4 3+3 3+4 4+4 4+5 4+7 5+5 6+6 7+7 8+8 1 2 3 4 1 2 3 4 5 1 2 2,4 3 4 5 5,4 6 7 8 9

none acute shallow acute medium acute deep obtuse shallow obtuse medium obtuse deep absent slight moderate heavy white cream cream + green cream + yellow green green + yellow yellow yellow + orange yellow + red orange pink red rose red soft semihard firm hard aromatic sweet subacid acid astringent light green green green yellow light yellow yellow orange orange yellow brown pink red purple (continued)

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(continued)

Category/ Descriptor No.a

Descriptor

Descriptor Defined

Morphology/28

Fruit Length (FRTLENGTH)

Morphology/34

Fruit over Color (FRTOVERCOL)

Fruit Length (Ave of 10 Fruits from Vigorous Tree) in mm. Fruit Over Color at Maturity

Morphology/35

Fruit Russet Intensity Percent of Fruit (FRTRUSSET) Surface with Russet (0–100%) Fruit Russet Loc. Location of Russet (FRTRUSSLOC) on Fruit

Morphology/36

Morphology/37

Fruit Russet Type (FRTRUSSTYP)

General Type of Russetting

Morphology/38

Fruit Shape (FRUITSHAPE)

Fruit Shape (Overall)

Morphology/41

Fruit Size Uniformity (FRTSIZUNIF) Fruit Stem Length (FRTSTEMLEN)

Consistency of Fruit Size Stem Length (Average of 5) in mm

Morphology/42

16

Code/Defined mm at maturity

1 none 2 green 2,5 green brown 3 yellow 3,2 yellow green 4 orange 5 brown 5,7 brown red 5,9 brown purple 6 pink 7 red 7,5 red brown 8 dark red 9 purple % of surface with russet (0–100%) 1 pedicel end only 2 calyx end only 3 both pedicel and calyx end 4 entire fruit 1 extremely fine 2 medium heavy 3 surface cracks 1.0 globose 1.1 globose-conical 1.2 short-globoseconical 2.0 flat 2.1 flat-globose (oblate) 3.0 conical 3.1 long-conical 3.2 intermediateconical 4.0 ellipsoid 4.1 ellipsoid-conical 5.0 oblong 5.1 oblong-conical 5.2 oblong-waisted 1 uniform 2 variable mm at maturity

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Morphology/43

Fruit Stem Thickness (FRTSTEMTHK)

Rating of Stem Thickness

Morphology/44

Fruit Texture (FRTTEXTURE)

Morphology/45

Fruit Width (FRUITWIDTH)

Morphology/60

Overcolor Intensity (OVERCOLOR) Stem Cavity (STEMCAVITY)

A Rating of Fruit Flesh Texture at Maturity Fruit Width (Ave of 10 Fruits from Vigorous Tree) in mm Percent of Overcolor on Fruit 0–100% Appearance of Calyx Basin

Morphology/91

Tenacity of Fruit (FRUITTENAC)

Tendency of Fruit to Abscise or Hang

Phenology/4

Harvest Season (HARVSEASON)

Harvest Season (days before or after Red Delicious— code no. 6)

Production/21

Tree Bearing Habit (TREEBEAR)

Tree Bearing Habit

Growth/1

Tree Vigor (TREEVIGOR)

Tree Vigor from Small to Large When Grown on a Vigorous Rootsock

Morphology/82

1 2 3 1 2 3

slender medium stout fine medium coarse mm at maturity

% overcolor on fruit at maturity 1 none 2 acute shallow 3 acute medium 4 acute deep 5 obtuse shallow 6 obtuse deep 1 drops before mature 2 holds past maturity 3 persists into winter 1 >60 extremely early 2 50–60 very early 3 30–50 early 4 20–30 medium early 5 10 medium 6 RD season med. late 7 10 late 8 >30 extremely late 1 columnar 2 Type I spur 3 Type II semispur 4 Type III standard 5 Type IV tipbearer 6 weeping 1 small 2 medium 3 vigorous 4 very vigorous

a

As listed on the Web at: http://www.ars-grin.gov/cgi-bin/npgs/html/desclist.pl?115. 17

18 Gazni Kondra Local Lake Tashkent Aksay River Turgen, Kamenka Almatinka River Talgar Kuznetzov Kotur Bulak Issyk Botanical Garden Lepsinsk Konstantinovka #1 Konstantinovka #2 Topelevka, Low el. Topelevka, High el. Boraldy, Low el. Boraldy, High el. Kokbulak Fergansky #1 Fergansky #2 Alekseyevka site 05 Alekseyevka site 07 Alekseyevka site 09 Alekseyevka site 06 Alekseyevka site 08 Alekseyevka site 10 Bolshoye Aksu Aksu Jabagli

Tajikistan/—/1.00 Uzbekistan/—/2.00 Uzbekistan/—/2.01 Uzbekistan/—/2.02 Kazakhstan/Zailisky/3.00 Kazakhstan/Zailisky/3.01 Kazakhstan/Zailisky/3.02 Kazakhstan/Zailisky/3.03 Kazakhstan/Zailisky/3.04 Kazakhstan/Zailisky/3.05 Kazakhstan/Zailisky/8.00 Kazakhstan/Djungarsky/4.00 Kazakhstan/Djungarsky/4.01 Kazakhstan/Djungarsky/4.02 Kazakhstan/Djungarsky/5.00 Kazakhstan/Djungarsky/5.01 Kazakhstan/Karatau/6.00 Kazakhstan/Karatau/6.01 Kazakhstan/Karatau/11.00 Kyrgyzstan/—/7.00 Kyrgyzstan/—/7.01 Kazakhstan/Tarbagatai/9.00 Kazakhstan/Tarbagatai/9.01 Kazakhstan/Tarbagatai/9.02 Kazakhstan/Tarbagatai/9.03 Kazakhstan/Tarbagatai/9.04 Kazakhstan/Tarbagatai/9.05 Kazakhstan/Ketmen/10.00 Kazakkstan/Talasky/12.00

1490–1860 1100–1630 1100–1630 1100–1630 1260–1490 1200–1360 1440 1490 1550 1680 — 1110–1360 1260 1185–1260 1180–1460 1500–1760 600–620 910 780–1230 1335 1550 800–870 820–960 1030–1120 860–990 800–920 860–1100 1600–1660 1010–1025

Elevation (m)

1989 1989 1989 1989 1995, 1996 1993, 1995 1995 1995 1989, 1995 1993 1993 1993, 1995, 1996 1996 1996 1993, 1995, 1996 1995, 1996 1993, 1995 1995 1996 1993 1993 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1996 1996

Years collected

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As depicted in Fig. 1.1.

— — — — 43°07'N /77°47'E. 43°19'N /77°35'E. 43°06'N /76°54'E. 43°17'N /77°23'E. 43°21'N /77°40'E. 43°12'N /77°40'E. — 45°31'N /80°43E. 45°38'N /80°55'E. 45°41'N /80°52'E. 45°24'N /80°24'E. 45°24'N /80°25'E. 42°52'N /69°53'E. 42°52'N /69°53'E. 42°40'N /70°16'E. 41°23'N /73°06'E. 41°16'N /72°53'E. 47°14'N /81°34'E. 47°16'N /81°35'E. 47°16'N /81°34'E. 47°16'N /81°34'E. 47°15'N /81°34'E. 47°15'N /81°35'E. 43°18'N /79°31'E. 42°19'N /70°22'E.

Latitude/Longitude

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a

Site Name

Country/Mt. Range/Area Sitea

Table 1.4. Site descriptions for collection regions and years sites were accessed in Central Asia.

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1. COLLECTION AND UTILIZATION OF WILD APPLES OF CENTRAL ASIA Table 1.5.

19

Site descriptions for collection regions in Central Asia.

Country/Mt. Range/ Area Sitea Tajikistan/—/1.00 Uzbekistan/—/ 2.00-202 Kazakhstan/Zailisky/ 3.00–3.05 Kazakhstan/Djungarsky/ 4.00–4.02 Kazakhstan/Djungarsky/ 5.00 Kazakhstan/Djungarsky/ 5.01

Kazakhstan/Karatau/ 6.00 Kazakhstan/Karatau/ 6.01 Kazakhstan/Karatau/ 11.00 Kazakhstan/Tarbagatai/ 9.00–9.05 Kazakhstan/Ketmen/ 10.00 Kazakhstan/Talasky/ 12.00

Forest Type Xeric, mixed scrub forest Xeric, mixed scrub forest Humid-temperate, mixed forest Humid-temperate, mixed forest Humid-temperate, mixed forest Humid-temperate, mixed forest with diverse flora at high elevation Diverse stream habitat in xeric area Xeric, mixed scrub forest Xeric area and stream habitat Dry continental forest with –40°C common Semi-dry, temperate mixed forest Dry N. slope of canyon

Elevation (m)

Annual Precipitation (mm)

—

300

—

300

1170–1690

700

1190–1360

800

1170–1490

850

1500–1760

850

600–620

250

880–910 780–1230

250 250

870–1120

450

1600–1700

650

1000–1025

320

a

As depicted in Fig 1.1.

Table 1.6.

Total M. sieversii seeds collected, distributed, and stored.

Year

No. Accessions

No. Seeds

No. Orders Processed

No. Distributed

No. at NSSL

No. at PGRU

1989 1993 1995 1996

119 60 408 307

9,563 23,406 60,081 28,039

33 147 443 279

1,022 8,486 14,713 6,454

3,494 7,290 19,970 8,344

5,047 7,630 25,398 13,241

Total

894

121,089

902

30,675

39,098

51,316

No. Accession (trees harv). 19 18 8 20 53 35 1 2 57 3 3 76 10 21 117 51

Country/Mt. Range/Area Sitea

Tajikistan/—/1.00 Uzbekistan/—/2.00 Uzbekistan/—/2.01 Uzbekistan/—/2.01 Kazakhstan/Zailisky/3.00 Kazakhstan/Zailisky/3.01 Kazakhstan/Zailisky/3.02 Kazakhstan/Zailisky/3.03 Kazakhstan/Zailisky/3.04 Kazakhstan/Zailisky/3.05 Kazakhstan/Zailisky/8.00 Kazakhstan/Djungarsky/4.00

Kazakhstan/Djungarsky/4.01 Kazakhstan/Djungarsky/4.02 Kazakhstan/Djungarsky/5.00

Kazakhstan/Djungarsky/5.01

6,151

474 1,997 21,021

1,267 1,055 385 1,100 4,449 3,456 374 2,406 7,323 949 3,125 11,397

No. Collected

989

35 474 5,254

108 244 24 82 681 1,042 50 230 834 643 363 1,620

No. Distributed

2,176

155 679 7,027

449 259 130 404 1,525 992 160 1,100 2,912 125 1,350 4,343

No. at NSSL

2,986

284 844 8,740

710 552 231 614 2,243 1,422 164 1076 3,577 181 1,412 5,434

No. at PGRU

1989 1989 1989 1989 1995, 1996 1993, 1995 1995 1995 1989, 1995 1993 1993 1993, 1995, 1996 1996 1996 1993, 1995, 1996 1993, 1995, 1996

Years Collected

20

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Total collections, distribution, and storage of M. sieversii seeds from Central Asia.

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Table 1.7.

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P. FORSLINE, H. ALDWINCKLE, E. DICKSON, J. LUBY, AND S. HOKANSON

As depicted in Fig. 1.1.

a

121,809

5,705 8,620 6,102 3,862 1,883 4,656 4,411 4,482 2,973 1,878 6,440 2,118 1,050 30,675

1,348 3,437 1,195 997 861 1,506 1,255 1,448 1,069 1,025 3,143 513 205 39,098

1,756 2,337 1,949 1,450 535 1,339 1,076 1,177 860 400 1,543 613 277 51,316

2,601 2,846 2,958 1,415 487 1,811 2,080 1,857 1,044 433 1,754 992 568 1989, 1993, 1995, 1996

1993, 1995 1993, 1995 1996 1993 1993 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1995, 1996 1996 1996

4:19 PM

894

45 46 53 6 6 45 60 49 15 7 33 22 13

8/8/02

Total

Kazakhstan/Karatau/6.00 Kazakhstan/Karatau/6.01 Kazakhstan/Karatau/11.00 Kyrgyzstan/—/7.00 Kyrgyzstan/—/7.01 Kazakhstan/Tarbagatai/9.00 Kazakhstan/Tarbagatai/9.01 Kazakhstan/Tarbagatai/9.02 Kazakhstan/Tarbagatai/9.03 Kazakhstan/Tarbagatai/9.04 Kazakhstan/Tarbagatai/9.05 Kazakhstan/Ketmen/10.00 Kazakkstan/Talasky/12.00

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the cultivated apple in other quality traits. Likewise, some trees were nearly disease-free and had other desirable horticultural characteristics. Accessions were described with 25 priority descriptors (Table 1.3) at collection time. These records can be accessed on the Germplasm Resources Information Network (GRIN) on the Web at: http://www.arsgrin.gov/cgi-bin/npgs/html/search.pl (USDA 2000). For cataloging purposes, all collections received a sequential Geneva Malus (GMAL) number in order to distinguish our apple collection from others in the NPGS. Additionally, all permanent accessions receive a Plant Introduction (PI) number that can be used as well to search the GRIN database on the Web at http://www.ars-grin.gov/npgs/acc/acc_queries.html: 1989 collection 1993 collection 1995 collection 1996 collection

PI 600302–600409 (GMAL 3242–3360) PI 600423–600482 (GMAL 3526–3585) PI 600485–600584 (GMAL 3596–4003) PI 600585–600624 (GMAL 4010–4317)

We also maintain inventory, accession, and passport records for each accession. For named cultivars, pedigree, developer, and general descriptive narrative are recorded. For collections of wild material, site records, including GPS data are recorded along with observations of each accession collected in situ. The software used in the local data management is PARADOX by Borland International. Paradox Version 3.5 is a relational database management program that can be used either as a stand-alone system on a single computer or as a multi-user system on a network. This program is menu driven. Forms, reports, queries, and graphs are some of the capaTable 1.8. Mean range for fruit size in samples from random or elite Malus sieversii trees measured during collection in Kazakhstan. Fruit size mean (range) in mm Areaa 3 4 5 5 6 9 10 11 12 a

Year(s)

Elite Samples

Random Samples

1995/1996 1995/1996 1995 1996 1995 1995/1996 1996 1996 1996

45 (32–56) 44 (32–55) 46 (37–56) 58 (54–65) 42 (33–46) 56 (46–72) 49 (47–51) 55 (44–76) 65 (60–74)

34 (25–49) 34 (28–44) 36 (27–48) 35 (25–49) 41 (28–54) 43 (28–62) 40 (29–51) 42 (29–63) 42 (32–50)

See Fig. 1.1 and Table 1.2 for location and description of collection areas.

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bilities of the program, as well as multi-table links using forms, reports, queries, or graphs. All data are loaded to the NPGS database management system, Germplasm Resources Information Network (GRIN), which uses Oracle and SQL programs. Information about GRIN can be found on the Web at: http://www.ars-grin.gov/npgs/. The fruit size and quality varied each year in the sites that were visited. For example, in region 5, the size and quality of the fruit was much better in 1996 than in 1995 and fruiting was observed on a much higher percentage of the trees (Table 1.8). In addition to seed from 146 elite genotypes, vegetative material was collected from 44 of the elite accessions in the 1995 and 1996 expeditions and placed in quarantine at the USDA Plant Quarantine Facility in Beltsville, Maryland. Fruit color and fruit quality traits (apple flesh firmness and flavor) are highlighted in Tables 1.9 through 1.12 for collections made in 1995 and 1996. In 1995 and 1996, material was also collected specifically for molecular studies of population genetics and biodiversity to follow preliminary work by Lamboy et al. (1996). These included randomly selected Table 1.9. Red color of M. sieversii fruit from random populations collected from Kazakhstan. Red color

Mt. Range/ Area Sitea Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00 Total a

Total Accessions

No. Accessions with Red Colorb

Percent Accessions with Red Colorb

Percent Red Intensity of Those with Red Colorc

80

29

36

35

87

42

48

27

114

74

64

42

64

34

53

37

149

101

68

46

20 40 10

13 33 8

65 83 80

38 44 51

564

334

59

40

As depicted in Fig. 1.1. Code 4–9 in Morphology descriptor 34; fruit over color (Table 1.3). c Morphology descriptor 60; percent over color intensity (Table 1.3). b

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Table 1.10. Red color of M. sieversii fruit from elite selections collected from Kazakhstan. Red color

Mt. Range/ Area Sitea

Total Accessions

No. Accessions with Red Colorb

Percent Accessions with Red Colorb

Percent Red Intensity of Those with Red Colorc

11

10

91

63

14

10

71

41

Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00

24

19

79

46

21

15

68

45

58

55

91

60

2 13 3

1 8 3

50 62 100

50 44 44

Total

146

121

83

48

a

As depicted in Fig. 1.1. Code 4–9 in Morphology descriptor 34; fruit over color (Table 1.3). c Morphology descriptor 60; percent over color intensity (Table 1.3). b

seeds from 304 trees across areas 3, 4, 5, 6, and 9 in 1995 (Fig. 1.1) and from 261 trees across areas 3, 4, 5, 9, 10, 11, and 12 (Fig. 1.1) in 1996. Leaves from the 261 maternal trees in 1996 also were collected and dried in silica gel, with DNA extracted later at PGRU. III. CENTRAL ASIAN COLLECTIONS Collections made in four expeditions (1989, 1993, 1995, and 1996) produced 121,089 seeds from 894 individual trees (Table 1.6). Localities of all collections are also listed in Table 1.4. A summary of total collections made in each year is listed in Table 1.6 and a summary of collections made at each site is listed in Table 1.7. A Global Positioning System device was available for collections made in 1993, 1995, and 1996 permitting latitude and longitude to be associated with each site (Table 1.4).

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Table 1.11. Quality ratings of randomly collected fruit from Kazakhstan based on firmness and flavor. Distribution (%) Firmnessb

Flavorc

Mt. Range/ Area Sitea

Total Accessions

1

2

3

4

1

2

3

4

5

Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/ 10.00 Karatau/ 11.00 Talasky/ 12.00

80

39

49

10

3

10

3

48

14

25

87

26

60

14

0

4

5

31

38

23

114

21

45

29

5

7

11

32

23

28

64

22

50

28

0

34

22

33

6

3

149

23

42

34

1

10

5

30

9

36

20

20

45

35

0

0

20

25

40

15

40

20

35

40

5

35

20

15

15

15

10

20

50

30

0

0

0

50

30

20

564

25

47

26

2

15

9

33

18

25

Total a

As depicted in Fig. 1.1. Morphology descriptor 23; fruit flesh firmness (Table 1.3). c Morphology descriptor 24; fruit flesh flavor (Table 1.3). b

A. 1989 Collection The first trip to Central Asia included Calvin Sperling (leader) accompanied by Herb Aldwinckle and Elizabeth Dickson. Gratitude must be expressed to the opening of botanical exchange between the USSR and the United States (Tom Elias, formerly of Rancho Santa Ana Botanical Garden, Claremont, California, and Igor Smirnov, Moscow Botanical Garden). This trip took place between August 29 and September 14 in Tajikistan, Uzbekistan, and Kazakhstan identified as areas 1–3 (Fig. 1.1). In 1989, 119 seedlots (9563 seeds) were collected (Table 1.6), including 19 seedlots (1267 seeds from Tajikistan and 46 seedlots (2540 seeds) from Uzbekistan (Table 1.7). While in Kazakhstan, the collectors met Professor Aimak Dzhangaliev who is the author and coauthor of two works of this present volume. He provided access to Kazakhstan/

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Table 1.12. Quality ratings of elite fruit collections from Kazakhstan based on firmness and flavor. Distribution (%) Firmnessb Mt. Range/ Area Sitea Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00 Total

Flavorc

Total Accessions

1

2

3

4

1

2

3

4

5

11

27

46

9

18

36

0

55

9

0

11

7

21

72

0

21

7

57

14

0

24

8

25

58

8

38

8

46

4

0

21

0

19

48

33

43

5

38

10

5

58

7

26

55

12

79

7

10

2

2

2 13 3

0 8 0

50 23 0

50 54 0

0 15 100

0 69 67

0 0 0

100 31 33

0 0 0

0 0 0

146

8

25

51

16

56

5

32

5

2

a

As depicted in Fig. 1.1. Morphology descriptor 23; fruit flesh firmness (Table 1.3). c Morphology descriptor 24; fruit flesh flavor (Table 1.3). b

Zailisky/3.04 where 54 of the 57 seedlots (5756 of 7323 seeds) were collected in 1989. The remainder of the seedlots/seeds were collected in that same site in 1995 (Table 1.7). Fruit characterization records for collections in 1989 were not made, but the collectors observed that they ranged in size from small crabapples to commercial size, with color and quality being described as very diverse. Records and specific site data can be found on the Web. The data on the 1989 collection can be found by accessing catalog numbers PI 600302–600409 (GMAL 3242–3360). In October 1992, Professor Dzhangaliev traveled to PGRU to meet members of the U.S. Apple Crop Germplasm Committee at their annual meeting. Cooperative agreements with Professor Dzhangaliev and the Kazakhstan Academy of Sciences were developed whereby three additional collection trips to diverse areas of Kazakhstan were planned.

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B. 1993 Collection A second expedition to Kazakhstan took place September 6–28, 1993. Philip Forsline led a team of collectors including Elizabeth Dickson, Gaylord Mink, and Dominique Noiton. Professor Dzhangaliev was again the main host and was assisted by others as listed in Table 1.2. Despite a severe spring frost in many of the forested areas of Kazakhstan and Kyrgyzstan, we were able to collect 60 seedlots (23,406) seeds of M. sieversii (Table 1.6) from the eight area/sites (Table 1.4 and Fig. 1.1). Fruit characterization records for these collections were recorded using a narrative approach. These records, along with specific site data can be found on the Web at: http://www.ars-grin.gov/cgi-bin/npgs/html/ search.pl. The data on the 1993 collection can be found by accessing catalog numbers PI 600423–600482 (GMAL 3526–3585).

C. 1995 Collection The material collected in 1989 and 1993 provided clues to enable future collection trips to be more effective (Hokanson et al. 1997b, 1999). A study using isozyme analysis (Lamboy et al. 1996) to determine levels of genetic variation present in the 1989 and 1993 collections of M. sieversii was conducted. Based on genetic analysis of sib families from four areas, the populations of M. sieversii surveyed appear to constitute a single panmictic population with more than 85 percent of the total genetic variation due to differences among families and only 15 percent due to differences among regions. Thus we determined that the most efficient strategy to acquire generalized genetic diversity and potentially useful alleles would be to explore as many unique ecological niches as possible. This third expedition to Kazakhstan took place August 23 to September 16, 1995. Philip Forsline led a team of collectors including Elizabeth Dickson, James Luby, and two scientists from South Africa, Gary Britz representing that country’s apple industry, and Taaibos Human, fruit breeder. Again, Professor Dzhangaliev was the main host assisted by others. The main objectives of this expedition were to: (1) collect germplasm of Malus sieversii in its center of diversity, supplementing collections made in 1989 and 1993 by broadening the range of collections and returning to areas that had sparse fruiting in 1993; (2) collect other crop species as found in association with apple; and (3) expand contacts with Kazakh scientists to develop plans for further ex situ collections and develop strategies for in situ conservation.

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This third collection trip for M. sieversii was the most successful expedition based on the diversity and quantity of material and characterization data collected. Previous collections made in 1993 were limited due to low fruiting from that season’s spring frost. The 1995 collection was also timely since the social/political situation in Kazakhstan was deteriorating rapidly and future trips were jeopardized. Collections were made in four of the same areas as in 1993, but also included the northernmost site for M. sieversii (47°, 16′N) where wild apple trees were found with the largest size and highest quality fruit of any site explored thus far. Some elite selections were nearly 70 mm in diameter. Strategy for collection of apple in the wild forests was twofold: (1) random population sampling of 5 to 8 fruit/tree (average of 45 seed) from each of 30 trees in each site which appeared to be a unique ecosystem; and (2) selection of elite wild types (seed and occasionally scions) within these sites. Ten populations of 30 trees (5–8 fruit/tree) each were sampled which yielded 13,842 seeds. Additionally, 101 elite selections were made across all sites yielding 46,239 seeds and scions from 14 of the most elite types for a total of 60,081 seeds from 408 accessions (Table 1.6). In addition to the high-quality collections in the northern area, collections were made in a xerophytic area at 42°, 52′N (Karatau region) where trees were adapted to drought (Tables 1.4 and 1.5). Fruit at this site had excellent horticultural characteristics including the capacity for fruit to hang on the tree past maturity in an area with high heat units indicating potential for adaptation to areas with longer growing seasons. All collections were documented by 24 morphological descriptors (Table 1.3) along with associated site information. Those descriptors and codes for each can be found for the purpose of querying individual accessions on the Web at: http://www.ars-grin.gov/cgiin/npgs/html/desclist.pl?115. 1. Geographic Sites in 1995 1. Zailisky, Sites 3.00, 3.01–3.04. Day trips were taken August 25–27 to the five sites (Table 1.4) near Almaty, an area with extensive human encroachment that has led to a decline in native habitats. Some of these were the same areas observed in 1989 and 1993. Two populations of 30 accessions each (GMAL 3700–3729 from site 3.00 and GMAL 3730–3759 from site 3.01) were collected in this area. From these 60 accessions, 2121 seeds were collected. The mean diameter of apples found in these random populations was 34 mm (Table 1.8). In addition, 11 wild elite samples selections from sites 3.00 (GMAL 3596–3598), 3.01 (GMAL 3602–3603), 3.02 (GMAL 3599), 3.03 (GMAL 3600–3601) and 3.04

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(GMAL 3697–3699) averaged 46 mm (Table 1.8) from which 7878 seeds were collected. Complete data sets on all collections (populations and elites) with site and accession records can be accessed on the Web at: http://www.ars-grin.gov/cgi-bin/npgs/html/search.pl. We were able to observe extensive diversity in each site throughout the expedition with 60 to 90 percent of the trees in the forests fruiting. Forests in the Almaty area, Topelevka, and Lepsinsk areas (areas 3, 4, 5) were very similar in habitat and fruit types. Fruit was not of high quality, and few elite samples were collected in these general areas. Fruit quality traits for all collections made in 1995–1996 are listed in Tables 1.9–1.12. Tarbagatai, Sites 9.00–9.05. The second phase of our 1995 expeditions took us to the Tarbagatai mountain range (Fig. 1.1) which had not been visited in previous collection trips. Six sites (9.00–9.05) were examined from August 29 to September 1. An area of 3.5 × 3.0 km was covered, but different valleys and slopes were explored. Grazing has destroyed a large portion of this habitat. We discovered high-quality fruits (Plate 1A) in all sites within this area. Three populations of 30 accessions each: (1) GMAL 3760–3789 from site 9.00; GMAL 3790–3819 from site 9.01; and GMAL 3820–3849 from site 9.02 (Plate 2A) were collected in this area. The mean diameter of all fruits in the random populations (90 trees) was 43 mm (Table 1.8) of which 3630 seeds were collected. In addition, 14,414 seeds of 46 wild elites were collected including scions of the very best from sites: 9.00 (GMAL 3604–3607 with scions from GMAL 3607); 9.01 (GMAL 3616–3619 with scions from GMAL 3616 and GMAL 3619); 9.02 (GMAL 3626–3632); 9.03 (GMAL 3608–3615 with scions from GMALs 3608 and 3614); 9.04 (GMAL 3620–3625 with scions from GMALs 3623 and 3625); and 9.05 (GMAL 3633–3649 with scions from GMALs 3636, 3637, and 3643). The mean diameter of the wild elite selections was 56 mm (Table 1.8). The quality of some of these elite specimens was near to that of commercial apples. Therefore scions were collected for 10 of the 46 selections. The tree with the largest apples among these elites had fruits that averaged 72 mm in diameter. Most elite selections had aromatic qualities (Table 1.12), were extremely firm, and had little evidence of disease. Some red color was present on 68 percent of the random population (Table 1.9) and on 91 percent of the elites (Table 1.10). These figures are much higher than those in the forests at 43°N and 45°N latitude. Additionally, these apples have evolved in a harsh climate that often drops to –40°C. Spur-type trees, which have many horticultural advantages, were also prominent in this area and were seldom seen in the other forests. In this environment, we did not see trees that were greater than 50 years old as we had seen in other areas such as

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Djungarsky and Zailisky; they were much smaller and more widely spaced here also. Fruit shapes from most areas were of the globose to globose conical type (Table 1.13). Djungarsky, Sites 4.00, 5.00, 5.01. The third phase of our 1995 expedition took us to the Djungarsky mountain range where we had visited in 1993. Three sites (4.00, 5.00, and 5.01 were examined from September 2 to September 5. Djungarsky near Topelevka village (Fig. 1.1, area 5) was the same area that was visited in 1993, but fruiting was heavy this year. Two population samples of 30 accessions each were collected which included a lower elevation sample at 1170–1450 m (GMAL 3880–3909 from site 5.00) and a higher elevation sample from 1450– 1690 m (GMAL 3850–3879 from site 5.01). The mean diameter of the fruit from the 60 random accessions was 36 mm. From these 60 random accessions, 3014 seeds were collected. In addition, seeds of 14 wild elites were collected from sites 5.00 (GMAL 3654–3663 with scions collected from GMAL 3654) and 5.01 (GMAL 3650–3653). The mean diameter of the wild-elite selections was 46 mm at site 5.00 and 38 mm at site 5.01. From these 14 trees, 7618 seeds were collected. Although the fruit was of similar size from the different elevations, both the 60 random accessions and 14 elites had variable horticultural characters and levels of disease susceptibility depending on elevation. The Lepsinsk area (Fig. 1.1, Table 1.4, area/site 4.00) was visited in 1993 with minimal collections. One population of 30 accessions was collected along with eight elites in 1995. The random collection at site 4.00 included GMAL 3910–3939 with a mean fruit diameter of 34 mm with 1425 seed collected. A total of 6065 seeds from eight wild elites were collected from site 4.00 (GMAL 3664–3671), mean fruit diameter 37 mm. The diversity in this area was minimal with predominantly small, yellow, scab-infested fruit whose firmness and aromatics were low. Karatau, Sites 6.00 and 6.01. Expeditions in the Karatau Mountains (Fig. 1.1, area 6) were made September 8 to 10. In site 6.00, a random population of 30 apples (GMAL 3940–3969) was collected along a stream tributary along with 12 elites (GMAL 3672–3681 and GMAL 3694–3696). The mean diameter of the apples from site 6.00 was 39–41 mm for both random and elites. We collected 1476 seed from the 30 random trees in site 6.00 and 4081 seed from the 12 elites. Site 6.01 was a xerophytic site 300 m above the stream area. There we collected a population of 34 apples (Plate 2B) and 12 wild elites (GMAL 3682–3693 with scions also collected from GMAL 3686 and GMAL 3693) that were completely without scab infection. We collected 2177 seed from the 34 random trees in site 6.01 and 6113 seed from the 12 elites. Because the area was so dry,

27 38 5 21 0

86

207

22

53

13 28

8

30

55

34

23

20

22

<1

0

0

0

<1

0

0

0

1

1.2

b

As depicted in Fig. 1.1 Morphology descriptor 38; overall fruit shape (Table 1.3).

a

35

39

138

711

46

101

35

1.1

5

15

13

0

1

2

7

10

1

2.0

22

38

19

32

17

33

28

20

16

2.1

2

0

0

0

3

1

2

0

3

3.0

1

8

2

0

0

0

1

0

1

3.1

<1

0

0

0

<1

0

0

0

0

3.2

1

0

2

0

1

1

0

2

0

4.0

1

8

0

10

<1

0

2

0

0

4.1

Percent of accessions rated for 13 shapeb categories

2

0

4

0

1

9

1

0

1

5.0

3

23

8

0

2

3

2

1

1

5.1

<1

0

2

0

0

0

0

0

0

5.2

4:19 PM

Total

39

91

Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/ 10.00 Karatau/ 11.00 Talasky/ 12.00

1.0

No. Accessions

Mt. Range/ Area Sitea

Fruit shape of populations and elites from Kazakhstan.

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Table 1.13.

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lack of scab on the fruit does not necessarily indicate scab resistance. However, the fruit was of good horticultural quality. Fruit from the random population at site 6.01 had the highest level of firmness and aromatics of any of the populations. Trees exhibited excellent fruit retention and the high level of fruit firmness was surprising given the very high level of heat units common in the area. These apples would likely be late harvest types in most apple-growing districts. They may also be of interest as a drought resistant source for rootstocks. The apples collected along the stream at site 6.00 were almost all drops and were difficult to characterize for quality since they were earlier maturing. On September 13, we visited the ex situ collections of Professor Dzhangaliev and observed the collection of 600 apple accessions that he received over the last four years from PGRU in Geneva, New York, Corvallis, Oregon, and Davis, California, including 40 accessions that were recently grafted from our collection three weeks earlier. We observed other elite commercial types that his institute maintains as well as 70 wild ‘selections’ of M. sieversii from the Kazakhstan forests. In 1993, we imported five of these promising ‘selections’, which remain in quarantine in Beltsville, Maryland. A total of 60,081 seeds were collected from 408 individual trees in 1995 (Table 1.6). Fruit characterization records for these collections were recorded using a descriptor approach (Table 1.3). These records along with specific site data can be found on the Web at: http://www .ars-grin.gov/cgi-bin/npgs/html/search.pl. The data on the 1995 collection can be found by accessing catalog numbers: PI 600485–600584 (GMAL 3596–3699 elites) and GMAL 3700–4003 (random populations). D. 1996 Collection A fourth expedition to Kazakhstan took place August 24 to September 19, 1996. Collectors included Philip Forsline, Stan Hokanson, Thomas Unruh, and Harold Pellett. Professor Aimak Dzhangaliev was again the main host who was assisted by others. Our objectives were threefold: (1) collect seeds and leaf samples from trees at random throughout each site in order to characterize diverse populations using morphological and molecular methods; (2) collect seeds and scions of superior types within each population based on horticultural and disease characteristics; and (3) establish linkages with scientists in the Kazakh Academy of Sciences to continue cooperation on characterization of the Kazakh

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apple forests, including studies to establish frameworks for in situ reserves, translation of previous works, and scientific exchange. M. sieversii was collected in nine distinct ecosystems, five of which had never been accessed in previous years. In three of four sites previously visited, fruiting was heavier than in previous years allowing for more complete sampling. Only in the northernmost site, Tarbagatai, which was very productive in the previous year (1995), did we see sparse fruiting. In this expedition, collections included random samples of leaves and 14,000 seeds from 260 trees across the nine sites. Population sizes varied at each site from 10 to 60 trees depending on the size of the area and more significantly on the time allotted for collection at each site by our hosts. We collected an additional 14,500 seeds from 47 superior trees throughout all sites. Scions were collected for 30 of these to establish clones. All fruit and trees were characterized with up to 24 descriptors (Table 1.3) and associated site information can be found on the Web at: http://www.ars-grin.gov/cgi-bin/npgs/html/desclist.pl?115. The manuscript of an English translation of the book, The Wild Apple Trees of Kazakhstan was obtained from the author, our host, Professor Aimak Dzhangaliev and a revision of this book is published as part of this volume. 1. Geographic Sites in 1996 Ketmen, Site 10.00. On August 27, we left for our first expedition to the Ketmen mountains (Fig. 1.1, area 10) accompanied by Professor Dzhangaliev, Raisa Turehanova (botanist), and Valerie Sotnikov (horticulturist), Mukhamedjan Issayev (interpreter), Mr. Kashaganov (forestry director for the Ketmen area), and Dominique Noiton (apple breeder from New Zealand). We drove 350 km from Almaty, arriving in the village of Kolzhat in the Ketmen area near the Chinese border. We observed the Ketmen ‘selections’ in local gardens and then drove northwest 120 km to a Forestry Reserve that protects a rare species of ash (Fraxinus sogdiana). The next morning, August 28, we drove 65 km to the Ketmen Mountains west of Kolzhat that we visited the previous day and collected M. sieversii near the village of Kirghiz Sai where time was limited. Three of the 20 random accessions collected were the red flesh form of M. sieversii that the Kazakhs classify as Malus niedwetzkyana (Table 1.1). In our three previous expeditions to Central Asia, we had never seen this in the wild, but Professor Dzhangaliev reported this species is found in most of the habitats in Kazakhstan. He maintains some forms at the Main Botanical Garden in Almaty. The size of the area covered was 5.0 km × 3.0 km. One population of 20 accessions (GMAL 4057–4076) was

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collected in this area. From these 20 accessions, we collected leaf samples for DNA extraction along with 1273 seeds. The mean fruit diameter in these 20 accessions was 40 mm. In addition, two wild elite samples from site 10.00 (GMAL 4010–4011) averaged 49 mm in diameter (Table 1.8) and 845 seeds were collected. The fruit was of poor quality (Tables 1.9–1.12). Zailisky, Site 3.00. On August 30, collections were made in the Zailisky Mountains near Almaty, the same area as the collection in 1995 (site 3.00) along the Aksay River. This collection was valuable since 80 percent of the trees were fruiting in contrast to less than 50 percent in 1995. From a random population of 20 trees (GMAL 4077–4096) in a 0.5 km × 1.0 km area, we collected 1340 seeds and leaves; 35 percent of the trees were scab-free. Tarbagatai, Sites 9.00–9.05. September 1 to 4 was spent in the same area (Fig. 1.1, area 9) in the Tarbagatai mountains, near the city of Urdzhar where we had collected in 1995 (sites 9.00–9.05). This is the area that produced some excellent superior selections in 1995 when nearly 90 percent of the trees were fruiting. However, as a result of cold weather during pollination in 1996, only 15 percent of the trees were fruiting, but we were able to make a random collection of 2440 seeds and leaves from 60 trees (GMAL 4097–4156) in a 3.0 × 5.5 km area; 11 trees in site 9.00 (GMAL 4136–4146); 23 trees in site 9.01 (GMAL 4097–4119); 10 trees in site 9.02 (GMAL 4147–4156); 7 trees in site 9.03 (GMAL 4120–4126); no trees in site 9.04; and 9 trees in site 9.05 (GMAL 4127–4135). The mean fruit diameter of 60 trees was 41 mm. We collected seeds and scions from 13 superior selections in sites 9.00–9.05 (GMAL 4012–4024) ranging in size from 45 mm to 62 mm with the mean diameter being 54 mm. We also collected scions of 11 of 13 elite selections and 4295 seeds. In this same area in 1995, having more diversity to select from with more precocious fruiting, 46 superior types were selected ranging in size from 46 mm to 72 mm. Very high levels of aromatic flavor, flesh firmness, red color, flesh firmness (Tables 1.9–1.12) and freedom from scab were observed. Although we did not record scab susceptibility on the 60 randomly selected fruit in 1995, the trees were very clean throughout the entire population. The grazing sheep and goats observed in this area in 1995 were not present in 1996. Djungarsky, Sites 4.00, 4.01, 4.02, 5.00, and 5.01. On September 5 to 9, we made collections in four unique sites within the Djungarsky area (Fig. 1.1, areas 4 and 5). Sites 4.01 and 4.02, located near the city of Andreyevka were only 7 km apart but quite unique from each other and

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they had not been accessed in previous years. Collections were made very close to the village of Konstantinovka. Collections at site 4.01 were limited to seeds and leaves of 10 randomly selected trees (GMAL 4157–4166) from which we collected 474 seeds. These fruit had a mean diameter of 33 mm. We selected only those trees that were at least 50 years old since this is an area that Professor Dzhangaliev has reforested over the last 40 years with seedlings that may be native to other areas. This population area with a southwest aspect was 1.0 km × 1.0 km. Site 4.02 (not part of the reforesting project) was located on a steep northwest-facing slope that was 1.5 km × 200 m. We collected leaves and seed of 17 randomly selected trees (GMAL 4167–4183) from which we collected 815 seeds and found four superior selected types (GMAL 4025–4028) from which 1172 seeds were collected. Scions were collected from GMALs 4027 and 4028). Mean fruit size of the random population was 33 mm and for the 4 elites was 48 mm. High levels of apple scab infection were observed in sites 4.01 and 4.02. We observed a full range of fruit diversity in these populations with 60 to 80 percent of the trees fruiting. Site 4.00 in the Djungarsky range was located near the village of Lepsinsk 20 km southwest of sites 4.01 and 4.02. This was the same area accessed in 1993 and 1995. However, we were pleasantly surprised with the fruit diversity observed in 1996 since 80 percent of the trees were fruiting; in previous years, the fruiting intensity was less than 40 percent. The size of the population area was 1.5 km × 2.0 km with large variance in elevation, slope, and aspect. Thirty trees were sampled at random (GMAL 4184–4123) from which we collected 1535 seeds along with two superior selections (GMAL 4029–4030 with 641 seeds and scions were also collected for these two elite specimens). Mean fruit size of the random population was 34 mm and for the two elites was 44 mm. Apple scab was rather heavy in the area but 4 of 30 trees sampled along with the superior selections were impressively clean of scab. We completed our collections in the Djungarsky range by returning to the site near the village of Topelevka (sites 5.00 and 5.01) about 35 km southwest of site 4.00. This same site was accessed in 1993 and 1995. We were impressed with the diversity available in 1996 with 95 percent of the trees fruiting. We sampled 54 randomly selected trees (GMAL 4214–4232 and 4249–4267 from site 5.00; GMAL 4233–4248 from site 5.01) from an area of 3.0 km × 5.0 km with a very large variance in elevation along with diverse slopes and aspects. The first ten trees sampled in the random populations were near an area of seedling reforestation. Trees GMAL 4224–4267 were collected on the last two days from localities of this area that had not been visited in 1993 or 1995. The mean

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diameter of the fruit from these 54 trees was 35 mm from which we collected 2928 seeds. We also collected 10 superior selections (GMAL 4031–4040 with scions collected from GMAL 4031–4036 and 4038) based on fruit quality, tree form, and disease resistance. The size range of these was 54 mm to 65 mm (mean fruit diameter was 58 mm) in comparison to the size range in 1995 of 37 mm to 56 mm (Table 1.8). From these 10 accessions 3190 seeds were collected. Quality of these fruits was also much more impressive than those collected during the sparse fruiting years of 1993 and 1995. All superior selections were without scab and 35 percent of the randomly selected materials were scab-free. We also collected seeds and scions of two genetic dwarfs (GMAL 4033 and 4034) found at an elevation of 1500 m to 1600 m. We initially suspected that they were dwarfed as a result of growing at high elevation, but vigorous, full sized trees were found in close association. Karatau, Site 11.00. We arrived in the city of Chimkent 700 km west of Almaty on September 12 and drove 70 km northeast to the forestry camp named Kokbulak, meaning “blue spring.” This site in the Karatau Mountains is 50 km southeast of the Boraldy forestry camp (sites 6.00/6.01) that we visited in 1993 and 1995 (also in the Karatau range). The habitat in Kokbulak, very similar to that in Boraldy, is a xerophytic area receiving only 250 mm of annual precipitation. However, there is a spring-fed stream running through the area. From September 12 to 14, we collected seed and leaf samples from 40 randomly selected M. sieversii (GMAL 4268–4307) over an area of 5.0 km × 7.0 km. The horticultural quality based on size, firmness, and flavor of these collections was good (Tables 1.9–1.12). Trees were 60 percent scab-free but this may be avoidance rather than resistance since the dry area may not be conducive to scab outbreaks. A population of 30 trees (GMAL 4268–4297) was collected near the active stream but may not be xerophytic types, and may be similar to those found along a stream in Boraldy in 1995 (site 6.00). The remainder of the random population (GMAL 4298–4307) was collected on a dry north-facing slope and may have drought tolerance similar to those we found in Boraldy in 1995 (site 6.01). The mean diameter of fruit from the 40 random collections (2295 seeds) was 42 mm. We collected 3990 seed from 13 superior types (GMAL 4041–4053) and scions from GMALs 4042–4043, 4049, and 4051–4053. Mean fruit size was 55 mm (Table 1.8), similar to the large fruit found in Tarbagatai in 1995. One accession had an average fruit size of 76 mm (Plate 1B). The quality of these fruits parallels those from the northern area of Tarbagatai. Talasky, Site 12.00. We drove 60 km southeast of site 11.00 in Karatau (Fig. 1.1) to the Talasky Mountain area, an area that we had not previ-

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ously visited. This was a restricted area listed as a National Park called Aksu Jabagli. The collection area is a deep narrow canyon 1270 m at the top and 1010 m at the river level. We hiked the steep winding trails on the southern slope down to the river level where we found M. sieversii growing up the face of the northern slope to the top of the canyon. We collected only along the river at 1010 m and up to 1025 m on the northern slope including leaves and 741 seeds from 10 randomly dispersed M. sieversii (Plate 2C) and 319 seeds and scions from three superior types (GMAL 4054–4056 with scions of GMAL 4054–4055). The quality of the fruit in this area was impressive (Tables 1.9–1.12) with mean fruit diameter of 42 mm. Mean fruit diameter of the three elites was 65 mm (Table 1.8). Two of the superior types (Plate 1C) had long ellipsoid/conical shapes that we had rarely seen before. This site had the highest levels of ellipsoid and oblong-shaped fruit (Table 1.13). The site was quite similar to that in Karatau with low rainfall and high summer temperatures. Although it would be difficult to collect along the entire steep north face, this area should be visited again in order to sample the trees on the upper slopes. A total of 28,039 seeds were collected from 307 individual trees in 1996 (Table 1.6). Fruit characterization records for these collections were recorded using a descriptor approach (Table 1.3). These records, along with specific site data can be found on the Web at: http://www.arsgrin.gov/cgi-bin/npgs/html/search.pl. The data on the 1996 collection can be found by accessing these catalog numbers: PI 600585–600624 (GMAL 4010–4056 elites) and GMAL 4057–4317 (random populations). IV. MAINTENANCE A system has been developed to back up the PGRU clonal collection by storing dormant buds in liquid nitrogen (LN). This system was first described by Sakai (1960), and refined for apple by Sakai and Nishiyama (1978) and Tyler et al. (1988). Stushnoff (1991) described how this method could be useful as a practical approach for storing germplasm in the NPGS. The protocol for storing germplasm as part of the base collection at the National Seed Storage Laboratory (NSSL) in Fort Collins, Colorado was developed as a pilot project on 64 diverse accessions (Forsline et al. 1998). This trial was also set up to determine longevity of storage over 40 years. No decline in viability was observed after four years of storage (Forsline et al. 1998) including some accessions now stored for 12 years (unpublished). Nearly 85 percent of the clonal collection (2000 accessions) has been backed up in LN-storage at the base collection at NSSL. Of these, 1700

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have been tested for recovery by grafting using the methods developed by Forsline et al. (1998) and refined for efficient storage and recovery (Forsline et al. 1995, 1996a, 1998). We have obtained good results for nearly all Malus accessions stored in the base collection. Grafted buds of 98 percent of M. ×domestica have shown adequate recovery. At least five, 10-bud containers of each accession are cryogenically stored. Adequate recovery following grafting onto seedling rootstock as the baseline test for the cryogenically stored material is 40 percent (Forsline et al. 1995, 1996a, 1998). Research is being conducted to determine a protocol to cryopreserve a small group of recalcitrant (nonhardy) apples. We have found that some of these such as ‘Golden Delicious’ vary in cryopreservability from year to year probably due to variable cold acclimation conditions. We concluded that most accessions can be successfully cryopreserved. Storage of M. sievesii seed from Central Asia is done both at NSSL and PGRU (Table 1.7). Presently, 51,316 seeds are stored at PGRU at –20°C with 39,098 seed stored at NSSL in LN. In addition, over 5000 seedlings have been screened for disease resistance traits in a collaborative program with the Plant Pathology Department at Cornell University, Geneva. Many of these seedlings have been distributed to sites outside of PGRU. Seed germination for a subset of the stored seed was monitored (Table 1.14) with over 93 percent germination of M. sieversii seeds. In Table 1.14.

Germination of M. sieversii seeds from Kazakhstan.

Mt. Range/ Area Sitea

No. Accessions Germinated

Percent Accessions with More Than 90 Percent Germ.

No. Seeds Stratified

Percent Seed Germinated

22

72

157

92

43

58

319

92

26

50

300

89

44

88

663

97

98

64

1323

91

21 24 13

62 83 62

154 255 153

84 91 90

291

68

3297

93

Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00 Total a

As depicted in Fig. 1.1.

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addition, chilling requirement was monitored for this subset (Table 1.15) with variable levels from 83 to 127 days. Longer chilling requirement in seeds may correlate with late bloom in those seedlings when they eventually begin to flower (Mehlenbacher and Voordeckers 1991). Areas 6 and 9 (Fig. 1.1) had the longest chilling requirements. Those seedlings are now being characterized for flowering times to determine if this correlation proves to be true. Over 1600 of these seedlings are being maintained at PGRU for horticultural characterization. When traits of interest are found in some of these seedlings, they too, will become potentially useful clones and propagated on rootstocks, established in orchards and backed up in cryogenic storage. In addition, when the 44 clonal accessions of M. sieversii are released from quarantine, they will also be maintained in orchards and backed up in cryogenic storage. The size of the permanent field collection is being reduced utilizing cryogenic storage. Since all accessions in the permanent field collection will be backed up at NSSL, it is possible to consider reducing each accession to only one propagule in costly field plantings. Cryogenic storage is very cost-efficient. We have calculated that after an initial cost

Table 1.15. Chilling requirement for M. sieversii seed germination collected in Kazakhstan.

Mt. Range/ Area Sitea

No. Accessions Germinated

Range of Chilling Requirement (days)

Accessions with More than 110 Days Chilling (%)

Ave. No. Days Chilling Needed

22

83–106

0

96

43

88–116

2

101

26

85–108

0

98

44

106–127

52

114

98

83–127

22

108

21 24 13

80–108 88–114 83–106

0 4 0

94 101 97

291

83–127

16

104

Zailisky/ 3.00–3.04 Djungarsky/ 4.00–4.02 Djungarsky/ 5.00–5.01 Karatau/ 6.00–6.01 Tarbagatai/ 9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00 Total a

As depicted in Fig. 1.1.

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of $50 to process an accession and test for baseline recovery, the cost of LN for long-term storage is only one U.S. dollar per year, per accession, whereas the estimate to establish and maintain each accession in the field is $50 to $75 per year. To facilitate protocols for cryogenic storage, buds can be collected from January to March at PGRU, and can be stored at –4°C for up to six months before processing and placing in LN storage (Forsline et al. 1996b). This greatly facilitates the implementation of the protocol. Cryogenic storage has given us a convenient way to ensure against losses from abiotic and biotic stresses. Fire blight (pathogen: Erwinia amylovora Burrill) has been our greatest challenge. We have had some severe epidemics in the PGRU orchards and have maintained records on levels of susceptibility for each accession. All accessions with high levels of susceptibility are stored in LN. In addition, we are repropagating our orchards on EMLA 7 rootstock, which is tolerant of fire blight and imparts less vigor than on seedling rootstock, which tends to exacerbate the problem of fire blight. Seed accessions from collections of wild Malus are maintained in moisture-proof envelopes and stored at –20°C according to IBPGR guidelines (IBPGR 1985). Portions of original seed are stored at NSSL and seeds are distributed to collaborative evaluators from the active collection at PGRU. V. DISTRIBUTION Distributions include scions (dormant and summer bud wood), pollen, seeds, leaves for biochemical/molecular analysis, and recently, DNA samples from the core collection (Forsline 1996). The 206-member core collection has been distributed many times for characterization and evaluation. Distributions, begun in 1989, are tracked through GRIN (USDA 2000). The PGRU has distributed over 1300 seed populations (30,675 seeds, Table 1.6) of M. sieversii to 24 cooperators (Table 1.16) worldwide for evaluation and is summarized in Table 1.7. In addition, over 2000 seedlings that were resistant to apple scab in screening at Geneva, New York, were sent to five cooperators. Cooperators successfully germinated some 20,000 of the 30,765 distributed seeds. The stratification times reported for germination was highly variable, ranging from as few as 38 to over 200 days. Many cooperators noted that M. sieversii seeds required a longer stratification period than typically expected for domesticated apple. Following initial mortality and screening for disease resistance or vegetative traits, over 15,000 seedlings of M. sieversii

Plant breeding, pomology Plant breeding

Curt Rom Cecil Stushnoff Schuyler Korban Joe Hecksel David Bedford, James Luby John Kreutzigera

Joseph Goffreda

Philip Forsline, Stan Hokanson, Warren Lamboy, Jing Yu Herb Aldwinckle, Ki-Sung Ko, Sang-Bum Lee, Tim Momol

University of Arkansas Fayetteville, USA

Colorado State University Fort Collins, USA

University of Illinois Urbana, USA

Private Grower, Eaton Rapids, Michigan, USA

University of Minnesota, St. Paul, USA

Heartland Germplasm York, Nebraska, USA

Rutgers Fruit Research and Ext. Center, New Brunswick, New Jersey, USA

USDA, ARS, Plant Genetic Resources Unit, Geneva, New York, USA

Cornell University, Dept. of Plant Pathology, Geneva, New York, USA

Plant pathology fire blight, cedar apple rust,

Pomology, physiology, plant pathology, molecular genetics

Plant breeding

Pomology

Plant breeding

Plant breeding, physiology

(continued)

Resistance to apple scab, Rosellinia necatrix and Helicobasidium mompa

Resistance to apple scab, fire blight, and cedar apple rust, fruit traits, molecular genetic studies of genetic diversity

Resistance to apple scab, fire blight, and cedar apple rust, fruit traits and storage ability

Disease and pest resistance, late flowering and leafing dates, fruit traits

Resistance to apple scab and fire blight, cold hardiness, fruit traits

Horticultural traits

Project terminated

Cold hardiness

High chilling requirement, fruit traits

Cold hardiness

Traits of Special Interest

4:19 PM

Breeding, pomology

Horticulture

Jennifer McBeath

University of Alaska Fairbanks, USA

Expertise of Evaluators

Name

Researchers evaluating M. sieversii collected in Central Asia.

8/8/02

Organization and Location

Table 1.16.

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41

42 Name Susan Brown, Norman Weeden Ian Merwin Harvey Reissig David Ferree, Diane Miller Donald Hendricks

Mitch Lynd

Bruce Barritt

Kevin Bradley Brian Smith

Cornell University, Dept. of Hort Science, Geneva, New York, USA

Cornell University, Dept. of Fruit & Vegetable Science, Ithaca, New York, USA

Cornell University, Dept. of Entomology, Geneva, New York, USA

Ohio Agricultural Research and Development Center, Wooster, USA

The Dawes Arboretum Newark, Ohio, USA

Private Grower, Midwest Apple Improvement Association, Johnstown, Ohio, USA

Washington State University, Wenatchee, USA

Private Grower, Madison, Wisconsin, USA

University of Wisconsin River Falls, USA

(continued)

Organization and Location

Table 1.16.

Breeding, pomology

Pomology

Breeding, pomology

Pomology, botany

Botany

Pomology

Entomology

Disease resistance, yield, manageable tree stature and habit, winter hardiness, late bloom, short juvenility, early fruit maturity, fruit traits

Horticultural traits

Resistance to pests, fire blight and powdery mildew, juvenility, cold hardiness, sunburn tolerance, fruit traits

Resistance to apple scab, cedar apple rust, and fire blight, late flowering, fruit traits

Resistance to apple scab, cedar apple rust, and fire blight, late flowering, fruit traits

Resistance to apple scab, cedar apple rust, and fire blight, late flowering, fruit traits

Resistance to apple maggot

Resistance to soil borne apple replant pathogens

4:19 PM

Pomology

Dwarf growth habit

Traits of Special Interest

8/8/02

Plant breeding, molecular genetics

Expertise of Evaluators

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Cheryl Hampson, Harvey Quamme Campbell Davidson

Daryl Hunter Christianne Deslauriers, Charles Embree Rolf Büttner, Martin Geibel

Hideo Bessho, Junichi Soejima Vincent Bus, Nadozie Oraguzie

Dag Roen

Agriculture Canada Summerland, British Columbia, Canada

Agriculture and Agri-Food Canada, Morden, Manitoba, Canada

Kings Landing Historical Corporation, Fredericton, New Brunswick, Canada

Agriculture and Agri-Food Canada, Kentville, Nova Scotia, Canada

Genebank for Fruit, Dresden, Germany

Ministry of Agriculture, Forestry & Fisheries, Yamanishi, Japan

HortResearch Havelock North, New Zealand

The Norwegian Crop Research Institute, Hermansverk, Norway

Plant breeding

Plant breeding, genetics, plant pathology, entomology, biometrics

Plant breeding

Pomology, plant breeding

(continued)

Resistance to apple scab and powdery mildew, fruit traits and storage ability, fruit rots and disorders

Resistance to apple scab, powdery mildew, fire blight, woolly apple aphid, leafroller, apple leaf curling midge, leafing and flowering dates, juvenility, burr knots, stooling ability, tree habit and vigor, fruit traits

Horticultural traits

Resistance to apple scab and powdery mildew (including molecular markers), plant stature, length of vegetative period, fire blight, fruit characters, genetic diversity for molecular and morphological traits

Disease resistance, cold hardiness, growth habit, fruit traits

Horticultural traits

4:19 PM

Plant breeding, physiology

Resistance to fire blight, cold hardiness, fruit traits

Disease resistance, cold hardiness, drought tolerance

8/8/02

Pomology

Plant breeding

Plant breeding, cold hardiness physiology

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43

44 Barrie Juniper Taiiboos Human

University of Oxford, Oxford, UK

INFRUITEC, Stellenbosch, South Africa

Deceased June 1999

Plant breeding, pomology

Botany, molecular genetics

Botany, molecular genetics

Resistance to woolly apple aphid, Phytophthora, apple scab, powdery mildew and drought, chilling requirement, sunburn tolerance, fruit traits

Origin of domesticated apple

Genetic diversity

Traits of Special Interest

4:19 PM

a

Gary Britz Rita Farrel

University of Reading, Reading, UK

Expertise of Evaluators

8/8/02

MULTIFRUIT EPPINDUST, South Africa

Name

(continued)

Organization and Location

Table 1.16.

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remain in field plantings at PGRU and at sites of cooperators for further characterization. The 44 clonal elite accessions remained in quarantine as of the end of 2001; however, permission for provisional release of 39 of the 44 accessions to the PGRU and Washington State University has been granted. They have been screened for apple scab by Cornell University researchers in the greenhouse at the PGRU and were planted on dwarfing rootstocks in the field in 1999 and 2000 according to guidelines of the provisional release. After three years in a field planting at PGRU, we have observed fruiting of 13 of these elites (Plate 1D) and have begun to characterize them for comparison with the characterization that was made when they were collected in Kazakhstan.

VI. CHARACTERIZATION AND EVALUATION A. Core Collection and Main Collection A core collection of apple has been established for the purpose of efficient characterization and evaluation. Brown (1989) first discussed the concept of core collections. The value for germplasm collections in the NPGS was further discussed (Kresovich et al. 1995) and developed for fruit crops including apple (Grauke et al. 1995). The core collection for apple includes 206 diverse accessions out of the total collection of 2438 clonal accessions. The initial core collection was designated in 1991 (Forsline 1996). It consists of: (1) 78 M. ×domestica cultivars with diverse pedigrees, a range of known traits, some with known disease resistance, and old and new types; (2) 27 PRI selections from the Purdue-RutgersIllinois cooperative breeding program (Crosby et al. 1992) with multiple disease/insect resistance; (3) 10 Malus species accessions with disease resistance traits; (4) 91 Malus species accessions with two to five representatives of each of 30 species (Way et al. 1990) having diverse geographic representation. This list can be found through GRIN (USDA 2000) on the Web at: http://www.ars-grin.gov/cgi-bin/npgs/html/ desc_find.pl?crop=APPLE&115154=Y. Each accession in the core collection is replicated four times and planted in five climatically diverse sites in New York, North Carolina, Washington, Illinois, and Minnesota. The core collection is a dynamic entity whereby certain accessions may be removed or others added as we learn more about the genetic character of the entire collection including newly acquired accessions. The core collection has been completely characterized for 25 priority morphological descriptors as listed on the Web at: http://www.ars-grin.gov/

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cgi-bin/npgs/html/crop.pl?115. In addition, we have made images which can be accessed on the Web at: http://www.ars-grin.gov/ars/NoAtlantic/ Geneva/malus_proj.html for most of the core collection. The other members of the clonal collection (2208 accessions) have now been characterized using the same priority descriptors. To better understand and utilize plant genetic resources, biochemical characterization (Weeden and Lamb 1985) has been used. More recently Kresovich et al. (1993), using molecular techniques, found efficient and informative DNA markers for germplasm characterization. Simple sequence repeats (SSRs) have a high level of polymorphism and a highinterspersion rate making them an abundant and effective genetic marker for maintenance of genebank diversity. Because of the potential of SSRs for automated analysis and their codominant nature, they are more useful than other PCR-based marker systems. SSRs have been identified in the entire Malus core subset (Hokanson et al. 1998, 2001). Automated fluorescence-based detection of polyacrylamide gels to improve analytical resolution, increase throughput and decrease unit cost, has been employed. In combination with passport and horticultural data, we intend to apply genetic information revealed by SSR primer pairs to better comprehend genetic identity and relatedness in the Malus germplasm collection, to help develop and refine the Malus core subset (Hokanson et al. 1997a, 1998, 2001; Szewc-McFadden et al. 1995 and 1996), and to advance our understanding of the extent and maintenance of genetic diversity of random Malus populations from Central Asia (Lamboy et al. 1996). The collection is evaluated through collaborative research with cooperators from around the world. By using the core collection as a test array, Howell et al. (1996) found superior virus indicators for the apple stem grooving virus: (1) M. ×micromalus, Mak. (GMAL 273a, PI 594092); (2) M. yunnanensis (French.) Schneid. (GMAL 2342, PI 589758); and (3) M. tschonoskii (Maxim.) Schneid. (GMAL 1834, PI 589395). The core collection has also been evaluated for: (1) disease resistance including: fire blight (Erwinia amylovora Burrill), apple scab (Venturia inaequalis (Cooke) Wint.), cedar apple rust (Gymnosporangium juniperivirginianae Schwein.), sooty blotch (Gloeodes pomigena Schwein.), flyspeck (Zygophiala jamaicensis Mason), bitter rot (Glomerella cingularia Stonem.), black rot (Botryosphaeria obtusa Schwein.), and white rot (Botryosphaeria dothidea, Moug.); and (2) arthropod resistance including: European red mite (Panonychus ulmi, Koch), apple maggot (Rhagoletis pomonella, Walsh), and woolly apple aphid (Eriosoma lanigerum Hausm).

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B. Central Asian Collections Over 50 scientists (Luby et al. 2001) and technicians with diverse expertise have been evaluating M. sieversii accessions at locations in several countries (Table 1.16). Because most trees held by cooperators are not yet fruiting, evaluation has focused on disease resistance or vegetative traits. Currently HortResearch in New Zealand and the PGRU are the only sites with large numbers of fruiting trees. Most evaluators intend to evaluate for standard fruit traits such as size, color, texture, aroma, and flavor and vegetative traits such as growth habit and vigor. At PGRU, we have characterized many of the earliest fruiting seedlings (Plate 2D). Most evaluators also are targeting disease and pest resistances and other traits related to adaptation in their regions. These plans are summarized in Table 1.16. Progress for some traits is discussed in the following section. 1. Disease and Pest Resistance Apple scab. Several cooperators are screening young seedlings for apple scab resistance. In New Jersey, nearly 2000 seedlings were screened for apple scab, and J. Goffreda (pers. comm. 1999) reported that a high proportion exhibited resistance similar to that conferred by the Vr gene. In New York, nearly 5000 seedlings representing 12 regions (Fig. 1.1 and Table 1.17) have been evaluated for apple scab resistance with a total of 27 percent of those screened being resistant (Aldwinckle et al. 1997). Combining data from screenings of material collected in all the expeditions revealed major variation in the incidence of apple scab resistance related to region of origin (Table 1.17). This trend was also observed previously with a smaller subset. Area 4 had the highest proportion (49%) of the resistant seedlings while area 1 had the lowest proportion (5%). In New Zealand, over 2500 seedlings from 60 seed lots from the 1995 and 1996 expeditions were evaluated in the greenhouse (Bus et al. 2002). Approximately 24 percent of the seedlings were resistant in these evaluations with a range among half-sib families from 0 to 70 percent. Areas 3, 10, and 11 had less than 10 percent resistant seedlings while areas 4, 5, 6, and 9 had greater than 25 percent resistant seedlings (Table 1.17). In addition, Vincent Bus (pers. comm.) also evaluated over 1400 trees from 52 seed lots from the 1993 expedition, following natural apple scab infection in the field. These trees form part of a large apple breeding population aimed at increasing genetic diversity for many traits (Noiton and Shelbourne 1992). Using estimated variance components, they found a low heritability (0.13 on family mean basis), due mainly to high levels of resistance and little variation among these families. They did

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Table 1.17. Apple scab resistance of Malus sieversii seedlings from 12 regions in Central Asia screened in greenhouses in New York (NY, 1993, 1995, and 1996 collections), New Zealand (NZ, 1995 and 1996 collections), and Minnesota (MN, 1995 collection). No. of seedlings screened Areaa

Percent of seedlings resistant

NY

NZ

MN

NY

NZ

MN

1 2 3 4 5 6 7 8b 9 10 11 12

21 101 450 369 1175 705 383 151 1125 123 226 133

– – 262 287 277 133 – – 684 86 244 –

– – – – – 362 – – 1171 – – –

5 24 17 49 27 37 25 07 27 06 23 14

– – 8 28 45 25 – – 29 2 5 –

– – – – – 74 – – 62 – – –

Total

4971

1973

1533

27

24

65

a

Areas described in Figure 1.1 and Table 1.2. Open pollinated seedlings of Professor Dzhangaliev’s forms at Almaty (Kazakhstan) Botanical Garden. b

not detect any significant differences for scab incidence among the collection regions. In Minnesota, seedlings from areas 6 and 9 were screened and 65 percent were resistant (Table 1.17). The results from the two areas were similar, but individual families ranged from 0 to 88 percent resistant seedlings. Several seed lots have been screened at multiple sites (Table 1.18). Although some families consistently produced high proportions of resistant seedlings (e.g., GMAL 3607, 3631, 4024, 4089, and 4177) or high proportions of susceptible seedlings (GMAL 3609, 3643, 4011, 4068, 4071, 4086, 4171, 4309, and 4315) over multiple sites, others were quite variable. Inconsistencies in the seedling screenings may be due to different local inoculum sources, or to sampling effects from small sample sizes for some accessions at some locations. Alternatively, they may be due to a variation in age or physiological state of seedlings or to different test conditions that could influence infection success. Of the 39 elite clonal accessions granted provisional release from quarantine, 30 have been evaluated for apple scab resistance in the

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Table 1.18. Apple scab resistance of Malus sieversii seedlings in Central Asia from seed lots evaluated in greenhouses in New York (NY), New Zealand (NZ), and Minnesota (MN) after inoculation with local strains. Seedlings screened

Seedlings resistant (%)

GMAL/PI Accession No.a

NY

NZ

MN

NY

NZ

MN

3604/600493 3605/600494 3607/600496 3608/600497 3609/600498 3618/600507 3625/600514 3627/600516 3631/600520 3634/600523 3636/600525 3643/600532 3683/600571 3688/600574 3691/600577 4011/600586 4024/600598 4032/600606 4038/600609 4068/— 4071/— 4086/— 4089/— 4171/— 4177/— 4190/— 4209/— 4302/— 4309/— 4315/—

8 11 15 14 14 8 15 13 14 14 14 15 15 13 15 24 26 24 24 7 5 7 5 6 5 7 6 5 7 7

42 20 38 23 35 40 38 42 45 39 37 48 38 32 36 43 49 35 40 19 24 22 15 20 17 23 19 13 24 17

23 44 24 13 – 20 36 9 38 34 19 40 27 – – – – – – – – – – – – – – – – –

12 45 60 50 7 38 73 54 71 29 71 13 60 62 27 0 62 33 38 14 0 0 80 0 100 29 67 0 0 0

26 40 68 26 9 8 8 55 62 49 24 0 45 19 25 2 59 46 48 0 4 0 53 5 65 70 32 23 0 0

78 55 92 0 – 60 53 56 58 85 90 57 78 – – – – – – – – – – – – – – – – –

a

GMAL 4068–4315 are from the random collections with small seedlot quantities. No PI numbers have been assigned to those.

greenhouse using replicate, grafted plants. Four of these accessions show apple scab resistance on all ten replicate trees. Five others were resistant on some trees. In addition, Mehlenbacher and Weeden (pers. comm.) extracted DNA from five elite accessions that had 40 percent or more of their offspring resistant to apple scab and screened for the presence of markers that had been linked to apple scab resistance genes in

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other Malus accessions. All five M. sieversii had RAPD markers P415B and UBC562 markers for the Vr resistance gene. However, since these markers were also present in many other accessions, including susceptible ones, they may be of little value. Four of these five accessions also contained one or more markers that were quite rare in germplasm that did not exhibit apple scab resistance. GMAL 4326 (Q35771), GMAL 4331 (Q35777), GMAL 4334 (Q35780) had OPB12 for Vm. GMAL 4327 (Q35772) had OPB12 for Vm, the CS5 RAPD for Vf, and the UBC220 marker for Vb. Since the elite selections were initially identified when collected in Kazakhstan based on their superior fruit traits, these accessions represent potentially new sources of multiple resistance genes in a horticulturally desirable background. Fire Blight. Extensive evaluations for fire blight (Erwinia amylovora) resistance have been conducted in New York and New Zealand. In a cooperative program of Cornell University and PGRU (Momol et al. 1999), 1125 seedlings from the 1989 and 1993 expeditions were inoculated with fire blight, and 29 percent were resistant. The status of the accessions from the 1989 collection is listed in Table 1.19. Work is in progress to screen subsequent collections. One fire blight resistant seedling, GMAL 3280.h (a single seedling designated as ‘.h’ to distinguish from other half-sibs in the population), had desirable fruit quality and large fruit size (56 mm diameter). These results are significant because we have identified a resistant genotype with desirable horticultural traits. Natural occurrence of fire blight has been monitored in the seedling grow-out at PGRU. After 3 to 4 years in the field, 25 percent of the seedlings are showing significant infection (Table 1.20). This method does not allow us to define resistance but it does indicate levels of susceptibility. There is a pattern that shows seedlings from some sites in Kazakhstan are much more susceptible than from other sites. Table 1.19.

Fire blight resistance of M. sieversii seedlings with artificial inoculation. Resistant seedlings

Country/Area Site

No. Accessions

No. Inoculated

No.

Percent

Tajikistan/1.00 Uzbekistan/2.00–2.02 Kazakhstan/ 3.04a

3 9 34

24 69 266

19 11 80

79 16 30

Total

46

359

110

30

a

Zailisky mountain range.

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Table 1.20. Natural occurrence of fire blight in Kazakhstan seedlings at PGRU after three to four years in the orchard. Seedlings infected Mt. Range/Area Sitea

No. Seedlings

No.

Percent

Zailisky/3.00–3.04 Djungarsky/4.00–4.02 Djungarsky/5.00–5.01 Karatau/6.00–6.01 Kyrgyzstan/7.00 Tarbagatai/9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00

24 100 169 195 5 372 54 131 101

0 7 27 36 2 162 4 27 18

0 7 16 19 40 44 8 21 18

1151

283

25

Total a

See Fig. 1.1.

In New Zealand, 936 trees from the 1993 expedition were evaluated in their fifth leaf for fire blight incidence following natural infection in the orchard. Approximately 87 percent of the trees remained disease-free in this field evaluation while Momol et al. (1999) found that 33 percent of 775 seedlings from the same 1993 collection expedition were resistant when inoculated as young seedlings. Seedlings from areas 3 and 6 were more susceptible than those were from areas 4, 5, and 7; whereas, no differences were observed in New Zealand following natural infection. Two of the most susceptible families (PI 600479 and PI 600480) were from a single site in Kyrgyzstan (area 7). Of the six most resistant families, three (PI 600428, PI 600429, and PI 600444) were from region 4 in east central Kazakhstan, and two (PI 600468 and PI 600476) were from region 6 in south central Kazakhstan. Inoculations of young seedlings by Momol et al. (1999) showed that PI 600468 was among the more resistant families. However, in contrast to the New Zealand field results, PI 600480 was among the more resistant families, but PI 600429 and PI 600444 were among the more susceptible ones. These apparent contradictions may result from a difference in natural versus artificial inoculation techniques, physiological differences in the trees due to age, genetic differences in the seed lots due to sampling, or different pathogen races. Apple Replant Pathogens. In New York, Isutsa and Merwin (2000) evaluated some M. sieversii accessions, as well as other Malus species, for their resistance or tolerance to apple replant pathogens (ARP) based on

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their relative biomass accumulation when grown in orchard soils. The ARP included various species of Pythium, Cylindrocarpon, Fusarium, Rhizoctonia, and Phytophthora, as well as nematodes. Seedlings from two families (PI 600427 and PI 600563) were categorized as tolerant of ARP. Powdery Mildew. In Germany and New Zealand, all seedlings are being evaluated for resistance to powdery mildew (Podosphaera leucotricha (Ell. & Ev.) E. S. Salmon). In Germany, the juvenile susceptibility to powdery mildew decreased significantly with the age of the plants (Geibel et al. 2000). Nearly all plants were susceptible when evaluated in the first season of growth, but mildew infections in 1998 were only 70 percent in the 2-year-old plants and 45 percent in the 3-year-old plants over all populations. Depending on the accession, 10 to 60 percent of the plants in a family were resistant. Other Diseases. Seedling evaluations for cedar apple rust (Gymnosporangium juniperi-virginianae Schwein.) resistance are being conducted in New York and New Jersey. In New York, over 2200 seedlings were screened with approximately 50 percent of those resistant (Table 1.21). Resistance among the Kazakh sites ranged from 34 to 78 percent. In New York, Lee, Ko, and Aldwinckle (2000) screened seedlings for resistance to western white root rot (Rosellinia necatrix Prill.) and violet root rot (Helicobasidium mompa Tanaka) and identified some populations with apparent resistance.

Table 1.21.

Cedar apple rust resistance of M. sieversii. Resistant seedlings

Mt. Range/Area Sitea

No. Inoculated

No.

Percent

Zailisky/3.00–3.04 Djungarsky/4.00–4.02 Djungarsky/5.00–5.01 Karatau/6.00–6.01 Zailisky/8.00 Kyrgyzstan/7.00 Tarbagatai/9.00–9.05 Ketmen/10.00 Karatau/11.00 Talasky/12.00

97 194 610 207 90 164 520 66 162 104

40 107 245 115 31 78 404 25 64 36

41 55 40 56 34 48 78 38 40 35

2214

1145

52

Total a

See Fig. 1.1.

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Insect Resistance. In New York, Reissig (pers. comm. 1999) conducted laboratory studies to compare the oviposition preference and survival of the apple maggot (Rhagoletis pomonella Walsh) in fruit from seedlings of M. sieversii. During the two years of the study, fruit from 43 different seedlings were evaluated from material collected in 1989. Oviposition preference was compared by picking the fruit in July and exposing the selections, along with fruit from ‘McIntosh’, to groups of apple maggot females from a laboratory colony in clear Plexiglas cages and counting subsequent oviposition punctures and eggs in the fruit. Apple maggot females oviposited in fruit from all of the M. sieversii seedlings tested, but fruit was generally less infested than the ‘McIntosh’ fruit by 3 to 94 percent. Apple maggot females also infested fruit from all seedlings in a “nochoice” test. After infestation, fruit was incubated on racks over waterfilled dishes and exiting larvae were collected to compare survival rates. Larvae survived in fruit from all of the selections tested, but numbers of larvae surviving in the M. sieversii fruit were generally lower than in ‘McIntosh’ standards. Although many of the M. sieversii selections were less favorable for apple maggot oviposition and survival than ‘McIntosh’, most of the fruit was somewhat smaller (avg. 2.1 to 3.3 cm in diameter) compared to the ‘McIntosh’ fruit (average 4.3 cm in diameter). Additional studies are needed to determine if the observed differences between these selections and the standard fruit are due to physical characteristics or chemical factors. In New Zealand, seedlings from the 1993 collection were evaluated for resistance to woolly apple aphid (Eriosoma lanigerum Hausmann) in order to estimate heritabilities and combining abilities and identify resistant genotypes. In a preliminary study on trees grown from root cuttings from a subset of the 1993 seedlings, resistance to light brown apple moth (Epiphyas postvittana Walker) and apple leaf curling midge (Dasyneura mali Kieff.) were investigated. The findings were inconclusive, perhaps because the screening technique was based on leaf damage rather than development or survival rates (Wearing and Colhoun, 1999). However, the indication of a strong genetic component in the variation for incidence of apple leaf curling midge warrants further research. Multiple Resistance. Several cooperators are screening seedlings for resistance to multiple diseases and pests. Some genotypes with resistance to multiple diseases have already been identified. In New York, for example, 775 seedlings from 33 seed lots collected in 1993 from regions 3, 4, 5, 6, 7, and 8 were inoculated with apple scab, cedar apple rust, and fire blight. Of these seedlings, 23 percent were resistant to apple

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scab (Aldwinckle et al. 1997), 38 percent were resistant to cedar apple rust and 33 percent were resistant to fire blight (Momol et al. 1997). As a result of these multiple disease resistance screens, 207 seedlings with putative multiple resistance were selected from the original 775 for further horticultural evaluation. We suspect that this group may be late blooming or have a mechanism to survive spring frosts since this 1993 collection was made after a severe spring frost in the wild apple forests of Central Asia that resulted in only 2 percent of the wild trees producing fruit. The majority of these selected seedlings were sent to the research group in Ohio (Table 1.16) where they are being observed at the Dawes Arboretum. Ohio was chosen since this is an area that is often challenged by early spring frosts. Researchers in Germany noted that every population evaluated included some plants that were resistant to apple scab and powdery mildew. After two years of evaluation a total of 64 plants from nearly 1100 seedlings had neither scab nor powdery mildew symptoms (Geibel et al. 2000). 2. Environmental Stress Tolerance Cold Hardiness. Several cooperators are interested in cold hardiness and will be evaluating material in the field at high latitude sites. Most material is too young to be evaluated, but all seedlings planted at Fairbanks, Alaska, were killed in their first winter. In Colorado, Stushnoff (pers. comm. 1999) has screened young seedlings using the following acclimation protocol followed by a laboratory-freezing test (Stushnoff et al. 1983). When seedlings were 3 to 5 cm tall they were exposed to short days (12 h) at 10°C for one week, followed by 2 weeks at 4°C, and an overnight frost of –5°C, and then returned to 10°C day/4°C night with 12 h daylength for one week. The entire flat was then frozen with the temperature decreased at a rate of 2°C/h to –30°C and held for 1 h. The seedlings were then grown in a greenhouse and evaluated for injury by comparing them to ‘Kerr’ open pollinated seedlings. All seedlings with 75 percent or greater die back were considered not hardy. Of 720 seedlings evaluated, 86 were considered hardy in this test and have been retained for field evaluation. Chilling Requirement and Late Bloom. In South Africa, Human (pers. comm. 1999) found that genotypes with shorter chilling requirements are sought for production areas with warm winters in contrast with the need in the central United States for material with a long chilling requirement to provide later bloom to avoid spring frosts. Researchers at these sites have been particularly interested in seedlings germinating

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after long periods in stratification, as this trait has been correlated with long chilling requirement and late bloom (Mehlenbacher and Voordeckers 1991). The chilling required for seed germination varied considerably as reported by the cooperators, ranging from 38 days in Germany to over 200 days in Nova Scotia. Material collected in 1993 may be especially appropriate for this objective since only about 2 percent of the trees in Kazakhstan bore fruit that year by escaping late spring frost. A total of 173 seedlings with multiple disease resistance screened at Cornell University (Aldwinckle et al. 1997; Momol et al. 1997) are under evaluation at the Dawes Arboretum in Newark, Ohio, under the auspices of the Midwest Apple Improvement Association (MAIA). An additional 787 seedlings from the 1995 and 1996 collections with long seed chilling requirement (104 to 127 days, Table 1.15) were sent to the Dawes Arboretum to evaluate late blooming. These seedlings were sent to Dawes after being screened for apple scab and cedar apple rust (Aldwinckle et al. 1997). Drought Tolerance and Sunburn Resistance. In warm, arid production regions such as Washington, British Columbia, and South Africa, cooperators indicated an interest in screening for drought tolerance. Malus sieversii is recognized as a drought tolerant rootstock in Kazakhstan and China. Regions 6, 11, and 12, in particular, have a hot, arid climate with high solar radiation (Table 1.5). Sunburn was surprisingly rare on fruit collected in these areas. 3. Plant Stature. Several cooperators indicated they were interested in obtaining manageable growth habits. Susan Brown in Geneva, New York, is interested specifically in obtaining genetic dwarfs and is evaluating seedlings from three accessions that Kazakh scientists described as genetic dwarfs. In New Zealand, the seedlings in the apple genetics population are rated for tree habit using the IBPGR descriptor and girth is assessed as a measure of vigor. 4. Molecular Genetic Diversity. Studies of molecular genetic diversity in the Central Asian apples at simple sequence repeat sites are continuing at the PGRU as a continuation of previous studies (Lamboy et al. 1996; Hokanson et al. 1998). In an initial study, Lamboy et al. (1996) reported that most allelic variation was among families within collection regions rather than among regions. The main objectives of the ongoing studies are to characterize (1) relative levels of diversity among and within populations, (2) variation between collecting years (1995 and 1996) at the same sites, and (3) diversity among maternal genotypes

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compared with the open-pollinated seedling populations derived from them using leaves and seeds collected in 1996. Several other cooperators plan studies of molecular genetic diversity (Table 1.16). The Oxford University, UK group is investigating the origin and migrations of the apple, including M. sieversii and its ancestors, in a comprehensive program using molecular markers as well as geological, historical, and anthropological approaches (Juniper et al. 1999, Robinson et al. 2001). VII. UTILIZATION The cooperators represent groups interested in cultivar development and genetic diversity. In addition to genebank programs in the United States, Germany, and New Zealand, there are several breeding programs at universities, government agencies, government-held corporations, and consortia of apple growers (Table 1.16). Since most seedlings held by cooperators were not yet fruiting in 2001, evaluation is ongoing and they have not yet been used in breeding. Many indicated their intentions for utilization, however. Nearly all cooperators viewed the M. sieversii germplasm as a means to broaden the genetic diversity in their breeding programs. Many cooperators indicated intentions to use M. sieversii selections in further breeding for rootstocks or scion cultivars. Most cooperators sought: (1) novel fruit quality characters such as color, texture, aroma, and flavor; (2) new sources for disease or pest resistance; and (3) stress tolerance for adaptation to their production regions. In addition, several were seeking easily managed growth habits for scion or rootstock cultivars. The success of utilization of the M. sieversii germplasm in cooperators’ breeding programs will not be known for many years. The ongoing evaluations will ensure that it is tested for critical traits in a large range of apple production regions. We anticipate that this germplasm will ultimately offer useful genetic diversity for several reasons. First, the ecological amplitude of the species in its native habitats is truly impressive. Samples were collected from diverse ecosystems ranging from lush, humid, temperate forests to sparse dry, cold northern forests to xeric, near-desert habitats (Table 1.5). Potential ecotypes from these regions should offer environmental adaptation as rootstock or scion for most apple production regions, except for subtropical areas. Second, M. sieversii in its native habitat has coevolved with several organisms that are pathogenic in orchards. Apple scab and codling moth (Cydia pomonella L.) were especially noted in collection sites. Other organisms such as ubiquitous apple replant pathogens are likely present in the montane apple forests, and natural selection for resistance may have resulted as

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a consequence of forest regeneration. Finally, some M. sieversii genotypes will be readily useful because they are already similar in phenotype to commercial cultivars for some critical horticultural traits. The elite clonal accessions (Plate 1D) and similar seedlings that will be discovered during evaluation may contribute to new cultivars without extensive back crossing. VIII. CONCLUSION Recent collections of wild apple in Central Asia follow the footsteps of early botanical explorers who first documented the geographic extent of the fruit forests of this region. Recognized as an important center of evolution by Vavilov and others, explorations, collections, and study of wild fruit species in the forests of Central Asia continue to this day. Scientists and workers from Russian and Central Asian botanical gardens (formerly within the USSR botanical system) shared their knowledge and experience of wild fruit relatives and led western explorers into their floristically rich forests. In particular, Professor Djangeliev and his associates in Almaty, Kazakhstan, guided the USDA-led teams to populations of apple adapted to various ecosystems in Kazakhstan. These areas are described in detail in the following sections of this volume. In former times, wild species of scientific interest or ornamental/horticultural value were maintained in orchards and gardens. With human population growth increasing stress on natural environments, the need to protect the genetic diversity of wild species from degradation has mounting significance. Central Asian apple genetic diversity is now being maintained within orchards of Central Asia, North America, South Africa, Europe, and New Zealand. In addition, technological advances have allowed preservation by freezing scions and DNA samples. These efforts help insure against future genetic erosion. Characterization of apple selections begun in the orchards and laboratories of Kazakhstan by Professor Djangeliev continue there and internationally. The applications for fruit breeding that result from recent wild fruit explorations in Central Asia demonstrate the synergistic effects of cooperative international endeavors. Distinct contributions by multiple disciplines from various societies and political systems enhance scientific dialogue by bringing together different views concerning genetics, breeding, and conservation. Apple cultivar development will benefit by this work, through the direct introduction of new selections and by introducing disease and insect resistant genes through breeding and gene transfer. The outcome has been cumulative, becoming larger than any single person’s vision.

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Through international agreements and collaboration, we envision apple cultivar improvement, model systems for collecting and preserving the genetic diversity of apple, and a need to develop plans for in situ conservation of these precious genetic resources. Conservation of apple as an “umbrella species” will conserve associated species and provide local economies with many benefits. LITERATURE CITED Aldwinckle, H. S., P. L. Forsline, H. L. Gustafson and S. C. Hokanson. 1997. Evaluation of apple scab resistance of Malus sieversii populations from Central Asia. HortScience 32:440 (Abstr. 075). Barton, D. W. 1975. Preserving genetic diversity. New York Food Life Sci. 9:18. Brooks, H. J., and D. W. Barton. 1977. A plan for national fruit and nut germplasm repositories. HortScience 12:298–300. Brown, A. H. D. 1989. The case for core collections. p. 136–156. In: A. H. D. Brown, O. H. Frankel, D. R. Marshall, and J. T. Williams (eds.). The use of plant genetic resources. Cambridge Univ. Press, Cambridge, UK. Bus, V. G. M., P. A. Alspach, M. E. Hofstee, and L. R. Brewer. 2002. Genetic varichility and preliminary heritability estimates of resistance to scale (Venturia inaequalis) in an Apple Genetics Population. NZJ. of Crop and HortScience 30:83–92. Calhoun, C. L. 1995. Old southern apples. McDonald and Woodward, Blacksburg, VA. Crosby, J. A., J. Janick, P. C. Pecknold, S. S. Korban, P. A. O’Connor, S. M. Ries, J. Goffreda, and A. Voordeckers. 1992. Breeding apples for scab resistance: 1945–1990. Fruit Var. J. 46:145–166. Dickson, E. E. and P. L. Forsline. 1994. Collection of wild apple in middle Asia. Malus 8:11–14. Dickson, E. E., S. Kresovich, and N. F. Weeden. 1991. Isozymes in North American Malus (Rosacea): Hybridization and species differentiation. Syst. Bot. 16:363–375. Dzhangaliev, A. D. 1977. The wild apple tree of Kazakhstan. Nauka Publishing House of Kazakh SSR. Alma Ata, Kazakhstan. (see present volume for English translation) Forsline, P. L. 1987. National Clonal Germplasm Repository for Apples and American Grapes at Geneva, New York. HortScience 22:1073 (Abstr. 272). Forsline, P. L. 1988. Progress in developing a national program for Malus and Vitis germplasm maintenance and evaluation in the USA. EUCARPIA Fruit Breeding Section Meeting, Hradec Kralove, Czechoslovakia. Acta Hort. 224:33–38. Forsline, P. L. 1992. Maintenance, evaluation and distribution of germplasm of apple and grape in the United States National Plant Germplasm System. Proc. of 21st Expert Committee on Plant Gene Resources, Trenton, Ontario, Canada, November 9–10, 1992. p. 84–87. Forsline, P. L.1995. Adding diversity to the National Apple Germplasm Collection; collecting wild apple in Kazakhstan. New York Fruit Quarterly 3:3–6. Forsline, P. L. 1996. Core subsets in the USDA/NPGS with apple as an example. Proc. of the 2nd Workshop on Clonal Genetic Resources, Ottawa, Ont. Canada, January 23–24, 1996. p. 172–175. Forsline, P. L. 2000. Procedures for collection, conservation, evaluation and documentation of germplasm using Malus as an example. Acta Hort. 522:223–234.

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Forsline, P. L., and R. D. Way. 1993. Apple accessions of low priority targeted for removal from the National Plant Germplasm System. Fruit Var. J. 47:204–214. Forsline, P. L., E. E. Dickson, and A. D. Dzhangaliev. 1994. Collection of wild Malus, Vitis and other fruit species genetic resources in Kazakhstan and neighboring republics. HortScience 29:433 (Abstr.). Forsline, P. L., J. R. McFerson, and W. F. Lamboy. 1996b. Optimizing bud harvest time and duration of temporary cold storage for cryopreservation of dormant apple buds. HortScience 31:646 (Abstr. 473). Forsline, P. L., L. E. Towill, J. Waddell, and L. Wiesner. 1995. Development of a base collection of Malus germplasm with cryopreserved buds. HortScience 30:872 (Abstr. 976). Forsline, P. L., L. E. Towill, J. Waddell, and L. Wiesner. 1996a. Il mantenimento delle risore genetiche del melo mediante “crio-conservazione” delle gemme. Rivista di Frutticoltura 11:37–40. Forsline, P. L., J. R. McFerson, W. F. Lamboy, and L. E. Towill. 1999. Development of base and active collections of Malus germplasm with cryopreserved dormant buds. EUCARPIA Fruit Breeding Section Meeting, Oxford, England, September 1–6, 1996. Acta Hort. 484:75–78. Forsline, P. L., C. Stushnoff, L. E. Towill, J. W. Waddell, W. F. Lamboy and J. R. McFerson. 1998. Recovery and longevity of cryopreserved apple buds. J. Am. Soc. Hort. Sci. 123:365–370. Geibel, M., K. J. Dehmer, and P. L. Forsline. 2000. Biological diversity in Malus sieversii populations from Central Asia. Acta Hort. 538:43–49. Grauke, L. J., T. E. Thompson, P. L. Forsline, and K. Hummer. 1995. Use of Core subsets in developing germplasm collections of clonally propagated crops. HortScience 30:907 (Abstr. 1073). Hokanson, S. C., A. K. Szewc-McFadden, W. F. Lamboy, and J. R. McFerson. 1997a. Simple sequence repeat (SSR) variation in a collection of Malus species and hybrids. HortScience 32:440 (Abstr. 077). Hokanson, S. C., J. R. McFerson, P. L. Forsline, W. F. Lamboy, J. J. Luby, A. D. Dzhangaliev, and H. S. Aldwinckle. 1997b. Collecting and managing wild Malus germplasm in its center of diversity. HortScience 32:173–176. Hokanson, S. C., A. K. Szewc-McFadden, W. F. Lamboy, and J. R. McFerson. 1998. Microsatellite (SSR) markers reveal genetic identities, genetic diversity and relationships in a Malus ×domestica Borkh. core subset collection. Theoret. Appl. Genet. 97:671–683. Hokanson, S. C., P. L. Forsline, J. R. McFerson, W. F. Lamboy, H. S. Aldwinckle, J. J. Luby, and A. D. Dzhangaliev. 1999. Ex situ and in situ conservation strategies for wild Malus germplasm in Kazakhstan. Acta Hort. 484:85–91. Hokanson, S. C., A. K. Szewc-McFadden, W. F. Lamboy, and J. R. McFerson. 2001. Microsatellite (SSR) variation in a collection of Malus (apple) species and hybrids. Euphytica 118:281–294. Howell, W. E., G. I. Mink, S. S. Hurtt, J. A. Foster, and J. D. Postman. 1996. Select Malus clones for rapid detection of Apple Stem Grooving Virus. Plant Dis. 80:1200–1202. International Board for Plant Genetic Resources. 1985. Long-term seed storage of major temperate fruits. IBPGR, Rome. Isutsa, D. K., and I. A. Merwin. 2000. Malus germplasm varies in resistance or tolerance to apple replant pathogens in a mixture of New York orchard soils. HortScience 35:262–268. Janick, J. 1974. The apple in Java. HortScience 9:13–15.

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Janick, J. 1989. The national plant germplasm system. Plant Breed. Rev. 7: p. 1–219. Janick, J., J. N. Cummins, S. K. Brown, and M. Hemmat. 1996. Apples, p. 1–76. In: J. Janick and J. Moore (eds.), Fruit breeding. vol II. Tree and tropical fruits. Wiley, New York. Juniper, B. E., R. Watkins, and S. A. Harris. 1999. The origin of the apple. Acta Hort. 484:27–33. Korban, S. S. 1986. Interspecific hybridization in Malus. HortScience 21:41–48. Korban, S. S., and R. M. Skirvin. 1984. Nomenclature of the cultivated apple. HortScience 19:177–180. Kresovich, S., E. E. Dickson, and N. F. Weeden. 1988. Assessment, acquisition, and preservation of the genetic diversity of Malus spp. Genome 30 (suppl):406. Kresovich, S., D. S. Demmin, M. R. Nolan and P. L. Forsline. 1990. Characterization of diversity within the U.S. National Malus collection. Ann. Mtg. Soc. Econ. Bot., Madison, WI. Kresovich, S., W. F. Lamboy, A. M. Szewc-McFadden, J. R. McFerson and P. L. Forsline. 1993. Molecular diagnostics and plant genetic resources conservation. Agr. Biotech News and Inform. 5:255–258. Kresovich, S., W. F. Lamboy, J. R. McFerson, and P. L. Forsline. 1995. Integrating different types of information to develop core collections, with particular reference to Brassica oleraceae and Malus ×domestica. p. 147–154. In: T. Hodgkin, A. H. D. Brown, T. J. L. van Hintum, and E. A. V. Morales (ed.), Core collections of plant genetic resources; IBPGR/CGN/CENARGEN workshop on core collections: improving the management and use of plant germplasm collections. Wiley, New York. Lamb, R. C. 1974. Future germplasm reserves of pome fruits. Fruit Var. J. 28:75–79. Lamboy, W. F., J. Yu, P. L. Forsline, and N. F. Weeden. 1996. Partitioning of allozyme diversity in wild populations of Malus sieversii L. and implications for germplasm collection. J. Am. Soc. Hort. Sci. 121:982–987. V. Langenfelds. 1991. Apple tree systematics. (In Russian) Rija, Zinatne. p. 119–195. Lee, S. B., K. Ko, and H. S. Aldwinckle. 2000. Resistance of selected Malus germplasm to Rosellinia necatrix. Fruit Var. J. 54:219–228. Li, Yu-nong. 1989. An investigation of the genetic center of Malus pumila and Malus in the world. (In Chinese) Acta Hort. Sinica 16:101–108. Li, Yu-nong. 1996. A critical review of the species and the classification of genus Malus Mill. in the world. (In Chinese) J. of Fruit Science 13:63–81. Luby, J., P. L. Forsline, H. S. Aldwinckle, V. Bus, and M. Geibel. 2001. Silk road apples— Collection, evaluation, and utilization of Malus sieversii from Central Asia. Workshop 11 “Collection, utilization, and preservation of fruit crop genetic resources—Some case studies.” HortScience 36:225–231. Mehlenbacher, S. A. and A. M. Voordeckers. 1991. Relationship of flowering time, rate of seed germination, and time of leaf budbreak and usefulness in selecting for lateflowering apples. J. Am. Soc. Hort. Sci. 116:565–568. Momol, M. T., W. F. Lamboy, P. L. Forsline, and H. S. Aldwinckle. 1997. Evaluation of fire blight resistance of Malus sieversii populations from Central Asia. HortScience 32:467 (Abstr. 234). Momol, M. T., P. L. Forsline, H. S. Aldwinckle, and W. F. Lamboy. 1999. Fire blight resistance and horticultural evaluation of wild Malus populations from Central Asia. Acta Hort. 489:229–234. Morgan, J., and A. Richards. 1993. The book of apples. Ebury Press, London, UK. Noiton, D., and P. Alspach. 1996. Founding clones, inbreeding, coancestry, and status number of modern apple cultivars. J. Am. Soc. Hort. Sci. 121:773–782. Noiton D., and C. G. A. Shelbourne. 1992. Quantitative genetics in an apple breeding strategy. Euphytica 60:213–219.

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Norelli, J. L., and H. S. Aldwinckle. 2000. Transgenic resistant varieties and rootstocks resistant to fire blight. p. 275–292. In: J. L. Vanneste (ed.), Fire blight: The disease and its causative agent Erwinia amylovora. CABI Publishing, New York. Ponomarenko, V. 1987. History of Malus domestica Borkh., origin and evolution. (In Russian) Bot. J. USSR. 176:10–18. Ponomarenko, V. 1992. Critical review of the system of the genus Malus Mill. (Rosaceae) species. In: Bulletin of applied botany, genetics and plant breeding, Vol. 146. Russian Acad. Agric. Sci. p. 1–10. Robinson, J. P., S. A. Harris, and B. E. Juniper. 2001. Taxonomy of the genus Malus Mill. (Rosaceae) with emphasis on the cultivated apple, Malus domestica Borkh. Plant Syst. Evol. 226:35–58. Sakai, A. 1960. Survival of the twigs of woody plants at –196°C. Nature 185:393–394. Sakai, A., and Y. Nishiyama. 1978. Cryopreservation of winter vegetative buds of hardy fruit trees in liquid nitrogen. HortScience 13:225–227. Smith, M. G. 1971. National apple register of the United Kingdom. London, Ministry of Agriculture, Fisheries and Food. Stushnoff, C. 1991. Cryopreservation of fruit crop genetic resources—implications for maintenance and diversity during conservation. HortScience 26:518–522. Stushnoff, C., O. Junttila, A. Kaurin, A. T. Ward, and N. Tyler. 1983. Screening for frost hardiness in ten-week-old apple seedlings. Rep. Proc. West. Can. Soc. Hort. Banff. 39:82–85. Szewc-McFadden, A. K., C. G. Alpha, S. M. Bliek, W. F. Lamboy, and J. R. McFerson. 1995. Identification of simple sequence repeats (SSRs) in Malus (apple). HortScience 30:855 (Abstr. 918). Szewc-McFadden, A. K., W. F. Lamboy, and J. R. McFerson. 1996. Utilization of identified simple sequence repeats (SSRs) in Malus ×domestica for germplasm characterization. HortScience 31:619 (Abstr. 312). Tyler, N. J., C. Stushnoff, and L. V. Gusta. 1988. Freezing of water in dormant vegetative apple buds in relation to cryopreservation. Plant Physiol. 87:201–205. USDA, ARS, National Genetic Resources Program. Germplasm Resources Information Network—(GRIN). 2000. [Online Database] National Germplasm Resources Laboratory, Beltsville, MD. Vavilov, N. I. 1930. Five continents. Trans. by Doris Love (1997). IPGRI/VIR, Italy, Rome. Wagner, I., and N. F. Weeden. 2000. Isozymes in Malus sylvestris, Malus ×domestica and in related Malus species. Acta Hort. 538:51–56. Way, R. D. 1976. The largest apple variety collection in the United States. New York’s Food Life Sci. 9:11–13. Way, R. D., H. S. Aldwinckle, R. C. Lamb, A. Rejman, S. Sansavini, T. Shen, R. Watkins, M. N. Westwood, and Y. Yoshida. 1990. Apples. (Malus) p. 3–62. In: J. N. Moore and J. R. Ballington Jr., (eds.), Genetic resources of temperate fruit and nut crops. Acta Hort 290. Wearing, C. H., and K. Colhoun. 1999. Bioassays for measuring the development of different apple cultivars to the development of leafrollers (Lepidoptera: Tortricidae). New Zealand J. Crop Hort. Sci. 27:91–99. Weeden, N. F., and R. C. Lamb. 1985. Identification of apple cultivars by isozyme phenotypes. J. Am. Soc. Hort. Sci. 110:509–515. Yü, D. 1979. Taxonomy of fruit trees in China. p. 1–55. Pome Fruits. No. 1 Malus. Agricultural Publishing House, Beijing. (English translation; translated by Geti Saad for OICD, ARS, USDA).

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2 The Wild Apple Tree of Kazakhstan* A. D. Dzhangaliev** Kazakhstan Academy of Science Interbranch Laboratory for the Protection of Germplasm Main Botanical Garden Almaty, Republic of Kazakhstan

I. INTRODUCTION II. HISTORICAL REVIEW A. The Genus Malus B. History of the Wild Apple of Kazakhstan III. THE ROLE OF WILD APPLE IN THE VEGETATIVE COVER OF MOUNTAIN REGIONS IN KAZAKHSTAN A. Physical and Geographical Assay of Apple Forest Distribution 1. Tarbagatai 2. Dzhungarskei Alatau 3. Zailijskei Alatau 4. Kirghizskei 5. Karatau 6. Talasskei *Originally translated from Russian by I. N. Rutkovskaya. Funded by U.S. Department of Agriculture, Agricultural Research Service. **I express my gratitude to my American colleagues from the U.S. Department of Agriculture, Agricultural Research Service and especially to the late Calvin R. Sperling, R. Soper, E. Rosenquist, and Joseph R. Ellis for providing me with the opportunity to continue the investigations on the genetic resources of apple forests of Kazakhstan. I gratefully acknowledge the American scientists and participants of our joint expeditions into the forests of the Kazakhstan mountain ranges under difficult conditions. These include Philip L. Forsline, the leader of the American group, Herb S. Aldwinckle, Gaylord I. Mink, Elizabeth E. Dickson, James J. Luby, Harold Pellett, Stan C. Hokanson, Thomas K. Unruh, Maxime M. Thompson, and David W. Ramming. The success of our investigations was greatly enhanced by a special visit to Almaty of Frank Browning, a journalist of National Public Radio.

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A. DZHANGALIEV B. Features of Forests 1. Hydrography 2. Climate 3. Soil C. Wild Apple Role in Plant Cover of the Mountain System 1. Tarbagatai 2. Dzhungarskei Alatau 3. Zailijskei Alatau 4. Kirghizskei Alatau 5. Karatau 6. Talasskei Alatau D. Classification of Wild Apples in Kazakhstan 1. Very Dry Growth Conditions 2. Dry Growth Conditions 3. Semi-moist Growth Conditions on Slopes 4. Semi-moist Growth Conditions Along Valleys 5. Moist Growth Conditions THE INFLUENCE OF WILD APPLES ON THE STRUCTURE OF THE ENVIRONMENT A. Influence of Apple Stands on Microclimate and Apple Response to Vertical Zone Conditions 1. Precipitation 2. Soil Temperature 3. Air Temperature 4. Air Humidity 5. Wind 6. Apple Growth and Development in Relation to Meteorological Conditions 7. Climatic Phase Changes as Affected by Different Altitude Zones 8. Setting Stages and Differentiation of Apple Flower Buds 9. Evaluation of Thermal Resources of the Territory B. Dependence of Apple Stands on Soil Conditions 1. Chemical and Water-holding Properties of Soil and Their Changes Under the Influence of Apple Forests 2. Accumulation and Ash Composition of Leaf Fall and Litter in Apple Forests CHARACTERISTICS OF WILD APPLE GROWTH AND DEVELOPMENT A. Natural Renewal 1. Vegetative Renewal 2. Renewal by Seed 3. The Vitality of Forest Stands from Vegetative Origin B. The Influence of Tree Thinning and Apple Cultivation on Apple Renewal and Stand Preservation C. Growth and Development of Wild Apple Trees in Relation to Natural Renewal D. Structural Features of Apple Root Systems in Relation to Growth Conditions E. Growth, Productivity, and Fruit Bearing of Wild Apple INTRASPECIFIC POLYMORPHISM OF WILD APPLE UTILITY AND BIOCHEMICAL CHARACTERIZATION OF WILD APPLE FRUIT A. Composition of Apple Fruit Forms with Different Ripening Dates B. Composition of Wild Apple Fruits in Relation to Environmental Growing Conditions

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Composition of Wild Apple Fruits That Have Different Flavor Types Processing Fruit Characteristics of Promising Wild Apple Forms Changes in the Chemical Composition of Wild Apples During Storage Processing Characteristics of Wild Apple Fruits 1. Cider 2. Calvados (Hard Cider) 3. Natural Juices and Juices with Sugar 4. Canned Fruits VIII. PRESERVATION OF WILD APPLES IX. CONCLUSION LITERATURE CITED

I. INTRODUCTION The conditions leading to the development of our work on wild apples are the natural and historical settings of the Republic of Kazakhstan. The enormous territory of Kazakhstan is situated in the center of Eurasia within an area of 272 million hectares. Kazakhstan spreads from the west to the east for 2925 km (from the Lower Volga to the Altai and China), and from the north to the south for 1500 km (from the Western Siberian Plain and the Southern Urals to the Tien Shan ridges and the Kizil-Kum deserts). Within this territory almost all types of landscape of the terrestrial globe are present, from dry subtropics and hot deserts to highmountain cold tundra and glaciers. Wild apple trees, as forest-forming and forest-constituting woody plants, have since antiquity occupied considerable area here. A great variety of ecological conditions exist in these regions. The wide range of geographical latitudes and longitudes, and, the variable altitudes and natural barriers in the mountain regions led to the isolation of populations and the separation of interpopulation connections of apple forests and caused intraspecific types to evolve. Consequently, Kazakhstan is one of the most suitable countries for projects aimed at discovering high-quality germplasm for apple resistances. Great value for scientific and practical projects exists because Kazakhstan is the original genetic center of biological variability of wild apples and historically, has formed their rich gene pool. Here, the wild apple formed as a species with numerous subspecies before wider distribution via cultivation. Wild apple species in Kazakhstan, genetically kindred to cultivated varieties of the world, have always grown here and form natural apple forests on specific altitude zones of the mountains. The long-term expeditionary and experimental investigations of these forests, conducted by

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the author, indicate the enormous possibilities of selection in wild apple forests of Kazakhstan. First, natural hybrids of wild apple (economically valuable phenotypes) may be used directly in cultivated orchards as stress resistant cultivars, having good pomological qualities that are superior to cultivated varieties. Since olden times, the best wild apple forms are grown in local orchards. Especially important are the wild apple forms selected for growing raw material in orchards for the manufacture of high-quality products for the food industry and for the production of cider. The products of these wild apple forms greatly exceed those from cultivated varieties and are not inferior to the best French cider cultivars. Second, wild apple phenotypes having many different important qualities should be considered as carriers of genes for selection. Previously, wild apple species did not much attract the attention of botanists. The genes of intraspecific phenotypes were not taken into account during the choice of parents for crossing in the breeding of new cultivars. This omission may be explained by the fact that there have been no summarized investigations on the importance of wild species polymorphism for selection. Generally, pomologists seek resistant parents for crossing that are found within the world collections of cultivated varieties, although many of them have undergone genetic erosion. In order to increase disease or stress resistance of new cultivars, they resort to an accidental collection of wild forms with unknown characters. Many have noticed that in the resistant hybrids, wild features such as small fruits and the presence of numerous spines also dominate (Michurin 1948). These facts testify to the lack of scientific information about the individual peculiarities of intraspecific wild species. As our investigations show, there is a wide range in phenotypes of individuals that develop in natural apple forests, including trees with large fruits and without numerous spines. Wild apples are distributed in many countries of the world, but until recently, the investigators of the flora have paid little attention to the variability within and among species, and to the range of their ecosystems. It became evident to us that the intraspecific polymorphism of the wild apple species of Kazakhstan could not be revealed by merely using classical comparative morphology. For this reason, field investigations included a genetic inventory and taxonomy of apple forests according to the phenotypic characteristics of the populations formed in nature as well as selection. Particular individual phenotypes emerged in the forests that were selected according to economically valuable characters and were tested under cultivated conditions. The wild apple forests of

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Kazakhstan are characterized as new material for breeding and selection. Special attention is devoted to the discovery of intraspecific polymorphism of wild apples. By their natures, the mountain systems of Kazakhstan are unique evolution centers of singular importance for wild fruit plants. Here, 130 wild species of edible pome fruits, stone fruits, walnut, small fruits, and grape grow. According to the extent of area occupied, among the most important of these is the wild apple. Also important are the apple’s forest-forming capabilities, its functions in nature, its richness of biological resources, and especially, the wide variety and quality of its gene pool. Modern apple forests of Kazakhstan are referred to as relictual forest types, formed as a result of successive development of tertiary vegetation, giving evidence of the unique characteristics of their gene pool for self-regulation and self-reproduction during a long evolutionary period. These forests, in spite of the historical remoteness from their origin, preserve their reproductive abilities as a self-developing, natural biological system in their main areas which are little affected by the economic activity of man. N. I. Vavilov (1931), the founder of the concept of World Gene Centers of Cultivated Plants, who visited wild apple forests of Kazakhstan around Alma-Ata in September 1, 1929, wrote Alma-Ata translated means ‘Father of apples.’ Around the town and at a great distance along the mountain slopes, the overgrowths of wild apples are spread, creating massive forests here. . . . It is possible to see with our own eyes that we are in the remarkable center of origin of the cultivated apple.

The results of my investigations and participating scientists of the joint Kazakhstan–American expeditions testify to the fact that in one and the same population, apple species differentiate into various and qualitatively different tree phenotypes, sharply distinguished among themselves in biological and many economically important characters. The selected tree phenotypes of wild apple differ among themselves in the duration of vegetation, the time of flowering, quantity and quality of fruits, ripening periods, biochemical composition of fruits, tree structure, winter hardiness, and immunity from diseases. Widely spread spontaneous hybridization in heterogeneous apple forest populations in a variety of backgrounds and under fluctuating mountainous conditions played a decisive evolutionary role in the formative

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processes in these forests. New population characteristics stemming from the enormous variability of intraspecific apples have resulted from mutation, genetic recombination, and natural selection. Polymorphism, the presence of several clearly distinctive wild apple forms within a single species, has been steadily maintained in the population by variable habitats and the tendency for the development of new groups to form in this habitat. Hybridization used by breeders to select new cultivars essentially mimics the evolutionary process normally occurring in nature. However, nature spontaneously formed polyploid species with colossal intraspecific variety, which presaged the success of domestication that occurred many millennia ago. Therefore, our project on intraspecific variability had the objective not only of direct phenotype selection but also to gain knowledge of the regularity of form, species formation, and species evolution. Selected tree phenotypes in cultivation were found to have had engrained in them individual characters seen in natural populations. These characteristics were revealed as a normal response under new growing conditions and persisted in hybrids with cultivated varieties. These phenotype traits have a genetic basis and are not modifications arising under the influence of habitat or the changed conditions of environment in cultivated plots. The phenotypes tested under different ecological conditions exhibited greater flexibility than cultivated apple, which did not withstand the severe trial conditions and were dropped out of the experiment. This suggests that the genetic erosion that occurs with cultivated varieties may be overcome by using wild apple germplasm. Artificial selection of economically valuable apple forms among numerous phenotype variants in the original Kazakhstan gene center, where their endemic structural gene pool was historically established, produced practical results. On the basis of the best phenotypes, we selected unique genotypes from all parts of apple forest areas and established a map on a scale of 1:5000 showing their locations. These selections in nature (in situ) will serve as a seed and clone base for economically valuable phenotypes for forest plantations and the international gene pool exchange. Besides that, collections of many valuable phenotypes were grafted in experimental plantings (ex situ) for further study and for possible introduction. We also prepared summary maps of wild walnuts and small fruit plants of the mountain ranges of the Republic on a scale of 1:50,000, where numerous wild drug, edible and other grasses, woody plants, and shrubs are found.

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II. HISTORICAL REVIEW A. The Genus Malus Since antiquity apple has attracted the attention of a wide range of naturalists, from nature lovers to professional botanists, horticulturalists, and pomologists. It continues to be an important subject of various investigations in the field of the theoretical and applied botany. The great systematist Carolus Linnaeus (1767) combined pear, apple, and quince in the genus Pyrus. He proposed the name Pyrus malus for cultivated apple, and that wild apples should be referred to as species of the main genus Pyrus (e.g., P. baccata). Considering the fruit as a distinctive character, the botanist and fruit grower Philip Miller (1768) separated apple from the genus Pyrus and described it as having the characteristics of the genus Malus. Recognized as the taxonomic author for apple, Miller wrote: Apple should be distinguished as a separate genus and this difference is included in the Nature itself of these plants, as they are incompatible on budding and whip grafting. Therefore, I separate apple from pear as it was always done by the botanists before Linnaeus.

Subsequently, many outstanding plant systematists have verified the separation of genus Malus. Koehne (1893) and Zabel (1903) subsequently classified Malus into selections. Having assumed calyx lobe deciduousness as a basic species character within the genus, Koehne divided the genus Malus into two sections: Calycomeles Koehne, apples with persistent calyx lobes; and Gymnomeles Koehne, apples with deciduous calyx lobes after blooming. He also separated an intermediate hybrid section Calyomeles ×Gymnomeles. Zabel employed simple or lobed leaves as the basis of his system and divided genus Malus into two subgenera: Eumalus Zabel, true apples with simple leaves, and Sorbomalus Zabel, mountain ash apples with lobed leaves. He referred to the hybrid species Eumalus ×Sorbomalus as a special group. These two types of divisions of the genus Malus are the foundations of studies by many modern systematists. Schneider (1906a,b) used Zabel’s system and separated species into two sections of the genus: (1) with deciduous calyx lobes, and (2) with persistent calyx lobes. But this is not quite logical, because these groups fully characterize the whole complex

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of the species. In Rehder’s (1915) system there were actually five sections of the genus Malus: (1) Eumalus Zab.; (2) Sorbomalus Zab.; (3) Chloromeles Rehd.; (4) Eriolobus Schneid.; and (5) Docyniopsis Schneid. There are also a number of shortcomings to his system. For example, two of the subgroups, Baccatae Rehd. and Pumilae Rehd., were united into the section Eumalus, that is, large-fruited apples of the M. domestica type and smallfruited apples of the M. baccata type were both included in the section. The Japanese botanist Koidzumi (1934) somewhat revised Koehne’s system. Apple species growing in the USSR were described by S. V. Yuzepchuk (1939) in the book Flora of the USSR and later on by A. A. Fedorov and O. M. Poletiko (1954). An interesting system proposed by F. D. Likhonos (1964) further revised both Koehne’s and Koidzumi’s systems. The Soviet monographer V. T. Langenfeld (1970) proposed a very clear and harmonious system of the genus Malus and identified six sections: (1) Eriolobus, downy-lobed apples; (2) Docyniopsis, docynious apples; (3) Sorbomalus, mountain-ash apples; (4) Chloromeles, greenfruited apples; (5) Gymnomeles, berry apples; and (6) Malus, true apples. More recently Korban and Skirvin (1983) determined that the appropriate binomial for cultivated apple should be Malus ×domestica. The initial period in this wild apple study was characterized by geobotanical investigations and data accumulation about the main representatives of the genus Malus. The works of P. S. Pallas (1773), W. Aiton (1789), C. G. Willdenov (1794), A. Michaux (1803), M. B. Borkhausen (1803), P. W. Watson (1825), E. Regel (1859), D. Brandis (1874), Th. Wenzig (1874), C. S. Sargent (1880, 1888, 1894), A. Batalin (1893), J. Matsumura (1899), C. K. Schneider (1906a), T. Kawakami (1911), A. Rehder (1915, 1933, 1939), T. Nakai (1916), A. N. Uglitskikh (1932), G. P. Viktorovsky (1935), S. V. Yuzepchuk (1939), G. G. Tarasenko (1940), G. N. Sumnevich (1942, 1948), Al. A. Fedorov and M. A. Fedorov (1949), S. N. Kudryashev (1950), P. P. Polyakov (1950), I. T. Vasilichenko (1952, 1963), and V. V. Ponomarenko (1972, 1973, 1975) were devoted to the study of wild apple species and their biology, emphasizing morphological and botanical classifications. As a result of their investigations 50 species of wild apples were identified and described, among them: 14 to 15 in China, 13 to 18 in USSR, 3 in Europe, 3 to 4 in Middle Asia, 3 to 4 in Himalaya, 5 in Japan, and 9 in North America. It is necessary to note that the descriptions of new apple species given by different authors were not well grounded; they greatly differed in names and species numbers. For example, Koehne (1893) designated 23 apple species, 10 of which were of hybrid origin; while the total number of apple species given by Zabel (1903) was 30.

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Schneider (1906a) described 18 species, 11 botanical varieties, 1 form, and 13 hybrids of the Malus genus. Rehder (1949) divided the genus into 25 species, and Koidzumi (1934), into 36 species. Van Eseltine (1933) described 9 apple species for North America. S. V. Yuzepchuk (1939) distinguished 12 apple species, growing in the USSR. A. A. Fedorov and O. M. Poletiko (1954) listed 37 species, spreading naturally and in culture in the territory of the USSR. In the first botanical descriptions in the literature, several different names were assigned to a single wild apple species. So, the rare wild apple species M. trilobata (Dc.) Boiss., because of its deeply trilobate leaves, was even described as a different genus by some botanists. Boissier (1872) gave a short description of this species under the name Sorbus trilobata, but Schneider (1906b) assigned this apple to genus Eriolobus. The American apple, M. ioensis (Wood) Britt., was described by Wood (1872) as a variety, Pyrus coronaria var. ioensis, then later it was separated as an independent species by Britton and Brown (1913). A hupehensis apple was simultaneously described by two authors under different names: P. hupehensis (Pampanini 1910) and M. theifera (M. hupehensis) (Rehder 1915). There were many arguments against the names assigned to species and varieties of wild apples. So, S. V. Yuzepchuck (1939) separated four species in Siberia: (1) M. baccata (L.) Borkh.; (2) M. manshurica (Maxim.) Kom.; (3) M. sachalinensis Yuz.; and (4) M. pallasiana Yuz. R. Ya. Kordon (1956) considered such species separation as irrational, because the new species identified by S. V. Yuzepchuk did not possess distinctive species characters and their locations were not widely spread and isolated. A detailed comparison of M. sachalinensis and M. manshurica allowed V. V. Ponomarenko (1973) to identify them as subspecies, that is, M. manshurica sp. sachalinensis (Yuz.) Ponomarenko stat. nov. The investigators paid great attention to Malus baccata, which has a large number of different forms growing across a wide area: from Lake Baikal to the Pacific Ocean. On the border between Manchuria and Himalaya, the deviation from the species type was most clearly expressed. It made it possible for Maximowicz (1874) to designate two additional species: M. sibirica and M. manshurica. V. I. Komarova (1959) (quoted from Cordon 1956) labeled them as apple species. V. V. Ponomarenko (1972) pointed to the independence of these two species, different by their morphological, biological and ecological characters, separated rather well geographically and having their own growing areas. There is great disagreement about the species of wild apple in the Caucasus. Borkhausen (1803) described one under the name M. dasyphylla

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Borkh. Then, A. N. Uglitskikh (1932) designated two species, M. orientalis Uglitskikh. and M. montana. V. V. Pashkevich (1929) described three apple species in the Caucasus: M. sylvestris Mill., M. pumula Mill., and M. dasyphylla. E. A. Syubarov (1968) considered M. dasyphylla and M. orientalis to be identical. S. V. Yuzepchuk (1939), A. A. Grossgeim (1948), A. A. Fedorov and O. M. Poletiko (1954), and I. T. Vasilichenko (1963) pointed to only one wild apple for the Caucasus, M. orientalis. V. T. Langenfeld (1970), besides the typical M. orientalis, separated a subspecies M. orientalis sp. montana (Uglitzk.). V. V. Ponomarenko (1975) assigned the sub-alpine variety, M. orientalis var. subalpina Ponom. V. F. Vasilyev (1932), G. V. Trusevich (1936), and P. G. Sorokin (1947) contended that only one apple species, M. pumula, was growing in the Crimea. A. G. Iliena (1959) determined that there are two ecological types, M. orientalis and M. praecox (Pall.) Borkh. Such contradictions may be explained by the exceptional level of wild apple polymorphism, which leads to a great resemblance among species morphology and difficulty in differentiation among them. Many investigations were conducted to study wild apple forms, but until now there is no common opinion concerning the multiple forms within a species’ composition. V. V. Pashkevich and A. P. Sigov (1928) were surprised at the high quality of apple fruits of the Pyatirechje and tried to find an explanation for this fact. They came to a rather original, but far from correct, conclusion that local apple forests were the results of the action of military “culturtregers.” M. G. Popov and G. M. Popova (1925) and A. V. Vasilyev (1938) explained apple polymorphism by citing different local environments and hybridization. It is possible that the crossing of wild apple forms with cultivated varieties of apple took place somewhere, but it was only of a local and private character, and was never widely spread. Most investigators are inclined to explain the extraordinary polymorphism of wild apples by different ecological and environmental growing conditions of the apple tree, by interspecies and intraspecific hybridization, and by accidental seed introduction from cultivated apple varieties brought into local species areas. The characteristics of M. sylvestris in the European part of the USSR were studied by F. S. Salynskij (1937), F. F. Kirillov (1937), F. I. Pekhoto and V. V. Malychenko (1966). These authors noted the high variability of the growth characters, time of ripening, fruit flavor, and eating quality of M. sylvestris. Blaja and Ivan (1960) revealed a rather prominent polymorphism of M. sylvestris. Malus praecox, a xerophytic race of M. sylvestris, differing from it mainly by much deeper pubescent leaves and less growth, was found by N. Stoyanov and B. Stefanov (1933) in the

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forests of the Balkan Peninsula. Fischer and Schmidt (1938) described the many forms of M. sylvestris in different areas of its habitat. The characteristics of M. orientalis were studied by N. A. Troitskij (1930), A. V. Vasilyev (1938), V. A. Khetagurov (1956), P. P. Usatyj (1959), I. T. Vasilichenko (1959), and V. V. Ponomarenko (1975). They identified a number of forms, which differed by fruit appearance, form, size, color, flavor qualities, and ripening stages. A. D. Voejkov (1909) singled out 20 forms of M. sibirica. N. I. Kichunov (1937) described three forms, Siberian giant, Zabajkalskaya polukustovaya, and Tipichnaya melkoplodnaya. P. A. Zhavoronkov (1938), when studying M. sibirica forms, gave special emphasis to their winter hardiness, resistance to pests and diseases, fruit size, flavor, color, fruit shape, and ripening period. Foreign scientists, including Regel (1862), Lauche (1880), Schneider (1906b), Makino (1931), Seeliger (1934), and Koidzumi (1934) also studied the polymorphism of this apple species in various growing areas. Wild apple multiformity in Middle Asia was noted by M. G. Popov (1925): The quantity of forms, clearly different, is indeed tremendous here, so it will not be an exaggeration to say, that each apple tree may be described as a new form. (p. 100)

He differentiated the forms based on their morphological fruit characters (fruit stalk, receptacle, fruit flesh, flavor, aroma, and ripening period). The intraspecific diversity of M. hissarica S. Kudr. (12 botanical varieties) was studied by I. T. Vasilichenko (1948, 1952). Al. A. Fedorov and M. A. Fedorov (1949) described the polymorphism of Middle Asian apple species. According to their data, a great number of apple forms was observed in the area of West Tien Shan. They studied the variability of fruit form, size, weight, color, fruit flesh consistency, fruit stalk length, shape and arrangement of seed cavities, shape of seeds, and calyx in M. kirghisorum Al. et An. Theod. Varieties and forms of American apples were studied by Sargent (1905, 1913, 1922), Britton (1905), Small (1933), and Rehder (1949). Especially notable were a great number of winter hardy apple forms in M. fusca (Raf.) C. K. Schneid., with an ability to withstand low temperatures, spreading from Alaska and the Aleutian Islands to California. In 1931 Makino and Nemoto separated three forms (f. lutea asami, f. rubra asami, and f. Yammoto asami) and one botanical variety (rinki

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asami) of the species M. asiatica Nakai., a wild Korean and Chinese species (Seeliger 1934). The importance and meaning of the dynamic origins of modern cultivated fruit cultivars from wild apple forms was presented in the scientific works of N. I. Vavilov (1931). Vavilov showed that wild fruit trees in the centers of their origin possessed wonderful polymorphism, but at the periphery of these areas, the multiformity and the number of dominant characters sharply decreased. N. I. Vavilov wrote: In the Caucasus and Middle Asia, it is possible to retrace step by step all of the stages of evolution. The so-called Circassian apple varieties, bred by ancient cultivators of the Caucasus, represent a kind of transition from wild forms to cultivated forms, distinguished by small fruits, great resistance to cold and damages, and sometimes by early ripening. Their fruits are often notable for their long storage and undamaged transportation. Among them one can find the high-quality varieties. Such “transits” from wild apples to cultivated ones were also noticed by Semirechje (p. 94). Factors of fruit tree introduction into culture and all stages of fruit growing evolution in the middle zone of the European part of the USSR and the Caucasus were presented by V. V. Pashkevich (1929).

Most investigators (Voejkov 1909; Pashkevich 1938; Skibinskaya 1951; Kordon and Pekhoto 1962; Zhukovsky 1971) consider the typical M. sylvestris and its subspecies M. praecox, to be the ancestor species for a large number of middle Russian apple cultivars. According to P. Z. Vinogradova-Nikitina (1929), A. V. Vasilyev (1938), N. V. Kovalev (1941), L. I. Prilipko (1954), and V. A. Khetagurov (1958), M. orientalis is the ancestor species of many local cultivated apple varieties in the Caucasus and in adjacent regions (e.g., Kekhura, Kitra, Abilaury). The Babaarabskaya apple, one of the forms of M. turkmenorum Yuz. et M. Pop., is widely distributed in apple orchards of the Ashkhabad region and is mainly represented by four cultivars: Turshy, Okcha, Juvan, and Kizilchi (Ostroukhova 1961). E. D. Taranova’s (1961) investigations on the origin of 13 apple cultivars and 1100 of their hybrid seedlings are interesting. On the basis of their morphological characters of vegetative organs, biological and sometimes physiological peculiarities, she found that they were very similar to the morphological characters of wild species, consisting of a form complex of the species M. pumila. Malus pumila, imported from Asia, is regarded to be the ancestor of all species, varieties, forms, and apple sorts growing at present in England (Brimble 1946) and the cider and table varieties in France (Cheva-

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lier 1953). Malus baccata is the ancestor of American crabs and of the cultivated forms on Formosa Island, M. formosana Kaw. et Koidz. (Pashkevich 1929). In China since antiquity, the apple assortment began to be created due to the introduction in culture of the best forms of M. asiatica, M. pumila, and others (Dragavtsev 1966). Thus, the wild apples group with their species and form variability undoubtedly may serve as a rich source for selection. The most valuable material for producing cultivated apple varieties with high winter hardiness is M. sibirica. The selection work with M. baccata, conducted by I. V. Michurin (1948), P. A. Zhavoronkov (1938), and other investigators, confirmed the high winter resistance of hybrids. Significant selection with M. baccata has been carried out in North America and Canada (Hansen 1924, 1929; Saunders 1896). M. prunifolia (Willd.) Borkh. was used in breeding new apple cultivars by I. V. Michurin (1948) and Saunders (1911). M. niedzwetzkyana Dieck., which was highly rated for its ornamental quality, was also used in breeding research (Michurin 1948; Schroder 1954). Research has been devoted to genetic investigations of wild apples. Nebel (1929) and Lincoln and McCann (1937) ascertained that most wild apple species have a somatic chromosome level, 2n = 34, being diploid plants (2x). But, M. coronaria (L.) Mill. and M. glaucescens Rehder are tetraploids 2n = 68, and M. spectabilis (Ait.) Borkh. is triploid 2n = 51. The same results concerning the quantitative content of chromosomes and polyploidy was obtained by Kobel (Knight 1963), Sax (1931, 1959), and Olden (1953). Sax noted that some Asiatic apple species having 17 pairs of chromosomes are diploid, but such apomicts as M. hupenensis (Pamp.) Rehd. and M. toringoides (Rehd.) Hughes are triploid. The crossing of triploid species with diploid apple varieties gave mostly triploid hybrids, but seldom tetraploid. Apomixis was a dominant factor in F1 progeny. Crane and Lawrence (1934) investigated the cytology and genetics of wild species and cultivars of apple. They determined chromosome number in wild apple species and demonstrated the dominant influence of wild apple genes on size, color, flavor, and flesh consistency of hybrid fruits. Anatomical, physiological, and morphological investigations on wild apple species in Germany were carried out for leaf color, leaf lamina, flower stalk length, flower number in the inflorescence, style number in a flower, fruit endocarp structure, and color and weight of fruit seeds (Seeliger 1934). M. halliana (Koehne.) and M. baccata were shown to have very low (0–40) latent cell numbers; M. fusca, M. floribunda, mean density (81–120), and M. coronaria, very high density (161–200).

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Genetic investigations by Henning (1947) showed that M. zumi (Mats.) Rehd., M. coronaria, and M. baccata are homozygous, and M. prunifolia is heterozygous for scab resistance incited by Venturia inaequalis. Malus prunifolia had a dominant influence on M. zumi hybrid progeny. As a result of hybridity analysis made by Williams et al. (1966), the allelic genes for resistance to Venturia inaequalis was found in M. floribunda Seib., M. prunifolia, and M. baccata. For this group of genes, Williams suggested the Vf symbol for M. floribunda. Genetic investigations on wild apple resistance to apple scab have been made (Stevenson and Jones 1953; Dayton 1954; Hunter and Sampson 1958). These studies found that the immunity from wild species was provided by one dominant gene (monogene) in M. floribunda, by a digenic dominant gene in M. micromalus Makino., and by a trigenic dominant gene in M. pumila. The geographical distributions of different wild apple species have been widely studied (Folgner 1897; Hempel and Wilhelm 1889; Collett 1902; Sargent 1905, 1922; Nikiforov 1911; Bailey 1916; Hegi 1919; Komarov 1922; Silva and Schneider 1922; Hayek 1927; Shepetov 1932; Murzaev 1934; Dyagilev 1936; Rubtsov 1940; Unchiev 1947; Parsa 1948; Uvarov 1949; Kabluchko 1950; Vashchenko 1958; Usenko 1953; Dobrochaeva 1954; Kobendza 1955; Sokolov 1955; Peterson 1966; Lisavenko 1959; Fedorov 1959; Trusevich 1962; Salamatov 1965; Syubarov 1963; Likhonos 1964; Aliev 1965; Pisarenko and Drozhalov 1966; Mlokosevich 1962). But there are some disputable and unsettled questions concerning apple distributions. Thus, until now the original habitat of the wild apple M. prunifolia has not been determined. Bailey (1949) and Rehder (1949) considered the plant known under the Japanese name, Rinki (M. prunifolia var. Rinki) and discovered by Wilson in Sichuan (Southwestern China), as a wild Chinese apple. N. I. Kichunov (1937) reported that a Siberian fruit grower M. G. Nikiforov had discovered groves of wild Chinese apples in the Kalgan Mountains in Northern China. According to Chow’s (1934) data, the true wild apple M. prunifolia was distributed in the northern and central regions of China. Schneider (1906b) supposed that M. prunifolia is a hybrid species of M. pumila × M. baccata; but G. G. Tarasenko (1940) believed this to be hardly well-grounded because the species has no natural region of its own and is known only in culture. Many publications have been devoted to the study of the biology, ecological conditions, and associations of wild apple. The research of Ascherson and Graebner (1906–1910), Schiemann (1932), Kobendza (1955) and I. T. Vasilichenko (1963) ascertained that M. sylvestris in Europe is one component of the deciduous forests consisting of beech,

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oak, birch, and hornbeam. M. manshurica in China is a part of the cedar broad leafed and cedar fir (Manchurian) forests, as well as oak woods and shrubs (Vasilichenko 1963; Dragavtsev 1966). The wild apple of the Crimea is mainly incorporated within oak groves (Iliena 1959). According to the data of Al. A. Fedorov and M. A. Fedorov (1949), M. kirghisorum grows along with residual relict walnut forests. I. S. Lupinovich (1947) studied the dependence of nut-apple forest development on hydrothermal conditions. V. I. Zapryagaeva (1964) investigated climate and soils (water ensuring) in broad-leaved forests of Tajikistan. Blaja and Ivan (1960) studied the ecological and biological growing conditions of M. silvestris in Romania. They described forest soils, relief, and climate characteristics (active accumulated temperatures [heat units], precipitation) in the regions of apple distribution. G. V. Trusevich (1938) studied the influence of relief and soil hydrobiological conditions on the stages of development of wild apple. The soils in areas of M. sylvestris distribution were described by A. D. Danilov (1950) and Hadfield (1955); M. orientalis, by V. F. Vasilyev (1932); M. manshurica, by N. V. Kovalev (1936), and N. A. Borodina and V. I. Nekrasov (1966); and M. pumula, by M. G. Popov (1929). Sargent (1922), Hartmann (1929), and Bailey (1949) described the soils of American wild apples. A. I. Olonichenko (1936), V. A. Moryakina (1965), M. N. Salamatov (1965), N. I. Denisov (1973), Saunders (1911), Macoun (1915), and Hansen (1924) studied some ecological and biological characteristics of M. sibirica. The biology and ecology of M. kirghisorum and M. sieversii (Ldb.) M. Roem. were studied by Al. A. Fedorov and M. A. Fedorov (1949) and Vorobieva (1966); M. hissarica, by B. Z. Sabirov (1959) and Kh. Azimov (1973); M. turkmenorum, by S. A. Ostroukhova (1970); M. orientalis, by P. P. Usatyj (1959) and V. V. Ponomarenko (1975). The valuable qualities of M. ioensis such as early ripening, late flowering, escaping from spring frosts, good fruit keeping, dwarfness, and resistance to Erwinia amylovara were discovered through the biological investigations of Hansen (1927) and Fischer and Schmidt (1938). The root systems of the wild apple were examined by Ja. S. Medvedev (1919), T. K. Kvarazkhelia (1927), V. V. Pashkevich (1930), N. K. Vekhov (1932), L. F. Pravdin (1938), A. A. Fedorov et al. (1945), P. K. Krasilnikov (1949), A. P. Dragavzev (1956), and L. A. Apoyan (1964). V. I. Zapryagaeva (1964) who noted the surface distribution of the main root mass of wild apple and their exceptional ability to form root suckers. McClintock (1931) discovered that under the conditions of South America, wild apples do not form lateral roots and usually have one straight long tap root, penetrating deeply into soil. Propagation of wild apple was considered by Al. A. Fedorov and M. A. Fedorov (1949) who noted that

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only M. kirghisorum and M. sieversii were able to propagate by root suckers. N. V. Kovalev and D. S. Tupitsin (1956) believed that this property was possessed by all wild apple species. A number of studies have been devoted to wild apple as rootstocks. Rootstock qualities of different wild apple species appear quite variable. Thus, Hansen (1899) considered M. baccata to be the best rootstock for cultivated apple varieties of North America. Regel (1862) and Kremer and Schultz (1941) noted incompatibility between scion and rootstock. V. V. Pashkevich (1938) recommended M. sylvestris as a rootstock for northern regions of the European part of the USSR, while V. I. Budagovskij (1949) noted insufficient winter and drought resistances of its seedlings. N. I. Kichunov (1937), indicating the positive features of M. prunifolia (fibrous root system, unaffectedness to soil conditions) and recommended it as an apple rootstock. A. I. Zhuravskij (1914) and G. V. Trusevich (1947), taking into account the weak growth and numerous spines of M. orientalis considered it unacceptable as a rootstock. However, I. V. Vasilichenko (1963) bearing in mind the grafting method of the Majkop Experimental Station and of the local population, pointed to M. orientalis as a rootstock for cultivated apple. Many studies have been devoted to the problem of apple population utilization. Recommendations on wild apple management were given by G. V. Trusevich (1938), G. I. Monashev (1947), P. P. Kostyk (1950), L. P. Savchenko (1957) and Ju. E. Pulko (1965). The authors A. G. Araratyan (1940), Ju. E. Pukhtinsky (1948), N. F. Batygin (1950), G. A. Sokolov (1955), E. I. Kharchenko (1952), Kh. Z. Gubajdullin and V. I. Dyachkova (1953), M. A. Fedorov (1959), B. Z. Sabirov (1958), P. P. Usatyj (1959), S. N. Krajnov (1961), A. G. Gusejnov et al. (1964), A. I. Pisarenko and M. N. Drozhalov (1966), and A. S. Abeev and N. T. Bondarev (1967) wrote about cultivation of apple forests. In these authors’ opinion, the best method of cultivating wild apple stands in forest orchards is regrafting, that is, using them as rootstocks for cultivated apple grafted right in the forest. Regrafting accelerates the time of apple variety bearing, increases their longevity, yield, resistance to pests, diseases, and unfavorable environments and above all, enhances fruit quality because domesticated fruits significantly exceed wild apples in flavor. But the method of mass grafting of wild apple trees did not prove to be successful, the orchards were not developed, and the wild fruit stands greatly declined (Avsaragov 1965; Vasilichenko 1971). There are some studies on the economics of wild apple use. P. S. Pallas (1786) wrote about the use of boiled and baked wild apples as well as their use for medical purposes. There are many instructions on using wild apple fruits for producing high-quality juices and apple sauce

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(Dobrin 1931; Dyrdin 1933; Kirillov 1937; Danilov 1950; Demyanenko 1951; Sokolov 1955; Fedorov 1959; Bogoyavlenskaya 1962; Tkhagushev 1962; and Dolidze et al. 1966); for production of syrup and jam (Fedorov 1959; Zapryagaeva 1964); for production of jam and stewed fruits (Savitskij 1938; Sumnevich 1942; Bezzubov 1949; Komarov 1953; Usenko 1953; Nikitin 1957; Vashchenko 1958; Khegaturov 1958); and for pastille production (Grzhivo 1932; Komarov 1953; Sokolov 1955; Osadchij 1956; Vaschenko 1958). Fruits of wild apples are widely used in the confectionery industry for marmalade production (Sorokin 1947) and sweets filling (Maslovsky and Dombkovskaya 1937; Bezzubov 1949; Lisavenko 1959). They are good for pickling and soaking (Unchiev 1947), cider production (Kilchevsky 1933; Savitskij 1938), and wine (Murzaev 1934). The waste (pomace) of the wine industry is utilized for the production of high-quality apple pectin used in the confectionery industry (Saburov and Grzhivo 1931; Kilchevsky 1933; Turkin 1935; Sorokin 1947). Fruits of some wild apple forms are used for fresh consumption (Vorobiev 1935; Kirillov 1937; Speranskij 1936; Vasilichenko 1963; Zapryagaeva 1964), and almost all wild apples are suitable for dried fruit production. The use of wild fruits in the food industry is brought about not only by a necessity to use natural resources, but also because some cultivated apples do not always meet production requirements for such products as cider. In this connection, some countries with a developed fruit processing industry (France and Spain) deliberately breed specific, wild apple varieties for this purpose (Vecher and Bukin 1940). The use of wild apple fruits in the fruit processing industry is extensively reported in foreign publications (Mieville 1920; Brimble 1946; Chevalier 1953; Iorga 1964). B. History of the Wild Apple of Kazakhstan The first record of wild apple trees within the modern territory of Kazakh SSR were found in the letters of I. Sievers in 1796 from Barnaul, a chemist and correspondent of the Volni Economical Society (Koidzumi 1934). In that publication, he reported that on his travels to “Kirghizes” and on to Aligal Lake (apparently Alakol) he found a very “slender apple tree” on a southern macroslope of Central Tarbagatai in the Urdzhar River Valley. The fruits of this tree were like the well known “Ryazan apples of hen-egg size” in Russia. Subsequently, after working on the herbarium specimens of the apples from Tarbagatai, Ledebour (1846) gave them the name of Pyrus sieversii Ledeb. Later on, the modern name M. sieversii (Ldb.) M. Roem. became

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legitimate. A detailed study of mountain vegetation in Central Asia and Kazakhstan began in the second half of the nineteenth century. In that period of time, publications issued by P. P. Semenov (1858, 1867) rather precisely described the vegetation zones of Tien Shan. In addition, works published by K. Struve and G. Potanin (1864), N. A. Severtsov (1873), A. N. Krasnov (1887, 1888), and S. I. Korzhinsky (1896) characterized the zonal vegetation of Tien Shan and Tarbagatai. Pereselencheskoe Upravlenie, who organized a number of botanical–geographical expeditions to Zailijskei and Dzhungarskei Alatau, played a great role in botanical investigations of the mountains in modern southeastern Kazakhstan. The results of these investigations were presented in many publications, in which separate data about fruit forests of wild apples could be found (Lipsky 1906; Prasolov 1909; Bezsonov 1910; Rozhevits 1910; Mikhelivson 1913; Sapozhnikov and Shishkin 1914). Many sided and detailed investigations of wild fruit forests of Kazakhstan began only after the October 1918 Revolution. The aim of studying the Kazakhstan flora and plant resources for their economic development became possible at that time. For the first time, the flora of fruit forests was described by M. G. Popov (1929, 1934, 1940) whose activity had an enormous influence on the development of botanical investigations in Kazakhstan. A famous naturalist and rare expert on flora, he already by that time had extensively evaluated the wild apple of Kazakhstan, and referred to it as one of the best geographical species. The outstanding botanist N. I. Vavilov (1931) was interested in the wild apple of Kazakhstan, and he especially emphasized the necessity of preserving, studying, and selecting wild fruits of this genus. The mountain vegetation of Middle Asia and Kazakhstan attracted the attention of many scientists. Therefore, it is not accidental that much research was devoted to the history of mountain forests and their genesis (Krasnov 1888; Korzhinsky 1896; Lavrenko 1938; Popov 1940; Bykov 1956, 1957, 1966; Korovin 1961, 1962; Kamelin 1967, 1973, and others). Almost all investigators agreed that the modern forests of Middle Asia and southern Kazakhstan are the result of successive developments of tertiary vegetation and represent “the impoverished forest refugium” (Lavrenko 1938). The most complete and proven idea about the leading role in the xerophillic process of tertiary vegetation and the transmutation of subtropical flora (e.g. Ginkgo) into xerophillic forests of the ancient Mediterranean which resulted in wild fruit forests, including apples, was stated by M. S. Popov (1940). According to his concept, subtropical forests in the Neogene period underwent three stages of transformation into xerophillic forests: nut, oak, juniper/pistachio. However,

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Bykov’s data (1956, 1966) indicated, and paleobotany findings confirmed, that in the tertiary period in the territory of Tien Shan, the Pamirs-Alay and their spurs, the forest flora was of the Turgai type on the one hand and the caatinga flora or xerophillic thin forests on the other. Consequently, different homogenous and mixed forests could have been developed already by the tertiary period in the Tien Shan territory. Side by side with pure floral investigations, work was conducted also on separate wild apple species, forms, and their participation in plant associations. A number of publications were devoted to the wild apple species of Kazakhstan mountain forests. S. V. Yuzepchuk (1939), during his work on wild apple material for the book, “Flora of the USSR,” considered among all apple types, two species, M. sieversii and M. niedzwetzkyana, as most similar to cultivated forms. M. G. Popov (1934) designated the apples of the Zailijskei Alatau foothills to be a separate species M. dasyphylla, and M. niedzwetzkyana as a botanical variety of M. dasyphylla; M. niedzwetzkyana representing a variant with anthocyanin pigment. In 1949 Al. A. Fedorov and M. A. Fedorov described M. kirghisorum which has a number of features distinctive from M. sieversii. The mesophyllousness and growing of M. kirghisorum adjacent to relict walnut forests with walnut residues served as factors in recognizing it to be ancient in middle Asian flora and close to the initial tertiary apples. F. D. Likhonos (1964, 1968) held the same opinion; he considered M. sieversii as a subspecies of M. kirghisorum Al. et An. Theod. subsp. sieversii. A. M. Skibinskaya (1966) raised a serious objection against this opinion and referred to M. sieversii as a more ancient species than M. kirghisorum in the evolutionary process. The Soviet monographer of the genus Malus, V. T. Langenfeld (1970), presented a new scheme of phylogenetic relationships for the genus, according to which all middle Asian and Kazakhstan species were included in the series Kirghisores ser nova of subsection Malus, the true wild apples. Malus kirghisorum was recognized as the type of this series. Extraordinary polymorphism of the wild apple in areas of southeastern Kazakhstan served as a basis for separating a number of variations into independent species and forms. Thus, P. P. Polyakov (1948) side by side with M. sieversii described M. jarmolenkoi P. Pol. and M. schichkini P. Pol. in Zailijskei Alatau and M. linczevskii P. Pol. in Talasskei Alatau. V. V. Romanovich (1957) placed them in a category of conditional species and more or less denied them their own distinctive classes. A number of varieties were distinguished by B. A. Bykov (1956); in Zailijskei Alatau (var. sphenocarpa B. Bykov), and in Zailijskei and Talasskei Alatau (var. longicarpa B. Bykov). But most authors are inclined to

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believe that two apple species, M. sieversii and M. kirghisorum, and their numerous forms, are growing in the apple forests of Kazakhstan (Al. A. Fedorov and M. A. Fedorov 1949; Shitt 1952; Rubtsov 1956; Mushegyan 1962; Vasilichenko 1963; Vintergoller 1964, 1965; Likhonos 1964; Zhukovsky 1971, and others). The main component of apple plant communities within the borders of Kazakhstan is M. sieversii. The data on apple forests in the Kazakhstan mountain ranges are very variable. The main apple forest stands were mostly studied in Zailijskei Alatau (Popov et al. 1935; Popov 1940; Rubtsov 1941b; Polyakov 1948, 1950; Komarova 1959; Bukejkhanov 1960; Vintergoller 1964) and in Dzhungarskei Alatau (Rubtsov 1948; Polyakov 1950; Infantiev 1955, 1957, 1959; Komarova 1959). They were less studied in Tarbagatai (Stepanova 1959, 1962; Mushegyan 1962), in Talasskei Alatau (Popov 1940; Pavlov 1947; Karmysheva 1973), and in Karatau (Pavlov 1942, 1947). The irregular distribution of apple areas in Kazakhstan, their adaptation to different soils and hydrothermal regimes, and the complexity of their structures explain the considerable variety of their types. The first attempt to classify fruit forests of Zailijskei Alatau was undertaken by A. G. Klabukov (quote of Popov et al. 1935). Forest typology schemes of wild apples in Zailijskei and Dzhungarskei Alatau were proposed by P. P. Polyakov (1948), K. A. Pashkovsky and M. P. Yashchenko (1953), I. A. Bezpoludenov (1954), and V. I. Infantiev (1957). A series of investigations was devoted to studying the soils of the apple forest regions in Zailijskei Alatau (Prasolov 1909; Bezsonov 1910; Nadezhdin 1954; Glazovskaya 1944; Gerasimov and Matusevich 1945; Serpikov 1961; Sokolov 1962). Some authors (Nadezhdin 1954; Serpikov 1961) compared these soils to mountain forest, the others (Bezsonov 1910), to chernozem, and the third group (Gerasimov and Matusevich 1945; Sokolov 1962), to podzolized-chernozem soils. I. A. Bezpoludenov (1954) conducted rather detailed investigations of apple forest soils in Dzhungarskei Alatau. Forest taxonomy and the structure of apple forests in Dzhungarskei Alatau were presented by V. A. Infantiev (1957, 1959). The results of a root system study and natural restoration of M. sieversii were presented by A. A. Fedorov et al. (1945), Al. A. Fedorov and M. A. Fedorov (1949), V. A. Infantiev (1957), and I. T. Vasilichenko (1963). These authors pointed to the surface distribution of the wild apple root system, limited seed production, and wide root sucker formation. M. G. Popov and G. M. Popov (1925), V. V. Pashkevich and A. P. Sigov (1928), M. G. Popov (1929), M. G. Popov et al. (1935), N. I. Momot (1940), S. S. Kalmykov (1952, 1956), and V. I. Infantiev (1957) pointed to the wide variety of wild apple forms in Zailijskei Alatau and in the

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other ranges of northern and western Tien Shan. They noted the exceptional polymorphism of wild apple fruits. M. P. Dyrdin (1933), M. G. Popov et al. (1935), V. I. Infantiev (1957, 1959) and B. A. Vintergoller (1964) reported on using wild apple fruits of Kazakhstan for the production of syrup, applesauce, and dried fruits. Problems of apple forest exploitation in Kazakhstan were considered by A. G. Klabukov (Popov et al. 1935), V. A. Infantiev (1957, 1959), M. I. Gajdin (1962), A. N. Mushegyan (1962), M. K. Kalmykov et al. (1962), and B. A. Vintergoller (1964, 1965). Along with recommendations for improving cutting methods, most of the authors believed in the value of a domesticating scheme, that is, using wild apple trees as rootstocks for grafting cultivars right in the forest. B. S. Rodionov (1974) and B. S. Rodionov et al. (1975a,b) studied the structure and productivity of wild apples in Zailijskei Alatau and referred to the domination of common apple populations in woody plant stands, consisting of 71.8 percent by mass of the total community plant biomass. III. THE ROLE OF WILD APPLE IN THE VEGETATIVE COVER OF MOUNTAIN REGIONS IN KAZAKHSTAN Polymorphic forms of species comprise both the floral and community composition in the various natural environments and habitats of the natural fruit forests in the Kazakhstan mountain regions. All fruit trees and shrubs belonging to the mountain range flora of Kazakhstan are evidently oreaphytes. Only Elaeagnus angustifolia L. (Russian olive), which spreads within the low forest belt of wild fruit plant communities of Dzhungarskei and Zailijskei Alatau and Karatau, forming small areas of shrub (675 ha), may be referred to as a typical representative of deserts. Among the other woody species of these deserts are two shrubs: Nitraria schoberi L. and Lycium ruthenicum Murr. (box-thorn), growing on ledges with berrylike, partially edible fruits, but without food usage. Consequently, the mountains mainly in the central parts of Tarbagatai, Dzhungarskei, Zailijskei, Talasskei, and Kirghizskei Alatau are the natural centers of focus for almost all wood and shrub fruit plants and the communities formed by them in southeastern Kazakhstan. In these extensive mountain regions that differ by geological history, the mountainous geographical structure and natural conditions of these forests have produced their own composition and distribution features of systematic mixtures of fruit trees, other woody plants and shrubs, and the ecosystems formed by them. The total area of the forest mass including wild apples in mountain leskhozes is 1541.17 thousand ha (Table 2.1). Forest management of leskhozes was carried out by Kazlesproject

84

Total

Nonforested Total Agricultural Special use Abandoned

Forested Total Forested Unforested

125,034

3,082 10 155 2,917

121,952 119,908 2,043

863,434

498,846 291,348 4,173 203,325

364,588 249,202 115,386

Dzhungarskei Alatau

325,526

158,333 88,104 3,170 67,059

167,192 104,173 63,020

Zailijskei Alatau

147,600

112,837 49,225 723 62,889

34,763 20,438 14,324

Talasskei Alatau

Area (ha)

79,585

53,625 14,239 268 39,118

25,960 14,979 10,981

Karatau

1,541,179 (100%)

826,724 (54%) 442,927 (29%) 8,489 (<1%) 375,308 (24%)

714,455 (46%) 508,700 (33%) 205,755 (13%)

Total

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Tarbagatai

The mountain leskhoz territory in main land categories.

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Area

Table 2.1.

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from 1958 to 1959 until 1971 to 1973 (State Committee Materials of Forestry Economy). The forest area, including brushwoods is 714.4 thousand ha (46.4%). The area more heavily covered with forest is 508.7 thousand ha (33.0%). The forest area that is not heavily covered with forest is 205.7 thousand ha (13.4%), including thin forests (9.5%), burntover land (0.5%), and glades (3.4%). The total nonforested area is 826.7 thousand ha (54%). The area in agriculture is 442.9 ha (28.7%), lands of special use are 8.5 thousand ha (0.6%), and abandoned lands are 375.3 thousand ha (24.3%), including abrupt slopes and gullies, 229.4 thousand ha (14.8%), rocks and ice, 55.0 thousand ha (3.6%), and mounds and precipices, 55.0 thousand ha (3.6%). The remaining area consists of 35.9 thousand ha (2.3%). Besides state forests, there are some mountain forests mixed with wild fruit plants belonging to kolkhozes, state farms, and other economic organizations. The areas of these organizations are not given in the statistical data, and therefore, the forest areas cited here are slightly reduced. According to B. A. Bykov’s report (1966) shrub area, including fruits was reduced more than twice. The area under forest stands with wild apples within the borders of leskhozes is 143.9 thousand ha (20.1% of the total forest mass area), including conifers, 83.0 thousand ha and deciduous trees, 60.3 thousand ha (Table 2.2). Shrubs (barberry, currant, honeysuckle, pearlbush, spiraea, sweetbriar, and others) are widely distributed in the mountains covering an area of 146,599 ha (46.3% of the forest covered area). In Tarbagatai they constitute 33.5 percent of the forested area, in Dzhungarskei Alatau 48.6 percent, in Zailijskei Alatau 39.8 percent, in Karatau 69.2 percent, and in Talasskei Alatau, 54.1 percent. The fraction of fruit trees in coniferous forests is insignificant. Now and then isolated trees of M. sieversii appear within the lower borders of Tien Shan spruce forests in Zailijskei and fir forests in Dzhungarskei Alatau. In Karatau, single specimens of Crataegus pontica C. Koch., Pyrus regelii Rehd., and Amygdalus spinosissima Bge., sometimes grow in the lower part of the xerophyllous thin forests of Juniperus semiglobosa Rgl. Apple participation in forest associations is characterized by the following data. The area under apples in the mountain leskhozes of Kazakhstan is 12083.0 ha (71.6 percent of the total area of natural fruits). The area under apples (2411.1 ha) in sovkhozes and kolkhoses is not included. Within the mountain ranges, the percentage of wild apple distribution is the following: in Tarbagatai (2.0%), Dzhungarskei Alatau (48.8%), Zailijskei (25.4%), Karatau (12.1%), and Talasskei Alatau (11.7%) (Table 2.2).

86

ha

– – 191 257 – – 4102 8980 115 – – – – – – 73 – –

Acacia Alder Almond Apple Apricot Ash Aspen Birch Bird cherry Cherry Elaeagnus Elm Fir Grape Hackberry Hawthorn Juniper Lime

– – 0.3 0.5 – – 7.6 7.6 0.2 – – – – – – 0.1 – –

% – – – 5891 86 – 8635 11762 – – 623 380 7545 – – 20 – –

ha – – – 3.8 – – 5.6 7.6 – – 0.4 0.2 4.9 – – – – –

%

Dzhungarskei Alatau

1 – 3064 609 6 2784 673 – – 52 102 – – – 40 – 1

–

ha – – – 4.8 1.0 – 4.4 1.0 – – 0.1 0.2 – – – – – –

%

Zailijskei Alatau

2 22 1452 790 502 3 142 – 181 – 113 – 2 554 669 1850 –

–

ha

Karatau

– – 0.1 5.6 3.0 – – 0.6 – 0.7 – – – – 2.2 2.6 19.1 –

%

– – – 1419 190 – – 114 – 50 – 4 – – 110 842 3370 –

ha

% – – – 8.0 1.1 – – 0.6 – 0.3 – – – – 0.6 7.6 19.1 –

Talasskei Alatau

2 1 213 12083 1675 508 15524 21671 115 231 675 599 7545 2 664 1644 5220 1

ha

– – 0.1 3.8 0.5 0.2 4.9 6.8 – 0.1 0.1 0.2 2.4 – 0.2 0.5 1.7 –

%

Total on mountain ranges

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Tarbagatai

Distribution of the predominate species within the forest-covered areas of leskhoz mountain ranges.

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Species

Table 2.2.

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– – – – – – – 401 21926 18151 – – – –

54196

Maple Mountain ash Other woods Pear Pine Pistachio Plum Poplar Purple willow Shrubbery Spruce Sweet cherry Walnut Willow

Total

100.0

– – – – 0.1 – – 1.2 2.3 48.6 25.3 – – – 63990

– 656 – 3 38 339 25453 30022 – 9 64

15 2 57

100.0

– – 0.1 – 1.0 – – – 0.5 39.8 46.9 – – 0.1 25656

1127 42 190 8 – 19 – 21 61 17751 – 1 1 153 100.0

1.4 0.2 0.8 – – 0.1 – 0.1 0.2 69.2 – – – 0.6 17684

247 – 834 – – 58 23 – 769 9564 – – 42 48 100.0

1.4 – 4.7 – – 0.3 0.1 – 4.4 54.1 – – 0.3 0.3 317140

1421 44 1081 8 858 77 26 2315 26604 146599 69389 1 52 292

100.0

0.5 – 0.3 – 0.3 – – 0.7 8.4 46.3 21.9 – – 0.1

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155614

32 – – – 202 – – 1855 3509 75680 39367 – – 27

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100.0

– – – – – – – 0.7 40.5 33.5 – – – –

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A. Physical and Geographical Assay of Apple Forest Distribution The mountains of southeastern Kazakhstan are at the periphery of the Central Asian mountain chains, which spread far to the West within the desert plains of Prebalkhash and Turan. The alternation of powerful upsurges and wide depressions, tracing within separate mountain systems, conforms to the general regularity in layout of the orographical structure of the mountain country in southwestern Kazakhstan. Thus, Zaisan depression separates the systems of Altai, Tarbagatai, and Saura. Balkhash-Alakul depression delimits the systems of Tarbagatai, Dzhungarskei Alatau and the northern Tien Shan, spreading far to desert plains of the West Karatau mountain range. Mountain chains of Tarbagatai, Dzhungarskei, Zailijskei, Kirghizskei, Talasskei Alatau, and Karatau are the main regions of wild apple (Fig. 2.1). 1. Tarbagatai. This is a mountain chain stretching between 46° to 48°N in latitude and 80° to 84°E in longitude. This is a monolithic mountain chain with a length of 250 km and average altitudes of 2000 to 2100 m. Glaciers and eternal snow are absent, but in some gorges on the mountain summits snow lies almost constantly. The layer structure of mountains is exposed rather clearly. The mountain surface is flat or slightly

Fig. 2.1. Distribution of Kazakhstan wild apples: 1 = Malus sieversii; 2 = M. kirghisorum; 3 = M. niedzwetzkyana.

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broken by plots to the north. To the south, the mountain gradually changes into middle-mountain relief with mild contours. The peripheral parts are vast sweeps of low mountains, among which the intermountain hollows are widely developed. The northern border of the wild apple of Kazakhstan extends through the southern macroslope of Tarbagatai. The Alakul hollow and the Dzhungar gate separate the mountain systems of Saura and Tarbagatai from Dzhungarskei Alatau. 2. Dzhungarskei Alatau. This is a complicated mountain country (the whole chain is about 400 km long), comprised of some stepped mountain ridges, divided by intermountain hollows and spread mainly in a latitudinal direction between 44° and 46°N latitude and 78° and 82°E longitude. Some of its spurs are extended far within the Balkhash-Alakul hollow. The orographical and geological structure of the Dzhungarskei Alatau is similar to more southern mountain chains of the Northern Tien Shan and is therefore often referred to as the last one (Rubtsov 1948). The River Koksu divides the Dzhungarskei Alatau into two masses, abrupt southern and ledge northern slopes. The region of the northern middle mountain of the Dzhungarskei Alatau is the habitat of wild apples, which form considerable stands here. 3. Zailijskei Alatau. This is the most northern ridge of the Tien Shan mountain range. Ilijskaya hollow separates it from Dzhungarskei Alatau, and divides deep tectonic valleys of the rivers Chilik and Kemin from Kungei Alatau, which is situated to the south. Chu-Ilijskie mountains, consisting of a number of eroded ridges, are spread toward the western tips of Zailijskei Alatau. The North Zailijskei Alatau gradually changes to hilly and slightly sloped premountain plains. The total extent of the ridge is 280 km long, and the largest width in the northern slope is 30 to 40 km wide. The low mountains and high mountains are widely developed into a geomorphological ridge. The modern area of wild apples is referred to here as the middle mountain belt. 4. Kirghizskei. The northern part of Kirghizskei Alatau spreads within Kazakhstan, the main mass of which is in Kirghizia. It is the extension of the Kungei Alatau and constitutes its western part. In the East, it is separated from Kungei and Zailijskei Alatau by the deep and narrow Boam gorge and the valley of the river Chu. In the West the river Talas separates it from Talasskei Alatau. Positioned almost in a latitudinal direction, Kirghizskei Alatau limits the wide Chujskaya valley, to which a longer and wider (40 to 50 km) northern slope faces to the South. It is rather steep and strongly cut, especially in the northeastern part,

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coming down by stepped ledges to the premountain valley. Wild apples also grow in the middle mountain zones, mainly along river terraces. 5. Karatau. The mountain chain Karatau is located in the western part of the northern arches of the Tien Shan ranges between 43° and 45°N Latitude and 67° and 70°E Longitude. This mountain chain separates from Talasskei Alatau in the low Chokpak region and stretches to the northwest for a distance of 420 km. The southern part of the chain is a complex low mountain system (1600 to 1800 m) consisting of a number of smooth plateaus, cut by small ridges in the northwest and northeast directions. Wild apple grows mainly on the southwestern slopes. 6. Talasskei. The main ridge of Talasskei Alatau is a northern branch of Tien Shan and extends almost in latitudinal direction with a few deviations to the northwest. From the main heights of Talasskei Alatau, mostly stretching within Kirghizia, some high ridges extend to the southwest. The region investigated by us is illustrated by a plot referring to the system of the northwestern ranges of Talasskei Alatau. The notion “west part of Talasskei Alatau” is similar to the notion “Kazakhstan part of Western Tien Shan.” The lack of outstanding peaks and high ridges with almost inaccessible slopes is characteristic of all vast spaces with spurs of Talasskei Alatau. The northern steep slopes have some smooth plots, but the southern ones are rocky and descend abruptly to the rivers. Here, wild apple grows exclusively on the northern slopes of ridges and in river valleys. B. Features of Forests 1. Hydrography. Most of Kazakhstan is situated in flowless basins and only the Irtish system is a part of the Arctic Ocean basin. Flowless basins are numerous. A large basin is in Balkhash-Alakul hollow, to where the rivers stream down from many parts of Northern Tien Shan, Dzhungarskei Alatau, and the southern slope of Tarbagatai. Many mountain rivers do not reach this large basin because on the plains they are used for irrigation and most of them are quickly lost in sands or in flowless hollows. From the southern Tarbagatai slope the rivers Eginsu, Kuzak, Urdzhar, Khatynsu, and others flow down. From the northern slope of Dzhungarskei Alatau to Lake Balkhash, the rivers Sarkan, Bien, Aksu, Ugensu, and others flow. To the river Ili from the northern slope of Zailijskei Alatau, the rivers Talgar, Kaskelen, Bolshaya and Malaya Almatinka, Issik, and Turgen

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flow down. The hydrographical network of Karatau consists of numerous rivers, springs, and lakes. The largest rivers are in the southern half of Karatau and belong on the one hand to the Syr Darya basin and on the other hand to the basin of the Bijlyukul lake groups. The numerous mountain rivers of Talasskei Alatau are Dzhabagli, Aksu, Kshi-Aksu, Bugulutur, and Bala-Baldarbek. In the valleys of the mountain rivers, apple trees grow more extensively than in other places of the mountain relief. 2. Climate. The climatic conditions are the result of complex and constant interactions of litter surface, solar radiation, atmospheric circulation, and other characteristics of the given country. The deep intercontinental situation of Kazakhstan mainly determines all the important features of its severe continental climate, including its low air humidity, insufficient quantity of precipitation on most of the territory (the steppes), hot summers, short winters in the South, and long, severe winters in the North. The mountains of eastern and southeastern Kazakhstan are in different latitude zones. Tarbagatai is within a desertsteppe zone. The rest of the mountain regions are within deserts (in the northern zone, Dzhungarskei Alatau). In the southern part of temperate zone, it changes into a subtropical zone. The Kazakhstan western Tien Shan is the most southern part of the whole mountain chain. Extensively cleaved relief of the mountain system (rugged high ridges, mid-mountain and low mountain regions with their own characteristics of structure, geology) is the main reason for the exceptional differentiation of mountainous ecological environments. The great differences in climatic conditions of the mountain regions have a significant influence on the plant cover, determining the variety of vegetation from the desert to the high mountains. All climatic elements change according to the place and altitude. For instance, the mean monthly air temperature in Zailijskei Alatau decreases on the average by 1°C with each drop of 100 m in altitude (Dzhangaliev 1972). The daily temperature may change from 1°C to 2.5°C depending on the weather and the season of year. The soil thermal gradient decreases according to soil depth. For example, at a layer of 0 to 5 cm, the temperature is 4.0 to 4.5°C, at a layer of 5 to 10 cm, 3.0 to 3.5°C; and at a depth of 40 cm, 2.0 to 2.5°C. Precipitation gradients and elevations at 1000 to 1700 m above sea level are positively related, that is, as the elevation increases, the quantity of rain also rises. The precipitation gradient at each 100 m of altitude is the following: 1000 to 1100 m, 80 mm; 1200 to 1300 m, 50 mm; 1400 to 1500 m, 25 mm, and 1500 to 1600 m, 10 mm. With altitude increasing, solar radiation also raises accordingly:

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at a height of 850 to 1100 m direct solar radiation is 2.74; 1100 to 1600 m, 2.49; 1600 to 1800 m, 2.04 kcal/cm2 per year for each 100 m of mountain height. The large exposure of a slope and the slope’s steepness greatly influence the microclimate of a territory. A different slope orientation at all altitudes causes a difference in air and soil temperatures, moisture regimes, and so on. Thus, the average daily air temperature of southern slopes with steepness 20 to 25° is 1.0 to 1.5° higher than those of northern slopes. The thermal conditions of mountain regions vary both with altitude above sea level and within one elevation, at different slope inclinations. Within one altitude, heat accumulation on slopes can be rated in the following way: the coldest are northern, followed by eastern, western, and southern slopes as the warmest. For the main wild apple habitat of Kazakhstan in the middle mountain zone, the climate is moderate continental. One of the important characteristics of a thermal regime is its average yearly air temperature, which for all forest foothills fluctuates from +4.1°C in the southeast (Tarbagatai) to +9.2°C in the southwest (Talasskei Alatau). The coldest month of the year is January with a mean temperature that ranges from –3.8°C (Zailjiskei Alatau) to –15.5°C (Tarbagatai). The mildest winters are in the middle mountain areas while the lowest temperatures are restricted to deep cleaved hollows (Lepsinskaya, absolute minimum of 52°C) where specific conditions for heat accumulation and cold radiation of air exist. July is the hottest month everywhere with mean temperatures ranging from 17.3°C (Dzhungarskei Alatau) to 22.4°C (Zailijskei Alatau). The absolute highest maximum is observed in Karatau and Talasskei Alatau (45° and 39°C, respectively). The ridges of southern and southeastern Kazakhstan belong to a mountain system with insufficient precipitation. The moisture contrast of different mountain ridges, and the regions of apple habitat referred to here, is extremely great. The annual precipitation fluctuates within large limits, from 343 (Karatau) to 888 mm (Zailijskei Alatau). The greatest part of the precipitation falls in the first part of the warm period, the maximum is May having 43 to 150 mm. The composition and distribution of apple forests of Kazakhstan are mostly related to the quantity of precipitation and air temperature. On the whole, the apple forest structure changes from the northeast to the southwest and becomes increasingly adapted to insufficient precipitation, arid conditions, and high summer temperatures. 3. Soil. The process of soil formation in mountain regions is related to the elevation of the location. At specific altitudes, soils are steppe, forest-steppe, and meadow-forest types. The soils of the steppe belt are chernozem, thin-humus, and weakly leached and the meadow forest

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regions have mountain meadow subalpine dark forest soils. The ecological and soil conditions of wild apple forests are rather varied, from patchily colored low mountain soils at elevations of 850 to 900 m, to rock ridges at more than 2000 m and from fertile mountain chernozem soils, where apples form medium dense stands, to mountain steppe residually calcareous soils under thin forests. Isolated trees grow on rocks, stone mounds, or spread within the subalpine zones, where they grow into dwarf bush shapes. Apple forests of Zailijskei and Dzhungarskei Alatau grow on deep and medium deep chernozem which are in different stages of leaching. Therefore, the trees here are very large. The wild apples of Tarbagatai, Talasskei Alatau, and Karatau grow mainly on stony, weakly developed, medium deep chestnut soils, and sometimes on mountain chernozem soils. Here the trees are smaller in size. C. Wild Apple Role in Plant Cover of the Mountain System The vegetative features of each mountain range are determined by a number of factors: latitude and longitude of the location, geological structure of the mountain ridges, and relief and slope characteristics. The amount of precipitation depends on the range’s main direction, its river valleys, slope exposure, and altitude above sea level. The soil mechanical structure, soil characteristics, and morphology are directly dependent on the abovementioned factors and on vegetation. Soils together with vegetation determine the surface runoff and the amount of water penetrating into soil, and consequently, the soil humidity. All of these factors influence the species composition of a plant community and the interactions among them. Under the pressure of these natural historical factors, which create the environmental conditions and the zone distribution of vegetation, landscapes are formed. Precisely expressed descriptions of natural zones in the Kazakhstan mountains were contributed by many investigators of Tarbagatai, Zailijskei, Dzhungarskei, and Talasskei Alatau (Struve and Potanin 1864; Krasnov 1887; Prasolov 1909; Krasheninnikov 1925; Abolin 1930; Rodin 1933; Glazovskaya 1944; Bykov 1956; Dragavtsev 1956; Sokolov 1959; Dzhangaliev 1967; Stanyukovich 1973, and others). The distribution and regularity of the mountain vegetation is initially conditioned by vertical zonality and is mainly related to the formation of high landscape belts. But the conditions for such zonality are not always available. The fullest expression of vegetative zonality is in the mountain ridges that have a latitudinal direction. Here, the main exposures usually dominate and the distinctively vegetated landscapes are unique (Bykov 1956). Local conditions (relief, exposure, slope steepness,

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amount of light and moisture content of different plots) greatly influence the plant composition and form a mosaic belt structure. Thus, significant differences in the structure of the landscape zonality on slopes are the results of distinctive climatic exposures. Mostly, it is expressed by the biogeo-development of low belts that spread to upper ones along hotter and drier southern slopes and then in reverse order, communities of the upper belts descend to low ones along cooler and wetter slopes of northern exposure. On southern slopes, here and there, separate belts are quite divided in comparison with northern slopes. The asymmetry of altitude zonality, especially peculiar to the mountains, exists in the regions of continental climate. But considerably less asymmetry is expressed in the mountain regions of oceanic climate, for instance in the Caucasus (Lavrenko and Sokolov 1949). In contrast to the boreal mountains (the Altai and the Sayani), the Kazakhstan mountains include in one belt a number of absolutely opposite plant formative elements: steppe, meadow, thin forest, rock vegetation, and so on. Zone borders are seldom clear and straight but are more rarely horizontal and spreading at one elevation. Twisting lines usually represent borders. For instance, the upper borders of a forest belt are seldom flat and above the border, separate trees usually grow, then small forests and finally dense forest appears. In some Kazakhstan mountain ranges, natural conditions are formed, in which the traits of zonal vegetation are clearly expressed. For example, the belt profile of Dzhungarskei Alatau is a typically normal expression of a boreal belt. E. P. Korovin (1962) described the structural types of high landscape zonality and referred to it as the northern Tien Shan group and distinguished it from the following belts: mountain semidesert, soddy steppes, defoliated forests and forest meadows, alpine meadows, and cold-loving cushions. The belting of the northern Tien Shan appears in some form in the mountains of Zailijskei Alatau. Here, the forest belts are well represented and a steppe belt has already been formed by semi-savanna vegetation. An analogous picture is observed on the northern slope of Kirghiz Alatau, where premountain plants change from steppe to mixedgrass semi-savannas. The profile of Kirghiz Alatau was characterized as a transitional one by R. V. Kamelin (1973), because vegetation traits of the northern and southern mountains are combined in it. The impoverished belt profile of Talasskei Alatau may serve as an example of a typical deviation from the normal belt picture of the northern Tien Shan group. Here, fir groves and deciduous forests do not take part in the belt characteristics. Situated in deep gullies and gorges, they do not spread to watersheds. In the western part, a special juniper belt

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is formed. Some anomalies are observed in the belt system of southern Karatau, where at the upper belt of the large grass subtropical steppes (semi-savannas), sod grass and feathered sheep’s fescue grass steppes are joined. When moving from the North to the South (from Tarbagatai, Saura to Talasskei Alatau) the common regularity in structure alters from altitude zonality to gradual rising, and a complex of structural characteristics is observed. Thus, the lower border of woody vegetation on the southern Tarbagatai landscape is at 600 to 700 m, in Dzhungarskei Alatau, 900 to 1100 m, on the northern slope of Zailijskei Alatau, 1200 to 1400 m, and Talasskei Alatau, 1500 to 1700 m. When moving to the South, the interlandscape structure of forest meadow steppe zone changes from dark coniferous forests (Saur) to the Tien Shan fir woods and Turkestan juniper forests. Vegetation zones (belts) in the different mountain Kazakhstan ranges, in meters above sea level, are: Vegetation Zone Tarbagatai (Stepanova 1962) Desert-steppe belt Steppe belt Shrub belt Subalpine belt Alpine belt Dzhungarskei Alatau (Stanyukovich 1973) Desert belt Steppe belt Forest-meadow belt Half cold-loving medium grass meadow and juniper belt Cold-loving low-grass meadow belt

Altitude (m) 500–700 700–1000 1000–1700 1700–2400 2400–3100 under 800–850 800–1500 1500–2300 2300–2800 above 2800

Zailijskei Alatau (Pavlov 1949) Desert belt Mixed belt Fir wood belt Half cold-loving sub-alpine belt Cold-loving alpine belt

under 600 600–1600 1800–2500 2500–2800 2800–3300

Zailijskei Alatau—fruit zone (Dzhangaliev 1967) Desert steppe zone Plain steppe zone

under 600 650–750

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Vegetation Zone Premountain zone Lowmountain zone Middlemountain zone Talasskei Alatau (Karmysheva 1973) Premountain semi-desert plain belt Semi-savanna belt Juniper and dry meadow belt Subalpine belt Alpine belt Snow belt

Altitude (m) 750–850 850–1100 1100–1700 1300–1400 1400–1600 1700–2200 2200–2800 2800–3300 3300–3400

In spite of various methods of different investigators (botanists, resource scientists, pedologists, or geographers) on selecting zones (belts), the picture of their landscapes is similar. So, in Zailijskei Alatau, almost all investigators separate a desert zone at altitudes below 600 to 700 m, a steppe zone below 1200 to 1400 m, coniferous forests higher than 1600 m, and subalpine, alpine and snow zones even higher. We agree with B. A. Bykov’s (1956) opinion, considering that the belt is revealed by formations situated on exposures corresponding to the main ridge slope, that is, under the concept “belt” horizon zone of mountain vegetation is meant, growing under definite physico-geographical conditions (altitude, soil, temperature, humidity, and relief) and characterized by a definite combination of vegetation types: zonal, intrazonal, and azonal. An obvious comparison draws attention to the differences in zone names expressed by soil scientists M. A. Glazovskaya (1949) and S. I. Sokolov (1959) and the zones proposed by fruit grower A. P. Dragavtsev (1956). A premountain zone at 650 to 900 m height (fruit grower’s term) corresponds to feather grass sheep’s fescue and grasswormwood steppes on mountain-chestnut soils (soil scientist’s term), a low mountain zone at 900 to 1200 m altitude conforms to bush steppes on mountain chernozems, and a middle-mountain fruit zone at the elevation of 1100 (1200) to 1700 (1800) m corresponds to motley grass meadow steppes and deciduous forests on gray forest soils and leached chernozems. The present day diversity of natural fruit forests of southeastern Kazakhstan is concentrated on middle mountains at an 800 to 2000 m altitude, in forest meadow, forest steppe and steppe belts of deciduous forest associations, at forest borders, and in shrub communities. In the

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same areas, the vegetation from motley grass steppe or high-grass meadow formation is also developed. In the southern part of Karatau, wild apples grow sporadically and in limited quantities, mainly as single trees and shrubs. In the central part of Tarbagatai, in Talasskei, and in Kirghiz Alatau, apple groves grow. In Zailijskei and Dzhungarskei Alatau, apple grows abundantly and frequently form apple forests. The ecological type of wild apple is regularly dominant in plant distribution. It is a plant that is more adaptive to the moderately frigid climate of the forest steppe region in the Northern Hemisphere and prefers the type of Eurasian vegetation supporting climate (Popov 1925) to hot and dry desert climates. This confirms P. V. Kamelin’s (1973) proposition that forest communities of Central Asia (southern Kazakhstan may be included) represent impoverished and more northern variants of forest flora in the eastern part of the Mediterranean region. Malus sieversii, M. kirghisorum, and M. niedzwetzkiana represent the wild apples of Kazakhstan. According to M. G. Popov (1927), V. T. Langenfeld (1971) and P. V. Kamelin (1973), one may believe that the former ancestor of wild apples existed in a subtropical flora of the upper Cretaceous and Pleocene ages. The most ancient and the closest to the original tertiary apple is M. kirghisorum, which has a mesophilous nature and grows together with the other relicts of the mesophilous Turgai forest flora, Juglans regia L. In the process of adapting to dry conditions, the descendants of the first mesophilous apple transformed and gained adaptive characteristics of a xeromorphological type, that is M. sieversii. Malus sieversii exists in a wider area than M. kirghisorum within the xerophyllous physical and geographical conditions of the Kazakhstan mountain regions and is more evolutionarily advanced. Under the present day conditions mentioned above, different apple species have different roles within the composition of plant community and vegetative cover. For example, M. sieversii has a wide and definite distribution in all places, forming large populations. In contrast, the role of M. kirghisorum in the modern landscape is minor and topographically limited to the middle mountains of Zailijskei and Dzhungarskei Alatau, along rivers and streams. Malus niedzwetzkyana, discovered by B. A. Bykov (1957) in Karatau and by I. I. Roldugin (1973) in Zailijskei Alatau (Pratov 1976), is few in number, restricted in area of distribution and nowhere forms large populations. Therefore, M. sieversii, because of its wide ecological range dominates as a mesophyllic, autotrophic tree species. The vast area of the species spreads by a rather wide but not uneven zone within Kazakhstan, extending and narrowing, depending on the different ecological conditions and natural factors of its habitats. The modern extent of M. sieversii in Kazakhstan is not continuous. Its

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most northeastern distribution is on the mountain macroslope in the central part of Tarbagatai within the Stepnoi forest reserve of Semipalatinsk region. A second distribution is on the lower part of the northern macroslope of Dzhungarskei Alatau and on its hilly modulating foothills, composed of powerful thick loess-like loams within the Andreevsky, Lepsinsky, and Sarkandsky forest reserve of Taldy-Kurgan region. Further to the southeast, in the low part of the northern macroslope of Zailijskei Alatau (Prigorodny and Alma-Ata forest farms of Alma-Ata region) is the third plot. The fourth plot is situated to the west of it, on the northern macroslope of Kirghiz Alatau (Dzhambulsky forest reserve of Dzhambul region). The fifth plot is on the northeastern slope of Karatau (Boraldaisky forest reserve) and the sixth, on the northwestern slopes (Ugamsky and Badamsky forest reserves) of Talasskei Alatau. Beyond the borders of Kazakhstan, M. sieversii grows in the territory of Kirghizia, where it penetrates into today’s areas of the more mesophillous M. kirghisorum on the slopes of the Fergana and Chatkalsk mountain ranges. Generally speaking, M. sieversii is the most widely distributed species of the genus Malus not only in the mountains of Kazakhstan but of Central Asia, western Tien Shan and the Pamirs-Alai, where its distribution extends to an altitude of 2300 to 2600 m (Pratov 1976). 1. Tarbagatai. The northern border of M. sieversii distribution in Kazakhstan is on the southern macroslope of the ridge. When comparing the vegetation cover of vertical belt schemes for Tarbagatai with other ridges (Saur, Dzhungarskei, and Zailijskei Alatau), it can be seen that there is a complete lack of the forest belt here. In the shrub belt of Tarbagatai, small aspen and apple forests appear, and though they have a landscape aspect, they do not form a belt. The lack of a forest belt here is related to a comparatively low altitude (2000 to 2100 m) of the ridge, the absence of glaciers, neighboring deserts, and dry climate. Tarbagatai woody vegetation is rather poor. There are no coniferous forests here and deciduous forests are characterized by impoverished plant fragments, growing mainly along valleys of the longest rivers and consisting chiefly of Salix kirilowiana Stschegl., mixed in various proportions with Populus laurifolia (Lam.) Gilib., Crataegus altaica Lange, and occasionally Betula microphylla Bge. In the central part of southern ridge slope of the shrub belt is wild apple (Fig. 2.2). The bush wood belt mainly spreads on the steep southern macroslope, growing on mountain steppe xerophytic leached soils in combination with mountain chernozems. Here, apple forests are distributed on slopes of different exposures, except for northern exposures, in valleys in the

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Malus sieversii in the Tarbagatai mountains.

beds of small rivers. Above the alluvial plain terraces at altitudes of 1000 to 1500 m, some apple stands are encountered at altitudes of 1700 m. Apple development is included in the following vegetative groups: almond-apple, sweetbriar (rose)-apple, and aspen-apple forests. O. F. Stepanova (1962) proposes a more detailed scheme of apple participation in plant associations, adding spirea-almond-apple and poplar apple forests. Wild apple exists as only one species M. sieversii, which forms plant communities with a 0.2 to 0.6 wood stand density. Apple trees of a 2 to 7 m height, depending on the microbiological conditions, are covered with strong prickly shoots. Apple stands are characterized by well developed secondary shrub canopies, consisting of Amygdalus ledebouriana Schlecht., not found growing in Zailijskei and Dzhungarskei Alatau, and Rosa spinosissima L. Spiraea hypericifolia L. often grows with an admixture of Daphne altaica Pall. Lonicera tatarica L., Rhamnus catharica L., and Berberis heteropoda Schrenk. Grass cover is rich in species number and is represented mainly by mesophillous Eurasian boreal species.

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2. Dzhungarskei Alatau. This is an isolated system of mountain ranges that are united under the same name and usually considered to be a part of northern Tien Shan. The Altai and Siberia vegetation have more growth here (except in Tarbagatai and Saura) than in other Kazakhstan mountains, because Dzhungarskei Alatau is situated at the point of union of the ancient Altai system and the geologically younger Tien Shan. Existing in Dzhungarskei Alatau, a number of boreal floral representatives, such as Abies sibirica Ledeb., Berberis sibirica Pall., Betula rezniczenkoana (Litv.) Schishchk., and Crataegus sanguinea Pall. survive beyond its borders as well. Sorbus tianschanica Rupr., Juniperus turkestanica Kom., Lonicera karelinii Bge. ex P. Kir., L. olgae Rgl. et Schmalh., L. altmannii Rgl. et Schmalh., and L. humilis Kar. et Kir. extend from the south and are established here. The great differences in community ratios of meadow steppe and steppe types of Dzhungarskei Alatau ridge slopes allowed N. I. Rubtsov (1948) to distinguish special regions, North Dzhungarskei and South Dzhungarskei. The wide spread species distributions of Tien Shan origin are related to the geobotanical South Dzhungarskei region where the landscape aspects are equal with those that are observed in Eastern and Central Tien Shan. In contrast, the influence of Siberia and the Altai, according to N. I. Rubtsov (1948), is mainly noted in the North Dzhungarskei region. Especially in the Lepsinsk geobotanical subregion, certain plant characters are concentrated that point to a genetic relationship of the flora and vegetation of Dzhungarskei Alatau with that of the Altai and Siberia. Here, on the slopes of Topolevsk and Lepsinsk intermountain hollows, significant apple stands are situated (Fig. 2.3). A vast territory with wild apples is in the eastern part of Konstantinovka village. Separate wild apple groves are seen in the west to the Zhaman-Terekti River and in the east to the Kiberti River and on southern slopes of the Bulan-Bai Mountain. Along river valleys, single apple trees are scattered a little more widely. I. I. Roldugin (1973) points to typical populations of M. sieversii in Tokti village near Dzhungar Gate at an altitude of 1300 to 1500 m. As a whole, wild apple in Dzhungarskei Alatau forms well expressed altitude-climatic zones at 1200 to 1600 m. Apple forests grow on the lower parts of forest-meadow belts that include the whole forest-steppe belt and later spread to the upper part of the steppe, forming the following plant groups: Forest-meadow belt: (1) apple groves (M. sieversii); (2) apple-aspen forests Forest-steppe belt: (1) apple forests (M. sieversii, M. kirghisorum); (2) apple forests (M. sieversii) with aspen; (3) forests in river val-

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leys (M. kirghisorum); (4) bushwood on southern slopes mixed with apple (M. sieversii) and hawthorn Steppe belt: (1) bushwoods with isolated apple trees (M. sieversii); (2) apple groves with hawthorn Forest-meadow Belt. This belt extends between the altitudes of 1500 to 2300 m. Here, M. sieversii is not widely distributed. Fir forests prevail in the upper part of the belt, mixing with Abies sibirica and Populus tremula. Sorbus tianschanica, Padus racemosa, and Crataegus songorica (C. Koch) are found in the understory. On the northern cold and wet slopes, M. sieversii is almost completely forced out by these species. On the southern warmer exposures, apple distribution is limited by unfavorable soil conditions such as strong relief cleavage and a constant surface exposure of natural rocks, mostly resulting from poorly developed soil formations, which definitely cannot provide vigorous growth and apple cropping. The trees here are small and often bushy and biennial apple bearing is sharply expressed.

Fig. 2.3. Distribution of wild apple forests on the northern slope of Dzhungarskei Alatau (Dzhangaliev, Zener, 1969): 1 = distribution of apple and apple-aspen forests; 2 = desertsteppe belt; 3 = steppe belt; 4 = forest-steppe belt; 5 = forest-meadow belt; 6 = spurs of the main ridges; 7 = borders of natural belts; 8 = thin groups and isolated wild apple trees; 9 = hilled watersheds (1200–1600 m), separating intermountain valleys from desert regions.

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Vegetation in the lower part of the belt exists in apple-aspen forests. On the southern slopes at a height of not more than 1800 m, apple populations are in good condition. The uniqueness of the forest-meadow belt is in its wide development of meadows not only on northern exposures but on southern slopes as well. The grass cover is vigorous (100–120 cm high) and consists of the usual mountain meadow plants: Dactylis glomerata L., Alchimilla sibirica Zam. and other species, to which the weedy species Urtica dioica L. and Artemisia vulgaris L. are admixed in significant quantities. Forest-Steppe Belt. This is situated at 1200 to 1600 m, and here, as a rule, M. sieversii and M. kirghisorum form closed forests. On richer soils a constant companion of wild apples is Populus tremula. In the upper part of the belt P. tremula frequently forces out apples and forms pure stands, with an insignificant admixture of Sorbus tianschanica and Acer semenovii Rgl. et Herd., which are derivative forest types. Crataegus songorica, C. altaica, Padus racemosa, and Sorbus tianschanica are secondary to wild apple. On the slopes of southern orientation where poorly developed soils prevail, aspen is excluded from apple communities. The soils here are predominantly sandy, lying right on stone mounds and on firm natural rocks that are frequently exposed during the day. In general, rocks and stone mounds are very characteristic elements of specific southern slope landscapes. Here, apple grows together with hawthorn and small understory species: Lonicera stenantha Pojark., L. tatarica, Berberis heteropoda, Rhamnus cathartica, Rosa platyacantha Schrenk., and Spiraea hypericifolia. Along the valleys of numerous small rivers, M. kirghisorum is a component of forests on alluvial plains. Forest stands of Populus laurifolia (true Altai plant) are seen here and were also found in Tarbagatai during our expedition in September 1972. Poa nemoralis L., Origanum vulgare L., Agrimonia asiatica Yuz., Hypericum perforatum L., Polygonum coriarium Grig., Urtica dioica, Humulus lupulus L., Aegopodium alpestre Ledeb., and Geranium rectum Trautv. are frequent components of grass cover. In this belt, apple forests grow and bear well on slopes of the northern and eastern exposures, which are on strongly and medium leached forest chernozem soils. Here, M. sieversii and M. kirghisorum form 0.6 to 0.8 density stands (Fig. 2.4). Steppe Belt. This belt lies between altitudes of 800 (900) to 1200 m. In the Lepsinsk subregion, it is considerably less well developed than in forest-meadows. Bordering on the slopes of the main range it is found between 800 and 1200 m and on eastern exposures between 800 and 1600 m. Consequently, the upper border of the steppe belt on the eastern slopes extends somewhat higher than on northern slopes. This is

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Formation of apple forests in Dzhungarskei Alatau (forest-steppe belt).

apparently related to a steep fall of the eastern range to the desert Alakol depression, the influence of which promotes a greater quantity of xerophyllous plants in the mountain profile (Rubtsov 1948). The existing mesophillous meadows do not play a significant role in the steppe belt landscape, which is found along river valleys and on steep northern slopes, and on plots well-wetted by surface subsoil water. The foundation of grass stands on motley grass meadow steppes is mainly composed of mesophillous species and such communities are referred to by N. I. Rubtsov (1948) not as steppes but as steppe-meadows. In the lower part of the belt, vegetation does not form a dense cover, existing as overgrown bushwoods of Spiraea hypericifolia, Rosa spinosissima, R. platyacantha, and Berberis heteropoda with often stunted, solitary apple trees. Grass cover is formed by Festuca sulcata Hack., Sedum hybridum L., Artemisia vulgaris, and Glycyrrhiza uralensis Fisch. A weakly developed root system and insufficient reserve of nutrient substances and moisture do not produce growth and fruit bearing in wild apples. In the upper part of the belt on northern slopes (1000–1100 m), M. sieversii develops mixed with Crataegus songorica and C. altaica.

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3. Zailijskei Alatau. The next apple forest stands of significant size in Kazakhstan are in Zailijskei Alatau, which resembles the Dzhungarskei vegetation cover, having the same meadow-forest aspect and a similar belt type. The Zailijskei Alatau flora consists of the southern Tien Shan species, most of which extend to the middle-mountain belts and especially to the high-mountain regions. At the same time, the Pamirs-Alai species are already widely distributed here. In the fir forests of the middle-mountain belt, a rich complex of mountain-taiga species has developed, though Abies sibirica is already disappearing. The southern border of the ancient arctic species Populus tremula, Padus recemosa, Rubus idaeus L., and others also extends to here. In the high climatic belt of apple forest stands, an important position is taken by Armeniaca vulgaris Lam., Acer semenovii, some hawthorn and honeysuckle species, and in the understory, Euonymus semenovii Rgl. et Herd. and many other shrubs, not growing in Dzhungarskei Alatau and Tarbagatai. Apple trees are distributed at an altitude of 1100 to 1800 m and stretch from the Bolshaya Almatinka River on the west to the Belchabdar River on the east. Separate wild apple populations spread a little bit wider on the west to the M. Dolan River, on the east to the Kiik-Bai River. On protected plots (slope bottoms, dry places) that receive additional moisture, and on isolated river valleys, wild apple trees are seen along the whole lower mountain part of the northern Zailijskei Alatau slopes (Fig. 2.5). According to species and variety zoning (Dzhangaliev 1967), apple forests are distributed in the low mountain zones (steppes), 800 to 1100 m, in middle mountains (forest-steppes), 1100 to 1500, and forest-

Fig. 2.5. Distribution of wild apple forests on the northern slope of Zailijskei Alatau (Dzhangaliev, Zener, 1965): 1 = apple forests; 2 = desert-steppe belt; 3 = steppe belt; 4 = forest-steppe belt; 5 = forest-meadow belt; 6 = spurs of main ridges; 7 = borders of natural belts; 8 = isolated wild apple trees.

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meadow-steppes, 1500 to 1800 m. Wild apple takes part in the following vegetation groups: Forest-meadow belt: aspen forests with admixture of M. sieversii Forest-steppe belt: (1) apple forest, (M. kirghisorum); (2) apple forest, (M. sieversii), mixed with Armeniaca vulgaris; (3) woodshrub overgrowths with M. sieversii Steppe belt: overgrowths with isolated stunted M. sieversii Apple overgrowths have the character of open park groves. Below, they are replaced by groves of Armeniaca vulgaris and Acer semenovii and above by Populus tremula overgrowths. In this connection, the species of border vegetative formation may be mixed with wild apples. Forest-Meadow Belt. The northern slopes of the upper parts of this belt are occupied with spruce forests of Picea schenkiana Fisch. et Mey. Below, Populus tremula grows to altitudes of 1100 to 1200 m and 1500 to 1800 m, where it forms pure and thick overgrowths, which resemble real multistoried forests. In spite of a closed canopy inside these forests, there is enough light for the development of vigorous grass vegetation of a meadow-steppe character. Sometimes Sorbus tianschanica, Acer semenovii, and M. sieversii mix with aspen trees, which, as a rule, are in a dwarfed state. Fruit tree felling in many respects promotes apple supplanting by aspen. Ribes meyeri Maxim. and sometimes Rubus idaeus are seen in interwoods. Forest-steppe Belt. On gradual slopes of the northern belt exposures, the wild apple forms tight (density 0.3–0.8) woodstands on thick humus leached chernozem soils (Fig. 2.6). On southern slopes with sandy soils, apple stands are thinner. Here, Armeniaca vulgaris appears more frequently. Though apricot groves are secondary in the mountain vegetative structure, apple trees in Zailijskei Alatau are very characteristic. The grass cover in Armeniaca vulgaris overgrowth is of a xerophyllous character. On stone mounds, Crataegus songorica stands are frequently seen with apple totally absent. Here, C. songorica forms only light overgrowths with a 0.5 to 0.6 density. In the apple understory, Rosa platyacantha, Lonicera stenantha, Rhamnus cathartica, Berberis heteropoda, and others appear rather frequently. A tight cover of rather vigorous grasses such as Melica altissima L., Brachypodium silvaticum (Huds) Beauv., Festuca gigantea (L.) Vill., and Aegopodium podagraria L. forms a grass layer. On forest-free plots in meadows, cock’s-foot motley grass and cock’s-foot short motley grass are distributed. “Tien Shan kurai” is formed on cattle pastures from long grasses: Liguralia macrophylla (Ledeb.) DC., Rumex tianschanicus

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Apple forest (Malus sieversii) in a forest-steppe belt in Zailijskei Alatau.

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Losinsk., Heracteum dessectum Ledeb., and Inula helenium L. (Pavlov 1940; Polyakov 1948). Steppe belt. The vegetative cover is of a steppe character, with a predominance of typical wormwood and motley grass wormwood communities. Significant areas are under sweetbriar and Armeniaca vulgaris, Craetaegus altaica, and M. sieversii, which mainly grow on northern slopes (1000–1100 m). Previously in this belt, apple had a much wider distribution. However, as a result of felling, the area under apples was significantly reduced. 4. Kirghizskei Alatau. Malus sieversii grows here on the northern macroslopes, the Kazakhstan part of the range. The flora is typical of the Middle Asian mountain province (Kamelin 1973). Here, side by side with the common Tien Shan vegetation, representatives of the Turkestan flora are distributed. The quantity of Pamirs-Alai species Lonicera korolkovii Stapf., L. cinerea Pojark., L. tianschanica Pojark., and Acer turkestanicum Pax. greatly increases. At the same time, there are still many boreal species. In Kirghizskei Alatau, many southern forms of Abelia corimbosa Rgl. et Schmalh find their northern borders, and some northern species also reach their limits. Plant communities of M. sieversii (Fig. 2.7) prefer a steppe of 800 to 1500 m and sometimes prefer meadow-steppe belts of 1500 to 2000 m. Under conditions of increasing xerophytic conditions, wild apple grows in small thin groups of trees, which develop on rather wet soils. Along with forms of M. sieversii are Pyrus regelii, Craetaegus pontica, Celtis caucasica Willd., Berberis heteropoda, Spiraea pilosa Franch., Cotoneaster melanocarpa Lodd., Rhamnus cathartica, Lonicera tatarica, L. microphylla Willd. ex Roem. et Schult., Rosa beggeriana Schrenk, and Amygdalus spinosissima. Short-lived perennials, such as Hordeum bulbosum L., and Prangos auscheri Boiss, accompany the grass cover. 5. Karatau. Malus sieversii (Fig. 2.8) and sometimes M. niedzwetzkiana are seen on the mountain slopes of the Karatau range. The vegetation cover here is to a great extent under the influence of neighboring deserts. On dry slopes of the range, junipers merge into pear-hawthorn thin forests consisting of Pyrus regelii, Crataegus pontica, Celtis caucasica, and Amygdalus spinosissima. Many Iranian and Iranian-Himalaya species are distributed in Karatau (Kamelin 1973). Here, the separate areas of northern species are known, that is, separate areas in Karatau, and dense distribution in northern Tien Shan including Populus nigra L., Rhamus cathartica, and others. The grass cover consists of short-lived

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Malus sieversii in Kirghizskei Alatau.

perennials: Hordeum bulbosum, Agropyrum trichoporum (Link.) Richt., and Prangos pabularia Lindl. Conforming to strain-variety zoning (Dzhangaliev 1967), wild apple distribution in Karatau is in premountain (700–900 m) and partially in middle-low mountain zones (900–1200 m). In such extremely dry conditions, wild apples form very resistant populations, and solitary dwarfed specimens are often seen. Trees are, as a rule, bushy with short, crooked and often numerous stems. 6. Talasskei Alatau. In the plant cover formation of this mountain range, most types of northern Tien Shan vegetation are found, though they are seen in specific combinations: spruce-fir and juniper forests, forest subalpine meadows and forest steppes, steppes and semideserts, meadowsteppe juniper thin forests and shrub semi-savannas. On slopes of northern exposure that are better provided with water and less exposed

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Wild apple in Karatau mountains.

to shading, the boreal type of vegetation is dominant. Groups of wild apple are distributed among shrub overgrowths on slopes with surface ground water and along river terraces at 1200 to 1800 m heights, which rise 200 m on the southern slopes (Karmysheva 1973). Malus sieversii comprise stands with a density of 0.4 to 0.5 (Fig. 2.9). Attendant species include: Crataegus turkestanica Pojark., Sorbus persica Held., Acer semenovii, Rhamnus cathartica, Euonymus koopmannii Lauche, Rosa alberti Rgl., R. fedischenkoana Rgl., Lonicera tianschanica, and Amygdalus petunnikowii Litv. According to N. Kh. Karmysheva’s data (1973), the grass cover under the forest canopy is thin and grows mainly near shady forest species including Poa nemoralis, Impatiens parviflora DC., Rumex tianschanicus, Ligularia songarica (Fisch.) Ling., Aegopodium podagraria, and others. Dactylis glomerata, Poa pratensis L., and Bromus inermis Leyss. are dominant in open spaces and glades. Within the ground cover, the most prevalent dicotyledons are Campanula glomerata L., Galium aparine L., Medicado tianschanica Vass., and others. According to this data, it is evident that the apple forests of the Kazakhstan mountain systems are subjected to the law of vertical (height)

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Fig. 2.9. Kara-Alma apple grove in Talasskei Alatau (northern slope of the River Aksu). Photo by A. F. Kovshar.

zonality and are formed within 800 to 2000 m limits. This belt, especially the middle mountains, is characterized by relatively optimal humidity conditions and a moderate average yearly temperature, which positively influences the renewal, growth, and productivity of apple forests (Dzhangaliev 1972, 1974a). The upper border of the wild apple distribution in Kazakhstan mountains is limited as a rule by temperature drops and a shortening of the vegetative period. The lower border is limited by a moisture deficiency. The highest altitude of wild apple distribution is in Zailijskei and Talasskei Alatau (up to 2000 m), and the lowest altitude of apple distribution is in Tarbagatai and Karatau (800–900 m). In Tarbagatai, apple grows as a rule on southern slopes because here the colder temperature is a limiting factor for its distribution. When moving toward the south, apple progressively occupies the northern slopes (Dzhungarskei and Zailijskei Alatau) under conditions of less light, air dryness, and moisture deficiency. Apples grow along river valleys and gorges, where surface ground water compensates for atmospheric humidity shortage. Here as a rule, they are spread on slopes with northern exposures (Kirghizskei and Talasskei Alatau, Karatau). The mountain apple forests of Kazakhstan are similar both to forestforming species and to resistant forest community types. According to E. P. Korovin’s data (1962), in the wild apple communities of Middle Asian and South Kazakhstan more than 30 species of shrubs and trees are registered. The list of grass vegetation includes about 100 species.

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Due to its ecology, apple is coupled with different hawthorn species, which take part in the formation of mixed apple-hawthorn associations. Thus, apple in Tarbagatai is associated with Crataegus altaica, which is typical of the boreal forests of Eurasia. Side by side with C. songorica, apple is widely spread in Dzhungarskei and Zailijskei Alatau. In apple forests of Talasskei Alatau, these species are substituted for pure middleAsian C. turkestanica and C. pontica (Table 2.3).

Table 2.3. Distribution of Malus and associated wood-shrub vegetative species in Kazakhstan mountains.

Plants Acer semenovii Rgl. et Herd. Amygdalus communis L. Amygdalus ledebouriana Schrenk Amygdalus petunnikowii Litv. Armeniaca vulgaris Lam Berberis heteropoda Schrenk Crataegus altaica Lange Crataegus pontica C. Koch Crataegus songorica C. Koch Crataegus turkestanica Pojark. Lonicera stenantha Pojark. Lonicera tatarica L. Lonicera tianschanica Pojark. Malus kirghisorum Al. et An. Theod Malus niedzwetzkyana Dieck. Malus sieversii (Ledeb.) Roem Padus racemosa (Lam.) Gilib. Populus laurifolia Ledeb. Populus talassica Kom. Populus tremula L. Rhamnus cathartica L. Ribes meyeri Maxim. Rosa fedtschenkoana Rgl. Rosa platyacantha Schrenk Rosa spinosissima L. Rubus idaeus L. Salix macropoda Stschegl. Sorbus persica Hedl. Sorbus tianschanica Rupr. Spiraea hypericifolia L. Spiraea pilosa Franch. + = present; – = absent

Tarbagatai

Dzhungarskei Alatau

Zailijskei Alatau

Talasskei Alatau

– –

– –

+ –

+ +

+ – – + + – – – + + –

– – – + + – + – + + –

– – + + + – + – + + –

– + – + – + + + – – +

– – + + + – + – – – – + – + – – + –

+ – + + + – + + + – + + + + – + + –

+ + + + – – + + + – + – + + – + – –

– – + – – + – + – + – – – – + – – +

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Representatives of the genus Populus are typical components of apple groves. Populus laurifolia is characteristic of Tarbagatai apples while in Dzhungarskei and Zailijskei Alatau, P. tremula is usual, but is absent in mountains of Kirghizskei and Talasskei Alatau. In Talasskei Alatau, the other representative of this genus, P. talassica Kom., is seen in apple stands. Almond, Amygdalus ledebouriana, which is found in Tarbagatai apple forests, is absent in Dzhungarskei and Zailijskei Alatau, and in the Talasskei apple-woods, it is replaced by the species A. petunnicowii and A. communus L. In the apple forests of the Kazakhstan mountain systems, the representatives of the shrub genera Lonicera and Rosa are typical, though their species composition undergoes a number of changes, depending on the geographical state of the apple distribution region. Thus, in the apple groves of Tarbagatai, Rosa spinosissima is very characteristic, which together with Amygdalus ledebouriana forms a clearly expressed, barely permeable second canopy. In the mountains of Zailijskei and Dzhungarskei Alatau, it is replaced by R. platyacantha and in Talasskei Alatau, hawthorn is associated with another more xerophyllous species, R. fedtchenkoana. The boreal species Lonicera stenanta and L. tatarica are specific for apple forests of Tarbagatai, Dzhungarskei and Zailijskei Alataus. In western Tien Shan, these species are replaced by another more drought-resistant form L. tianschanica. The whole complex of these cited materials, namely, the broad area of distribution, the abundance of apple tree habitats in the Kazakhstan mountains, the significant numbers of trees, often with large plant community associations and the presence of regularly repeated specific features of apple development gives grounds for considering them to be in a group of Aestilignosa forest types. C. Classification of Wild Apples in Kazakhstan The differentiation of mountain environments, the scattering of apple areas, their adaptation to different soil-climatic conditions, and the structural complexity both of the floral and of the plant composition determine the wide variety of community types. The vegetative cover mosaic and the presence in each forest stand of remnants of similar types with particular plant community interactions are characteristic of the Kazakhstan wild apples. In this connection, when describing wild apple types, we assign them somewhat generalized characteristics according to the combinations of growing conditions, which are a result of the biogeodevelopmental status of the forest types (Sukachev and Dylis 1961). Dzhangaliev (1974a) noted the merits and disadvantages of fruit forest classifications by A. G. Klabukov (quote of Popov et al. 1935), P. P.

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Polyakov (1948), I. A. Bezpoludenov (1954), K. A. Pashkovsky and M. P. Yashchenko (1953) and V. I. Infantiev (1955) and the necessity of a new classification. When working out the typological scheme for wild apples, they used the soil network of E. V. Alekseev and P. S. Pogrebnyak and the diagnostic indexes of the genetic classification by B. P. Kolesnikov and applied to Kazakhstan conditions by L. N. Gribanov (1964). The variety of growth conditions of wild apples is presented as a generalized scheme of ecologic-topographic habitat types on a geomorphological basis with an evaluation of their potentialities for wood mass and fruit productivity and taxonomic peculiarities of forest stands (composition and density). Within each general climatic type, two main geomorphological complexes are distinguished, mountain and valley wild apples. In our wild apple classification, a group of forest types was determined as a lower but not the main taxonomic unit. These forest types were selected according to established types of growing conditions. This provided a potentially similar forest-vegetative effect. When distinguishing forest types, the soil moisture content of the habitat was considered first, then soil fertility, grass cover, and species combinations of understory species and the forest canopy, reflecting the uniqueness of the forest process. The ecologic-topographic scheme of growth-condition types for wild apples and their potential productivity is given below. 1. Very Dry Growth Conditions (Fig. 2.10, 2.11). These conditions include vast areas of the regions in the steppe belt that are in the process of erosion. In the forest-steppe belt this condition is seen much less frequently. In the forest-meadow, it is only found in spots on rocky slopes. Very dry growth conditions are on rocky and upper plots of the southern slopes. The soils are mountain-steppe and residually calcareous, medium and poor chernozems. For such conditions, chronic moisture deficiency is characteristic. Outflow prevails over accumulation. In the second part of summer (July to August), in the upper part of the profile, humidity reduces to the point at which it is unavailable for plants. The humidity in the remaining part of the profile is almost unavailable. Average and readily available moisture appears with autumn (November) precipitation. The vegetative cover has a dry-steppe aspect, in which wormwood-type and thyme-type associations are widely distributed, including groups of low Spiraea hypericifolia, Rosa platyacantha, and Berberis heteropoda. Isolated apple trees seldom grow. Festuca sulcata, Sedum hybridum, Glycyrrhiza uralensis, Artemisia vulgaris, and Sophora laponica L. are seen in the grass cover, and occasionally Poa nemoralis. The forest-forming species, M. sieversii, at the age of 21 to 30

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Fig. 2.10.

A. DZHANGALIEV

Very dry type of wild apple growing conditions.

Fig. 2.11. Locations of soil moisture under apple stands in different types of habitats or growth conditions: A = very dry, B = dry, C = on slopes, D = semi-humid in valleys; 1 = unavailable, 2 = almost unavailable, 3 = average availability, 4 = readily available.

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years has a 2 to 3 m height with a trunk diameter of about 5 to 15 cm. The average annual increment of shoot growth does not exceed 6 cm. In this type of forest vegetative conditions of very dry spiraea-sweetbriar, wild apples develop. Having adapted to the most unfavorable habitats, they are naturally accompanied by low producing wood stands and the yield in favorable years does not exceed 400 kg/ha (Bonitet III). The growth and bearing characteristics point to the low productivity of this group of apple tree types. Such areas do not provide the soil moisture and nutrient substances necessary to maintain plantings of moisture-loving fruit plant forms. It is necessary to strictly preserve the existing forests because they protect the soil from destruction and supply valuable material for breeding, especially for selection of droughtresistant forms. These forms are interesting for cultivation under severe nonirrigated conditions. Under the unfavorable conditions of the rockslope, fruit communities, as vegetative pioneers, play an important role in developing the rock-stony mountain soils and taking part in their formation so that forest vegetation can become established. 2. Dry Growth Conditions (Fig. 2.12). These conditions are widely distributed in the steppe and forest-steppe belts. These conditions are present on slopes of southern and western exposures and watersheds. The

Fig. 2.12.

The dry type of wild apple growing conditions.

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soil cover is mountain chernozems, weakly leached, medium and shallow heavy loams. For such conditions, insufficient moisture is mainly characteristic, but at times it is sufficient. Moisture outflow and accumulation occur at approximately the same time. In the second half of the summer, moisture is almost unavailable in the soil profile. However complete moisture unavailability with soil withering seldom occurs. In the lower part of the profile at a depth of 180 to 200 cm, soil moisture is preserved within average availability. It is natural that in the second cycle of vegetation, apple withstands evident moisture shortages, although in contrast to very dry growth conditions, its shortage here may be compensated by the humidity from lower horizons. The vegetative cover is of a steppe character with sheep’s fescue-wormwood-motley grass and motley grass-wormwood communities prevailing and large areas are covered with steppe shrubs. The woody vegetation under dry conditions is associated with small groves of M. sieversii, Crataegus songorica, and C. altaica. Shrubs of Rosa platyacantha, Berberis heteropoda, and Rhamnus catharica often grow. Origanum vulgare, Vicia cracca L., Agrimonia asiatica, Hypericum perforatum, Achillea filipendulina Lam., and Dactylis glomerata exist in grass stands. The forest forming species, M. sieversii, has a height of 4 to 5 m, trunk diameter of 20 cm, and a yield of 750 kg/ha. Apple productivity, in comparison with very dry conditions, is somewhat higher. In the forest vegetative conditions of dry grass-shrub, apple trees with hawthorn develop. In order to increase apple-stand productivity, it is necessary to take some measures for moisture preservation, such as grass mowing and soil mulching under the tree crowns. During forest reclamation in such conditions, preference should be given to Armeniaca vulgaris as the most drought-resistant plant and to the selected drought-resistant Malus sieversii forms from the same plots. 3. Semi-moist Growth Conditions on Slopes (Fig. 2.13). These conditions dominate in the apple forests of Zailijskei and Dzhungarskei Alatau and are characteristic of forest-steppe and forest-meadow steppe belts. In the steppe belt, such conditions are not widely spread and are found at the foot of the steep northern slopes and more seldom on the eastern slopes. In the forest-steppe belt, they are observed on the northeastern, northwestern, and eastern slopes that have a 20 to 30° steepness. The soil cover is accompanied with mountain, strongly leached, thick and medium-thick chernozems. For such conditions, sufficient moisture is characteristic, but periodically, insufficient. As a whole, water accumulation is greater than outflow. In dry summer periods, the upper part of the profile is withered down to almost unavailable moisture but at the

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117

Semi-moist type of growth conditions of wild apples on slopes.

depth of 90 to 100 cm, an average available humidity is preserved in the soil (Fig. 2.11B). M. sieversii develops here in close masses with 0.6 to 0.8 density. Crataegus songorica, C. altaica, Populus tremula, Sorbus tianschanica, Ribes meyeri, Rubus idaeus, R. caesius L., Lonicera tatarica, and Rhamnus cathartica grow in the forest structure. Grass cover is represented by Brachypodium silvaticum, Urtica dioica, Polygonum coriarium, Aegopodium alpestre, Geranium rectum, Heracleum dissectum, Dactilis glomerata, Alchimilla sibirica, and others. Under these conditions, wild apple grows well and bears. Tree height is 6 to 8 m; trunk diameter, 20 to 30 cm; and yield, 4000 to 5000 kg/ha. Here, fresh grass-shrubs and wild apples grow with hawthorn and aspen. 4. Semi-moist Growth Conditions Along Valleys (Fig. 2.14). These conditions are not widely distributed in the mountains of Zailijskei and Dzhungarskei Alatau but extend onto precipices of ancient terraces and outflow hollows. The soils are meadow chernozems and degraded chernozems. Sufficient moisture prevails due to an additional flow from higher plots. Here, even under drought conditions of the summer period, at a depth of an average of 130 to 140 cm, humidity is preserved (Fig. 2.11D). Vegetation is represented by apple groups with a thick grass

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Semi-moist type of growth conditions of wild apples along valleys.

cover of Aegopodium alpestre, Polygonum coriarium, and Arctium tomentosum Mill. In the understory, Rhamnus cathartica and Rubus caesius are seen. Tree height is 9 to 10 m; the annual growth increment is 10 to 16 cm; yield is 300 kg/ha. In these forest-vegetative conditions, fresh grass-motley-grass wild apples are formed. It should be noted that in the third and the fourth types of growth conditions described, most of the highly productive wild apples are distributed. Here, are the best large-fruited wild apple forms and the area has important seed growing plots, this being a seed production source for nurseries in the southern part of Kazakhstan. The plots with moist growth conditions on slopes and intramountain valleys are highly favorable for planting large-fruited forms. To increase wild apple productivity, it is necessary to take measures to reduce surface drainage from adjacent slopes. For man-made plantings, it is recommended to reclaim plots with good air drainage. In locations remote from favorable conditions of the third and fourth types, apples can be moved down the alluvial plains of the mountain rivers where they are closer to ground water and they grow on a shallow soil with a surface bedding of pebble-stone

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layers. Apple can withstand short-term flooding because of additional root formation during aerated conditions. 5. Moist Growth Conditions (Fig. 2.15). These conditions occupy insignificant areas on the valley bottoms and the lower slopes, which are the places of additional ground moisture. Soils are soddy-humus, and are shallow or medium-deep. For any given growth condition, excessive moisture of a stagnant type is characteristic. In the steppe belt, vegetation is represented by shrub overgrowths of Rosa platyacantha and Berberis heteropoda, and rarely Armeniaca vulgaris and Crataegus songorica are seen. In the forest-steppe belt, wood vegetation is formed by groves of Malus, Armeniaca, Crataegus, and Populus tremula. Besides Rosa and Berberis, bush woods such as Lonicera tatarica and Salix macropoda Stschegl. are seen growing with exceptionally thick grass vegetation of meadow type of Dactylis glomerata, Heracleum sibiricum, Urtica dioica, and Geranium rectum. In these forest-vegetative conditions, moist herbaceous wild apples are found. Tree height is 7 to 10 m; trunk diameter is 30 to 40 cm; the annual growth increment is 10 to 20 cm; and the fruit yield is 200 to 300 kg/ha. In order to increase their productivity, it is necessary, in addition to common management practices, to take measures to reduce the flow of water from higher lying

Fig. 2.15.

Humid type of wild apple growth conditions.

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plots. The great diversity of soil conditions, stony soils and water regime differences, limit the mass development of artificial plant communities. Limiting the propagation of forms growing here and the preservation of existing plantings are the most rational courses of action. The common procedure of growing wild apples under conditions of the second and the fourth types should be sanitation, aiming at clearing them periodically of dying and faulty trees and cutting out dead branches in their crowns. In dry apple woods, besides general management, it is necessary to take measures to detain and conserve water by building tree pans, digging shallow gutters across the slopes for flow detention, early grass mowing, and mulching soils with grass under the crowns. In moist apple woods, additional measures should be taken to reduce water ground flows by ridging and terrace building. In apple forests growing in the first condition type described, the full implementation of any management system, including the improvement of felling and clearing of dying and faulty trees, is needed in relation to the erosion danger of the plot. Data on M. sieversii distribution, according to the type of growing conditions, characterize this species as a xeromesophyllous ecomorph and a xerophyte, the characteristics of which are most distinctly revealed under dry and very dry conditions. Under semi-moist and moist conditions, it usually discards its xerophyllous characteristics and in its morphological characteristics, it approximates a type of mesophyllous M. kirghisorum. Its ecological plasticity allows M. sieversii to preserve its own areas and to form clusters in new, quite different (dry and humid) habitats. Under conditions of an increasingly xerophyllous mode, ecological flexibility is more strongly expressed in the xeromorphous species, M. sieversii, than it is in the mesophyllous species, M. kirghisorum, which grows in moisture-rich habitats. Probably, the lower capacity to adapt to dry habitats by M. kirghisorum may be explained not as much by its loss of formative uniqueness, as a more ancient species, but because it grows in ecologically more mesophyllous and multiforming vegetation, where it is not able to fully realize its potential. The wide distribution of M. sieversii and growth in different habitats testify to its high vitality and adaptability to multiple ecological conditions, especially to drought, with a corresponding parallel variability of morphological characteristics, the most important of which is its ability to create many forms of natural renewal (Dzhangaliev 1973a). This characterizes M. sieversii as a promising species with a well-expressed adaptive capacity under dry climatic conditions of southeastern Kazakhstan.

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IV. THE INFLUENCE OF WILD APPLES ON THE STRUCTURE OF THE ENVIRONMENT The vital activity of apple stands is closely related to the environmental conditions. This relationship is mutual. On the one hand, distribution, growth, and development of wild apples depend on the environment as a whole as well as on separate ecological factors. On the other hand, because of their own vital activity, wild apples change the conditions of their environment. Additionally, in relation to root system activity and natural apple renewal, tree roots expend a considerable amount of mechanical work when they penetrate the soil. Apple stands also have a definite influence on the microclimate. While studying the interrelation of wild apples and their environmental conditions, we must consider the mutual relations among biotic and abiotic components. Because apple environments are heterogeneous (their composition includes many components closely connected with each other), and for consideration of the role of ecological growth and developmental factors, it was necessary to use experimental methods in field investigations. In spite of the great difficulty and tedious work of these investigations, the need to conduct them was dictated by the following two considerations. First, the materials relating to the role of ecological factors in wild apple life testify to the biological significance of wild apples and that they may be used for wild apple reintroduction into the vertical zones of the mountains. Second, such materials on natural fruit plantings were collected for the first time and they represent a scientific interest in their own right. A. Influence of Apple Stands on Microclimate and Apple Response to Vertical Zone Conditions Investigations were conducted in the central part of Zailijskei Alatau, on established experimental wild apple plots with crown closures (density) of 60 to 80 percent. The areas studied were 20 × 20 m in size, at specific altitudes of 1050, 1350, and 1900 m above sea level. Open treeless areas of the same size were situated nearby and served as controls. Observations were conducted on tree crowns at a height of 9 m and on inside crowns at 3 and 1.5 m above the soil surface. When laying out the stationary plots for microclimate investigations, we used the methods of V. N. Sukachev (1956), E. M. Lavrenko (1959), and B. P. Korol (1959). 1. Precipitation. The main source of apple-stand water was rain. In spite of considerable water evaporation from the canopy surface, the greater

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the rainfall, the greater is the amount of water available to the trees. During botanical investigations, it was important to measure the precipitation distributed at each level of altitude, which is an indirect index of the whole complex of ecological and hydrometeorological factors, including moisture indexes. Precipitation is precisely an objective moisture index which accurately reflects the regularity of humidity changes in the soil cover (Fig. 2.16). The curve of the long-term annual precipitation index is practically parallel to the distribution curve of mean moisture reserves in a meter of soil during the vegetative period. When necessary, coordination of these factors permits the use of precipitation data as a moisture index for the whole territory. The distribution curve of the average long-term soil moisture reserves during the vegetative period is an accurately expressed precipitation maximum, at a height of about 1600 m. There is a higher moisture reserve even with reduced precipitation. This is explained by much more gravel-sandy soils and consequently by less water-holding capacity at heights above 1600 m, and also by increasing total evaporation, which is caused by a lower atmosphere pressure. Changes in the mean annual precipitation over a 20-year period (1944–1963) and the mean moisture reserves during the vegetative stage in a meter soil layer (for 1962–1968) at each 100 m level of altitude are presented in Table 2.4. The precipitation gradients are positive over altitudes from 1000 to 1700 m, that is, with increasing altitudes, the quantity of rain increases accordingly. The average increase in gradient is 43 Moisture reserve (mm) 350

A

300 250 200 Precipitation (mm) 1000 B 900 800 1000

1200

1400 1600 1800 Altitude (m)

2000

Fig. 2.16. (A) Mean moisture reserve in a meter of soil layer for the vegetative period depending on altitude above sea level and (B) differences in long-term annual precipitation.

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Table 2.4. Altitude gradients of annual precipitation and moisture reserves in a meter of soil layer.

Altitude above sea level (m) 1000–1100 1100–1200 1200–1300 1300–1400 1400–1500 1500–1600 1600–1700 1700–1800 1800–1900 1900–2000

Annual precipitation (mm)

Gradient precipitation (mm, at 100 m)

Mean moisture reserves in soil (mm)

Moisture reserve gradient (mm, at 100 m)

690–770 770–830 830–880 880–915 915–940 940–950 950 950–940 940–910 910–860

+80 +60 +50 +35 +25 +10 0 –10 –30 –50

170 215 245 270 295 310 320 320 315 290

+45 +30 +25 +25 +15 +10 0 –50 –25 –45

Data of moisture reserves refer to the slopes of northern exposure.

mm for each 100 m in height increase. Above 1700 m, precipitation decreases and the gradients become negative. On average, precipitation decreases 30 mm for each 100 m increase in altitude. The greatest rainfall is at an altitude of 1600 to 1700 m. The change in moisture reserves in a meter of soil layer, depending on altitude, is analogous to the annual precipitation distribution. At the same time, the mean gradient for altitudes of 1000 to 1600 m is 25 mm for each 100 m of height. At altitudes higher than 1700 m, the gradient is the same, but with a negative index. These correlations provide a basis to use an average annual precipitation as a characteristic humidity for the territory as a whole. The rain in apple stands and in open places is unequally distributed. The canopy structure of apple woods promotes light and rain redistribution. Light in apple groves penetrates through a series of green organic filters, absorbing the main part of the physiologically active radiation. Atmospheric water, due to its downward flow and to evaporation from the multilayered vegetative cover, penetrates into the soil to a lesser amount. According to observations made in 1963 on a treeless plot from April to August, 417 mm of rain fell, but under the forest canopy only 343 mm or 18 percent less. Especially great differences were noted during the period of full tree defoliation when treeless areas received 20 to 25 mm more rain than defoliated areas. Moreover, the soil under apple stands has more infiltration capacity. Thus, apple woods increase the water absorption and water accumulation on slopes and thus provide soil conservation and fertility. This is one of the many apparent aspects

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of the erosion prevention and water preservation role of apple communities in the mountains. In the soil, various transformations of rain water take place. The moisture content of the soil mainly depends on the time and amount of water falling, the water-physical soil properties, the degree of slope, the composition of the vegetation, and its mass. Many investigators have studied soil humidity regimes in the forests and in open areas. G. N. Vysotsky (1952) on the basis of an analysis of literature data and his own observations concluded that the role of forests is declining. At the same time, the investigations conducted by S. V. Zonn et al. (1953) did not find that soils under forests are declining. A. A. Molchanov’s (1952) opinion is very important. He believes that moisture depletion by any culture is not totally conditioned by the current increment of rainfall but by the humidity regime, plus the plant community character, climate, soil, and other physical and geographical conditions. Forest vegetation under mountain conditions, in the opinion of most investigators, promotes soil moisture preservation and favorably influences the regime of river outflows. Our investigations (Table 2.5) showed that apple stands promote soil moisture preservation. During the whole vegetative period, the total moisture reserves in soils was rather high (724.2–1149.3 mm). In the second half of summer, reserves were slightly reduced, however water soil saturation did not fall lower than 80 percent of the full field water capacity (FFW). On treeless plots, the moisture content in the 0 to 300 cm soil layer was somewhat higher in comparison to apple stands. This is quite natural as water depletion by woody stands exceeds that required by grass vegetation. Thus, apple stands promote water regime improvement in the soil. This may be explained by the favorable moisture preserving microclimatic and soil conditions that are formed under the wood canopy. Table 2.5. The influence of apple stands on the total moisture reserves in strongly leached deep chernozem (0–300 cm), Zailijskei Alatau, 1963. Total moisture reserves (mm)

Stand density Apple-stand, aged 80 (density 0.3) Apple-stand, aged 80 (density 0.7) Glade

Full field water capacity

May 20

June 20

Aug 14

Sept 30

Oct 30

870.1

1149.3

1144.4

915.5

781.8

885.7

897.0

1085.4

1094.6

789.3

766.9

724.2

882.7

1201.9

1205.7

976.8

830.0

913.6

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2. Soil Temperature. The analysis of soil temperature under apple stands and treeless areas situated on northern slopes revealed the following regularity: during the vegetative period the soil temperature in open glades is considerably higher than that in apple stands (Dzhangaliev 1974b). The mentioned regularity is observed not only in the upper (5–20 cm), but also in the deeper (40–100 cm), soil layers (Table 2.6). During any 24-hour period, the soil in the tree stands was also considerably colder than it was in open places. The greatest temperature difference (from 2.6–3.8°C) was noted in the upper soil layers and in the daylight hours. In the morning and at night, the temperature decreases by 0.8 to 1.5°C. The difference due to depth differences during a twentyfour-hour period was 1.6 to 2.0°C. 3. Air Temperature. The air temperature, the main index for microclimate, during a 24-hour period is determined by solar radiation, degree of slope, vegetative humidity, and other factors (Dzhangaliev 1972). Vegetation greatly influences the air temperature (when other factors are equal), including the extent of tree cover (Table 2.7). At 1450 m altitude in forested northern slopes, the maximum air temperature was 0.6 to 1.0°C lower than on open slopes. Similarly, the mean monthly minimum temperatures differed by 0.1 to 0.8°C in the forested and open slopes. A converse picture was observed regarding the daily air temperature changes on open and forested areas, that is, it was colder on open plots (1.5–2.0°C). These differences were most distinctly revealed during the daylight hours from 10:00 until 18:00 h. Minimum temperatures at all heights on the trees (1.5, 3.0, and 9.0 m) were observed before sunrise at 4:00 to 6:00 h, and the maximum temperatures at 14:00 to 16:00 h. The Table 2.6. The influence of apple stands on an average ten-day period of soil temperature, Zailijskei Alatau, 1962, 14 hours per measurement. Average temperature (°C) June Area

August

III

I

Open area Apple stand Difference

23.3 17.3 6.0

22.3 18.3 4.0

Open area Apple stand Difference

19.2 14.8 4.4

20.0 15.6 4.4

II

September III

I

II

5–20 cm depth 23.2 21.2 18.0 16.9 4.0 4.3

20.1 16.5 3.6

18.5 15.7 2.8

40–100 cm depth 20.6 20.0 15.6 15.5 5.0 4.5

18.7 15.2 3.5

17.7 14.8 2.9

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Table 2.7. The influence of tree cover on the mean monthly extremes of air temperatures, Zailijskei Alatau, 1963. Temperature on northern slope, 1450 m (°C) Non-forested

Forested

Month

Min.

Max.

Min.

Max.

May June July August

8.2 12.1 14.4 12.0

17.5 22.3 24.9 23.0

7.6 12.0 13.6 12.4

17.7 21.7 24.0 22.0

daily air temperature change in apple stands is more gradual than it is on glades. Temperature drops are the greatest near the soil surfaces and this causes low humidity in the forest. The surface soil layer in the forests reaches lower minimums than in open spaces. 4. Air Humidity. The relative air humidity is closely related to temperature. As the temperature decreases, with other conditions constant, the relative humidity increases. Its maximum indexes occur during the night and in the early morning, but the minimums occur in the hottest time of the day. In apple stands, the relative air humidity during a twenty-fourhour period is 5 to 10 percent higher than it is on forest-free areas. 5. Wind. The wind direction during any twenty-four-hour period on all slopes is rather distinct and is determined by the mountain-valley circulation. In the daytime from 10:00 until 18:00 h, the wind blows upward across the slopes, that is, from the north to the south, and at night, in the opposite direction, The maximum wind velocity occurs in the evening and the minimum at night. In the forest-free areas, the wind is stronger than in the forest. The maximum wind velocity at a 9 m height above the soil surface on an open plot is 3.5 m/s., in forest, 1.4 m/s., and near the soil surface, the wind velocity decreases. Adjacent to the under-tree crowns during the daytime hours, at a 1.5 m height, its velocity does not exceed 0.9, and at 3 m height 1.1 m/s. Thus, climatic conditions are to a great extent regulated by apple community interactions. A large amount of the precipitation, draining down from the upper plots in spring, summer, and autumn are slowed by forest associations which are promoted by the structure and water-holding capacity of the soil, and by thick woody roots. All of this water then flows into the basins of numerous mountain rivers with their wide-branching tributaries that begin on the mountain slopes. Closely interlaced shoots of numer-

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ous kinds of bushes and the rather vigorous crowns of apple stands, along with their developed skeletons, fine twigs and foliage canopy, form a firm framework. This framework protects the soil cover and the soil from destruction by large raindrops and snowdrifts. Well-developed and interlaced root systems of these communities preserve mountain soils from erosion and landslides. Fruit plant associations have a great waterregulating influence. First, in winter, snow masses accumulate in them and thaw gradually in the spring. Second, along with fruit plantings are large areas of shrub communities, which have closed canopies (numerous sweetbriars, honeysuckles, barberries, blackthorns and others) and grass vegetation. These absorb a considerable amount of moisture, which comes from surface outflows and precipitation. In A. A. Uranov’s opinion (1974), a significant role of the vegetative cover is due to the fact that it returns most of its evaporating water to the atmosphere. This process is a special characteristic belonging to forests. Besides, the forest creates the conditions for a good accumulation of rainwater and this protects the soil from surface erosion. Thus, fruit plant communities create their own microclimate. Under their influence, temperature conditions, air humidity, soil moisture distribution, and wind velocity can change. 6. Apple Growth and Development in Relation to Meteorological Conditions. Observations were made on the northern slope of Zailijskei Alatau at altitudes of 1050, 1350, and 1900 m and on the northern and the southern slopes of Dzhungarskei Alatau between 1100 and 2000 m (at every 100 m). At established apple plots, climatic phases were recorded: times of bud-breaking, flowering (beginning, full flowering, end of flowering), shoot growth (beginning, end), fruit ripening, and defoliation. The times of these climatic phases were determined by the methods, suggested in the 1963 Manual for Hydrometeorological Stations and Posts, Agrometeorological observations. Part 1, 1, Leningrad 1963. These investigations revealed some apple growth and bearing characters affected by the different altitude zones of Zailijskei and Dzhungarskei Alatau. 7. Climatic Phase Changes as Affected by Different Altitude Zones. The dates of climatic phases begin to change significantly in relation to the height of the location, and within each height zone according to the time of year, depending on the developing weather conditions (Table 2.8). Thus, at 1050 to 1900 m altitude, the differences in the beginning dates of flower bud-breaking are 8 to 10 days, the beginning of flowering, 20 to 26 days, flowering ending 19 to 24 days, and the fruit ripening period,

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Table 2.8. Beginning dates of the main wild apple climatic phases according to altitudes and years. Altitude Flowering above sea level Bud(m) Year breaking Beginning Full

Shoot growth End

Beginning

End

Beginning of fruit ripening

1050

1966 Apr. 13 1967 Apr. 14

Apr. 30 Apr. 28

May 3 May 10 Apr. 30 May 8

Apr. 30 May 5

Jun. 17 Jun. 14

Aug. 24 Aug. 23

1350

1966 Apr. 14 1967 Apr. 19

May 7 May 5

May 14 May 22 May 6 May 17

May 7 May 6

Jun. 17 Jun. 16

Aug. 30 Aug. 25

1900

1966 Apr. 23 1967 Apr. 22

May 26 May 18

May 27 Jun. 4 May 19 May 29

May 26 Jun. 18

Jun. 26 Jun. 24

Sept. 14 Sept. 14

21 to 22 days. For each 100 m rise, bud-breaking was delayed on an average of 1.0 day, while flowering and fruit ripening were delayed for 2.5 to 3.0 days. The vegetative period, that is, from the start of bud-breaking to the start of fruit ripening, at a height of 1050 m is 133 days, at 1900 m, 144 to 145 days. Depending on the meteorological conditions that occur during the year, the differences in the beginning dates of the phenological phases was 5 days at an altitude 1050 m and 1 to 8 days at altitudes between 1350 and 1900 m. The greatest divergence was noted in the flowering phase. The initiation of climatic phases of apple at different altitudes is closely related to the total temperature accumulation (heat units). A comparison of the dates of the bud-breaking phase, beginning with the transition date of air temperature in the spring through different limits (0.5 and 8°C), showed that beginning date of this phase is approximately the same as the transition date for the mean daily air temperature of +5°C. Thus, at the altitudes of 1050 and 1350 m in 1966 and 1967, the divergence was 3 to 4 days, that is, the temperature transition took place 3 to 4 days earlier than the beginning of bud-breaking. At the altitude of 1900 m, this regularity was not observed. At this altitude, this phase begins at a lower temperature. Wild apple begins to bloom at the end of April or at the beginning of May between altitudes of 1050 to 1350 m and at the end of May at 1900 m (Table 2.9). The period from bud-breaking to the beginning of flowering at 1050 m is 14 to 17 days, and at 1900 m, 26 to 33 days. During this period, the total positive temperatures were 173 to 185 and 137 to 158 degree days respectively. Between the dates of the steady transition of the mean daily air temperature through 0° and the beginning of flowering at the

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Table 2.9. Meteorological conditions characteristic of the period from bud-breaking to the beginning of wild apple flowering. Phase Altitude above sea level (m)

Year

Date of budbreaking

Beginning of flowering

Duration (days)

Mean temperature (°C)

Total positive temperatures (degree days)

1050

1966 1967

Apr. 13 Apr. 14

Apr. 30 Apr. 28

17 14

10.1 13.2

173 185

1350

1966 1967

Apr. 14 Apr. 19

May 7 May 5

23 16

8.2 12.3

189 198

1900

1966 1967

Apr. 23 Apr. 22

May 26 May 18

33 26

4.1 6.0

137 158

height of 1050 m, the accumulated positive temperatures were 304 to 314 degree days, at 1350 m, 222 to 289 degree days, and at 1900 m, 174 to 194 degree days. Thus, in high mountain regions (at 1900 m), the wild apple begins flowering at a lesser accumulative temperature (by 120–130 degree days) than at 1050 m. The flowering period lasts 9 to 15 days. During this period at the height of 1050 m, positive temperatures accumulated at the rate of 112 to 117 degree days, at 1350 m, 118 to 121 degree days, and at 1900 m, 103 to 129 degree days (Table 2.10). The table data point to a constancy of the total accumulated temperatures, which are necessary for the initiation of each definite phase at a specific altitude but also to its sharp decrease with an increase in mountain height. Thus, the difference in the total positive temperatures for the period from temperature transition through 0° to the beginning of flowering between the 1050 and 1900 m altitude in 1966 was 130°C, and in 1967, 120°C; and at the end of flowering, 144 and 104°C respectively. Significant differences in the total temperatures according to altitude were also observed in the rest climatic phases of apple. The temperature levels during the period from bud-breaking to flowering start, depending on the habitats, are also different. The difference between the air mean daily temperatures at 1050 and 1900 m was 6 to 7°C (Table 2.10). By following climatic phases, the analysis of shoot growth periods corresponds with the regularities that were observed with flower initiation already described. These regularities include total temperature constancy, necessity for phase initiation within a specific altitude, decreasing values with an increase in mountain height, and a reduction in the interphase period with a rise in the temperature level (Table 2.11).

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Table 2.10. period.

Meteorological conditions characteristic during the apple flowering

Total degree days Altitude above sea level (m)

Flowering

Mean Duration temperature (days) (°C)

Year

Beginning

End

1050

1966 1967

Apr. 30 Apr. 28

May 10 May 8

10 10

1350

1966 1967

May 7 May 5

May 22 May 17

1900

1966 1967

May 26 May 18

Jun. 4 May 29

Flowering Beginning

Full

End

12.9 11.2

304 314

336 340

420 427

15 12

8.0 9.9

222 289

261 235

343 407

9 11

11.3 11.7

174 194

181 199

276 323

8. Setting Stages and Differentiation of Apple Flower Buds. The initiation and differentiation processes of fruit flower buds in connection with the ecological conditions of mountain regions at various height zones had not been studied. These dates of these stages are conditioned by the complex of environments in which the temperature under optimal moisture is one of the leading factors. Therefore, when studying this problem, we paid special attention to the temperature of environments. The investigations were conducted at the altitudes of 1050, 1350, and 1900 m. In order to determine the start of the initiation of apple flower buds, from June to September samples were taken and flower disbuddings were made. In June and at the end of July, samples were taken once every ten days and at the stage of the differentiation process (from the second Table 2.11. Meteorological conditions characteristic during the period of wild apple shoot growth.

Altitude above sea level (m)

Total degree days Shoot growth Year

Beginning

End

Period duration (days)

Mean temperature (°C)

Above 0°C

Above 5°C

1050

1966 1967

Apr. 30 May 5

Jun. 17 Jun. 14

48 39

12.5 15.0

603 587

383 399

1350

1966 1967

May 7 May 6

Jun. 17 Jun. 16

41 41

12.7 12.8

512 531

327 333

1900

1966 1967

May 26 May 18

Jun. 26 Jun. 24

31 37

12.9 11.4

402 427

254 239

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half of July to September), every five to eight days. The condition and the bud development stage were recorded after microscopic examination according to M. S. Navashin’s (1936) method. The results of these investigations showed that apple flower bud differentiation under the conditions of Zailijskei Alatau begins 29 to 34 days after the end of the shoot growth period (Table 2.12). At the altitudes of 1050 and 1350 m, flower bud differentiation began in mid-July, and at 1900 m, at the end of July. The difference between the beginning dates and ending of bud differentiation at these heights in 1965 was 13 days and in 1967, 10 days. In the period from shoot growth termination to the initiation of bud differentiation, the mean air temperature at 1050 and 1350 m was 16 to 19°C, and at the height of 1900 m, 13 to 14°C. The first signs of the beginning of bud differentiation appeared following shoot growth termination at 1050 and 1350 m, only after total positive temperatures had accumulated 543 to 589 degree days, and at 1900 m, 425 to 492 degree days. The recorded observations support the argument that time periods of bud differentiation initiation may be influenced by total temperatures (heat units). The beginning of bud differentiation under mountain conditions starts within 30 to 35 days after shoot growth ends. The recorded observations on vertical zones and years showed that the time period of flower bud differentiation in the summer–autumn seasons vary. In 1965 at altitudes of 1050–1350 m, the time period consisted of 42 days and at 1900 m, it was 10 days shorter. Under the hot conditions of 1967, which had higher air temperatures, these differences were three Table 2.12. Meteorological conditions characteristic during the period from shoot growth termination to the initiation of differentiation of wild apple flower buds. Total temperatures (°C)

Phase Altitude above sea level (m)

Year

Shoot growth End

Bud differentiation beginning

Period duration (days)

Above 0°C

Above 5°C

Mean temperature (°C)

1050

1965 1967

Jun. 10 Jun. 14

Jul. 13 Jul. 17

33 33

543 589

378 424

16.4 17.8

1350

1965 1966 1967

Jun. 15 Jun. 17 Jun. 16

Jul. 13 Jul. 19 Jul. 19

29 32 33

545 547 572

405 397 407

18.8 18.2 17.3

1900

1965 1967

Jun. 22 Jun. 24

Jul. 26 Jul. 27

34 33

492 425

322 260

14.4 12.8

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Table 2.13. Meteorological conditions during the summer–autumn differentiation stage of wild apple flower buds. Altitude above sea level (m)

Differentiation stage

Total degree days

No. days with temperature higher than

Year

Beginning

End

Above 0°C

Above 5°C

5°C

10°C

12°C

15°C

1050

1965 1967

Jul. 13 Jul. 17

Aug. 24 Aug. 23

796 670

586 485

42 37

42 37

41 37

38 28

1350

1965 1967

Jul. 13 Jul. 19

Aug. 24 Aug. 29

723 706

513 501

42 41

42 38

42 38

36 30

1900

1965 1967

Jul. 26 Jul. 27

Aug. 26 Aug. 31

465 431

312 256

32 34

27 20

27 20

14 9

to seven days less (Table 2.13). Depending on the meteorological conditions, the differentiation process at 1050 m height proceeded for five days, and at 1900 m, for two days. In the summer–autumn period, as in the spring, for flower bud differentiation the relationship between total temperatures and the termination of differentiation was recorded. In 1965 at elevations of 1050 and 1350 m, during the whole period of flower bud differentiation, the mean daily air temperature for almost all 42 days, was not lower than 12°C. The temperatures actually exceeded 15°C for 38 and 36 days at these elevations. At 1900 m, during this period, the temperature did not even reach 10°C everyday. Exceptionally few days were observed with a mean daily air temperature of 15°C (Table 2.13). An analysis of the 1967 data pointed to an even greater spread in temperature levels according to elevation. 9. Evaluation of Thermal Resources of the Territory. By using agrometeorological methods of analysis, the observed data on apple heat requirements in relation to phenological phases and vertical zones allowed us to evaluate the thermal resources of mountain region territories. This knowledge is necessary when new apple introductions are described. Such a complex evaluation of these investigated regions has resulted in an explanation for the first time. In Zailijskei Alatau, because of radiation differences at different altitudes, the spectral composition of solar light and some other factors (including fruit plant heat requirements) are variable. At higher mountain elevations, this index decreases. For instance, at a height of 900 m, from the start of vegetative growth until fruit ripening, the autumn–

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winter apple trees are exposed to mean accumulated effective (above 5°C) temperatures (heat-units) of 2000°C, and at 1700 m, only 1450 to 1500°C units. The difference in the total heat necessary for fruit ripening of autumn and autumn–winter apples, between the altitudes of 1550 to 1600 and 900 m, is 20 to 25 percent. A comparison of the total heat at points of equal height shows that the Zailijskei Alatau territory is richer in heat reserves than Dzhungarskei Alatau. The heat resources of Zailijskei Alatau regions, up to 1450 m, are exceptionally favorable for apple growing. Within the altitudes of 1500 to 1600 m, heat resources also enhance apple productivity, but above 1700 m in most years, only the early ripening apples may ripen. In Dzhungarskei Alatau regions, situated not above 1200 to 1300 m, heat is enough for the full development of summer, autumn and autumn– winter apples. Within the altitudes of 1300 to 1400 m, apple is provided with sufficient heat for the completion of the full vegetative cycle no more than six to seven years out of ten. Thus, the most favorable climatic conditions are observed within the altitudes of 1100 to 1500 m. Apple stands growing under extreme conditions present a special opportunity. Here, it is most rational to select stress-resistant forms. Winter-hardy types exist at the limits of apple growing (above 1700–1900 m). Also, single trees with a shorter vegetative period of development and with winter stages of fruit ripening are seen here. Drought-resistant types can be found in the low borders of steppe zones at a height below 900 m and in very dry conditions of growing where temperature inversions and moisture minimums are sharply expressed. B. Dependence of Apple Stands on Soil Conditions The forest, as a stable long-lasting community, forms together with its environment a correlative unit in which a substance cycle consisting of mineral nutrients and carbon dioxide and an energy transformation are in a state of dynamic balance. This unit, formed by plant communities and their environments, according to V. N. Suckachev (1945), is a biogeocoenosis, the inanimate part of which (ecotop) consists of atmosphere (climatop) and soil (edaphotop), and the animate part (biocoenosis) includes the vegetation (phytocoenosis), the microorganisms (microbiocoenosis) and the animal populations (zoocoenosis). It is natural that the basis of biogeocoenosis, which plays the main role in the process of the biological substance cycle, is made up of autotrophic plants, which form organic substances and heterotrophic organisms (fungi and bacteria),

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that decompose them. Forest biogeocoenoses are most complex and are highly influential in the nature. If we take into consideration that the forest represents the most powerful ‘life lamina’ (by V. N. Vernadsky) and that in other biogeocoenosis the influence of their vital substances on the inanimate nature is not so great and various as it is in the forest, then it would become clear that the forest has an exclusively great role on our planet’s existence and on the life of mankind. (The Program of Biogeocoenosis Investigations. 1966. Moscow)

Recently, great attention has been paid to investigating the interrelationships between plants and the soil, because the biological cycle of these building blocks is determined by the geochemical activity of living organisms. V. V. Dokutchaev (1899) in writing about the vegetative role of changes in soil fertility, attached great importance to forest type. In particular, he pointed out that, in the future, one would be able to distinguish soils under oak, lime, or birch trees and other forest groups. The forest influence on soil chemistry and development is independent of the soil’s origin. Either an artificial or natural change in vegetation, according to I. V. Tyurin’s data (1930), leads to a change in the physical and chemical characteristics of soils and in the soil morphology. Forest influence on soil is different. S. V. Sonn (1954), V. N. Mina (1954), D. F. Sokolov and S. S. Frantsevich (1954) pointed to soil’s enrichment of nitrogen and ash elements under forests and to an improvement of its water-holding properties. At the same time, S. I. Korzhinsky, V. V. Gemmerling and K. D. Glinka (Rosanova 1960) and later N. N. Stepanov (1937), Weir (1940), and Shawarby (1952) noted that rapid chernozem degradation occurred when under the influence of forest vegetation. P. S. Pogrebnyak (1948) and I. P. Pokhiton (1953, 1954) showed that the planting of forest vegetation on chernozem soils leads to some chernozem degradation, though their poor soil formation is not regressive in character. On the other hand, poor soil promotes greater mobility of a number of nutrient soil elements, which improves their potential absorption by wood vegetation under conditions of little moisture (Pogrebnyak 1970). A great number of ash elements are involved in the biological cycle. Broad-leaved forests prevent the process of poor soil development under their canopies and promote ash element accumulation in the upper soil horizon (Remezov et al. 1949; Evdokimova 1955). Soil enrichment of nutrient substances occurs from fallen litter. Annually, about 4 t/ha of oak leaves fall on chernozem surfaces (Afanasjev 1966). Such a mass

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of organic substances undoubtedly influences the water-holding and chemical properties of soils. The nature of the influence of woody vegetation depends on the composition, age, and density of the wood stands (Zonn and Kusmina 1960, 1964; Soloviev 1960, 1967; Metz 1954; Ovington 1954; Pelisek 1962). Data described in these publications are from broad-leaved forests of the European part of the USSR. Studies by D. G. Vilensky (1946), Ju. A. Liverovsky et al. (1949), P. A. Letunov (1953), and A. N. Rozanova (1953), show a positive role of walnut forests in Southern Kirghizia in the development of soil fertility. As far as apple forests are concerned, we did not find similar investigations in publications. The problems of our investigation were the following: studying the intercorrelations between apple communities (natural and artificial) and their soil conditions; determining the quantity and quality of differences of nitrogen and ash elements that return to soil with annual litter fall for different-density and different-aged apple woods; and identifying changes in water-holding and the chemical properties of the soil, which take place under the plant community canopy. In order to study the influence of apple forests on soil processes, a number of trial areas of 400 m2 each were laid out and model trees were chosen. Before laying out the experimental areas, the soil homogeneity was first tested from soil borings taken from eight to ten points using the following indexes: depth of the humus and compact soil horizons, depth of the calcium carbonate layers, and the humus content. The apple trees were divided according to age into groups: 20 to 40, 40 to 80, and 80 to 100 years. The last two groups were predominant in the tested conditions. For an investigation of the influence of different densities of apple trees on the soil properties of apple forests, densities of 0.3, 0.5, and 0.5 to 0.7 were selected. Treeless plots (glades, arable lands, and wastelands) of 400 m2 in size at a distance of 150 m from the experimental areas were studied for comparison. All trial areas were laid out in natural plant communities in middle mountain nonirrigated zones at an altitude of 1250 to 1400 m, on the middle part of slopes having a 15 to 20° incline and under conditions of smooth microrelief. On artificial plantations, experimental areas were laid out in irrigated zones at the altitude of 800 to 900 m. The laying out of trial areas and the model tree choices were conducted according to the method of N. P. Remesov et al. (1959). Chemical analyses of soils and plants were carried out at the agrochemistry laboratory of the Research Institute of Fruits and Viticulture of the Ministry of Agriculture of the Kazakh SSR, at the laboratory of soil

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chemistry and physics of the Soil Science Institute of the Academy of Sciences of the Kazakh SSR, and at the laboratory of the fruit plant department of the Main Botanical Garden of the Academy of Sciences of the Kazakh SSR. In Zailijskei Alatau, deep and medium-deep, strongly leached chernozems of the northern exposure (fresh growth conditions) were investigated, and in Dzhungarskei Alatau, strongly leached medium-deep (fresh growth conditions) and weakly leached (dry growth conditions) chernozems. The soil cover of wood forests (growing at the main Botanical Garden of the Academy of Science of the Kazakh SSR) consists of dark chestnut leached soils.

1. Chemical and Water-holding Properties of Soil and Their Changes Under the Influence of Apple Forests. Apple forests positively influence the properties of mountain chernozems by enhancing their potential fertility. In regard to the organic content of soils in apple forest belts, the humus content was in direct proportion to the density and age of the forest stands. The highest content (14.0–16.6%) was in soils of 80 to 100 year-old apple forests with a stand density of 0.5 to 0.7 (Table 2.14). The great depth of the humus horizon was also noted. It is evidently related to the parent soil-forming rock (loess soils) which allows for deep penetration and fixation of the humus. These humus reserves show especially clearly the differences between mountain chernozems under apple forests and in treeless areas. Thus, the total humus reserves in a meter layer of 80 to 100 year-old apple stands with a 0.5 to 0.7 density are more than 500 t/ha, and in the glade layer, 410 t/ha. It is natural that because of this deep humus accumulation in the mountain chernozems beneath apple forests, there is a relatively high nitrogen content in them also of 0.70 to 0.97 percent (Table 2.14). The carbon:nitrogen ratio in the upper horizons was 9.2 to 10.6, and decreased with depth to 6.3 to 7.4. This illustrates the great nitrogen increasing influence of the humus content of the lower horizons. The absorbing complex of mountain chernozems was fully base-saturated, since the saturation degree in the upper horizon was 85 to 95 percent. The total base-saturation was between 33.9 to 47.7 meq. in 100 g of soil. The main elements in the absorbing complex were Ca (94%) and Mg (5.2%). The Al and H contents were insignificant. In the upper horizons of chernozems of apple forests, soils are saturated by exchangeable Ca that is supplied by an intensive decomposition of apple fall litter which is highly rich in Ca. The reduction of exchangeable Ca and Mg in the lower horizons beneath apple trees is probably related to the deep

95 92 90 92 93 95 95

Plot 4. Applewood, 80-year old (density 0.7). Strongly leached deep chernozem 14.00 0.81 10.0 42.4 5.3 47.7 0.01 none 7.56 0.47 9.3 25.0 4.5 29.5 0.01 0.02 3.72 0.25 8.7 22.2 3.6 25.8 0.01 none 2.89 0.16 10.7 20.6 4.4 25.0 0.01 none 1.71 0.11 9.0 19.8 4.0 23.8 0.01 none 1.07 0.08 7.4 20.6 4.4 25.0 – – 0.96 – – 20.4 2.6 23.0 – –

2–12 20–30 40–50 60–70 80–90 100–110 130–140

N

95 91 89 89 92 87

Al

Plot 5. Applewood, 80-year old (density 0.5). Strongly leached deep chernozem 16.60 0.97 9.9 42.4 2.0 44.4 0.02 none 8.36 0.44 11.1 23.8 4.9 28.7 0.01 none 3.09 0.22 8.2 21.4 4.0 25.4 0.01 0.01 1.84 0.16 6.6 20.6 2.8 23.4 0.01 0.01 1.65 0.09 10.3 20.6 4.9 25.5 0.01 0.01 0.71 0.07 5.8 12.9 1.2 14.1 – –

Total

1–10 20–30 40–50 60–70 80–90 130–140

Mg

Saturation degree (%)

85 78 80 88 94 98

Ca

(Sokolov method)

Plot 3. Applewood, 80-year old (density 0.3). Strongly leached deep chernozem 12.83 0.70 10.6 31.1 2.8 33.9 0.02 none 6.68 0.35 7.7 16.2 4.1 20.3 0.01 none 3.38 0.27 7.4 16.2 2.0 18.2 0.01 0.01 2.20 0.16 7.7 17.4 2.1 19.5 0.01 0.01 0.98 0.06 9.0 15.8 3.2 19.0 0.01 0.03 0.53 – – 8.9 2.0 10.9 – –

C:N ratio

(Shmuk method)

1–10 20–30 35–45 60–70 90–100 130–140

N (%) (Kjeljdal Method)

Absorbed cations in 100 g soil (meq.)

(continued)

6.9 6.7 6.6 6.7 6.7 7.5 7.7

6.9 6.8 6.6 6.6 6.7 7.1

6.8 6.7 6.7 6.5 7.1 7.1

pH water suspension (potentiometer)

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Zailijskei Alatau, 1963

Sample depth (cm)

Humus, (%) (Tyurin method)

Apple forest influence on the chemical structure of soils.

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Table 2.14.

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138 12.70 5.38 2.91 2.35 1.36 1.07 0.43 11.90 7.46 4.01 3.12 1.75 1.09 0.86

Sample depth (cm)

0–10 20–30 40–50 60–70 80–90 100–110 130–140

0–10 20–30 40–50 60–70 80–90 100–110 130–140

Total

Al

N

0.02 None None 0.01 None – –

Mg

Plot 11. Arable land. Strongly leached deep chernozem 0.74 9.3 35.7 1.2 36.9 0.02 0.52 8.2 27.5 2.4 29.9 0.01 0.32 7.3 22.2 4.4 26.6 0.01 0.19 9.1 22.6 2.4 25.0 0.01 0.11 8.9 16.9 5.8 22.7 0.01 0.08 7.7 21.4 0.8 22.2 – – – 25.4 2.4 27.8 –

Ca

(Sokolov method)

0.01 0.01 0.01 0.01 0.01 0.01 0.01

C:N ratio

(Shmuk method)

Plot 10. Glade. Strongly leached deep chernozem 0.79 9.3 33.1 3.2 36.3 0.02 0.37 8.4 22.2 2.0 24.2 0.01 0.23 7.4 21.8 2.4 24.2 0.01 0.11 12.0 20.2 1.6 21.8 0.01 0.07 11.1 21.8 3.9 25.7 0.01 0.05 12.4 23.0 1.6 24.6 0.01 – – 21.8 1.8 23.6 0.01

N (%) (Kjeljdal Method)

Absorbed cations in 100 g soil (meq.)

88 87 86 86 87 92 96

85 72 87 89 94 93 96

Saturation degree (%)

– – – – – – –

6.6 6.6 6.7 6.7 6.8 7.0 7.6

pH water suspension (potentiometer)

4:19 PM

Humus, (%) (Tyurin method)

(continued)

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Table 2.14.

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4.5 3.3 2.3 1.4 1.0 3.1 2.3 1.8 1.9 1.1 2.9 3.0 1.8

0–10 15–25 35–45 55–65 75–85

0–10 20–30 35–45 55–65 72–77

0–10 15–25 35–45

9.56 5.05 3.56 2.27 1.36

– – – – – – – –

– – – – –

Plot 2. Grassland. Dark-chestnut leached soil 0.14 12.7 16.0 2.0 18.0 0.13 10.7 15.0 1.5 20.0 0.10 10.9 15.0 1.1 16.1 0.10 10.7 14.0 1.0 15.0 – – 9.5 2.5 12.0

Plot 5. Arable land (potatoes). Dark-chestnut leached soil 0.17 10.1 14.5 1.0 15.5 – 0.16 10.8 14.5 1.5 16.0 – 0.12 8.7 12.5 1.5 14.0 –

– – – – –

– – – – –

0.13 0.04 0.03 0.02 0.02

Plot 1. Apple orchard. Dark-chestnut leached soil 0.24 10.7 16.4 4.0 20.4 0.15 12.2 18.0 2.0 20.0 0.11 11.1 15.5 1.5 17.0 0.09 9.6 14.0 4.0 18.0 – – 12.5 1.0 13.5

Plot 15a. Glade. Strongly leached medium deep chernozem 0.63 8.8 30.3 4.4 34.7 0.06 0.39 7.4 24.7 3.2 27.9 0.05 0.25 8.3 23.5 2.8 26.3 0.05 0.18 7.1 21.5 4.0 25.5 0.05 0.15 5.2 18.7 2.8 21.5 0.04

– – –

– – – – –

– – – – –

85 85 84 91 96

94 90 87 88 90

7.7 7.7 7.6

8.2 8.1 8.5 8.2 8.1

7.7 7.8 8.1 8.3 8.9

6.2 6.2 6.5 6.7 6.8

6.6 6.8 6.8 6.9 7.0

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0–10 20–30 40–50 60–70 90–100

Plot 15. Applewood, 80-year old (density 0.8). Strongly leached medium deep chernozem 2–12 12.39 0.78 9.2 34.3 4.0 38.3 0.22 0.36 20–30 6.23 0.46 7.9 24.3 4.8 29.1 0.15 0.39 40–50 3.52 0.24 8.4 17.5 2.4 19.9 0.06 0.05 60–70 1.71 0.13 7.6 17.1 2.8 19.9 0.07 0.16 90–100 1.51 0.14 6.3 17.1 2.0 19.1 0.05 0.13

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penetration of root systems of different wood types and with a more intensive absorption of Ca and Mg at these depths. A. N. Rozanov (1953), B. V. Nadezhdin (1954), A. I. Pavlenko (1955) and I. M. Rozanova (1960) also pointed to a similar change in the character of exchangeable bases in field and in forest chernozems. In regard to the high base-saturation of mountain chernozems, the soil pH is close to neutral; the pH of a water suspension is between 6.6 to 6.9. Little difference in the soil reaction beneath apple stands and under treeless areas was observed. Even with an increase in the age of apple populations, soil acidification was not noted. The total P content in mountain chernozems and its distribution in soil beneath apple stands and in treeless areas was similar to that of N. The total amount of K in apple stand soils and on treeless areas was not very different. The K2O content in glade grass vegetation was rather high (4.76%) and because of this, a significant amount of this element annually returns to the soil. These data from chemical analyses of mountain chernozems indicate that apple wood has a favorable influence on soil fertility, as shown by an increase of content of humus, N, P, and Ca. Thus, it is evident that an accumulation of humus and nutrient substances in mountain chernozem soils is closely connected with the age and density of apple stand communities. In order to measure the apple stand influence on soil podzolization, the humus composition was studied (Table 2.15). The humus source under forests is fallen litter of wood and grass vegetation, annually dying off of feeder roots, and different organic residues in the soil. Three groups represent the humus substance of mountain chernozems: humic acids, fulvoacids, and soil humins. Readily available fulvoacids and slightly available humic acids are the most significant substances in humus. Different forest types are characterized by definite ratios between these acid contents CHA : CFA, the higher this ratio is, the more favorable are the edaphic conditions for plant development. In the upper part (0–10 cm) of the mountain chernozem profile beneath apple stands, the highest percent of the initial carbon content is made up of fulvoacids, and the ratio CHA : CFA is 0.76 to 0.93. In lower horizons, the amount of humic acid increases and its ratio is 1.15 to 2.26 (Table 2.15). Nevertheless, an increase in fulvoacid in the upper soil does not promote a more intensive podzol-forming process, because the high ash content and the increasing base content in apple fall litter neutralize the acidic organic substances. A total analysis of the data on apple forest belt soils (Fig. 2.17) points to a homogeneity of their profiles. We failed to differentiate between the horizon, which is impoverished by sesquioxides, and the horizon, which

Dzhungarskei Alatau, 1962

0–10 20–30 40–50 60–70

2–12 20–30 40–50 60–70

0–10 20–30 40–50 60–70

1–10 20–30 40–50 60–70

Nonhydrolysis residues Hydrolysate

Decalcinate

1st

2nd

3rd

Fraction Total

1st

22.0 18.2 16.6 18.6

Plot 10. Glade. Strongly leached deep chernozem 11.8 14.0 11.8 9.0 4.3 8.9 9.7 13.5 18.7 5.4 6.5 9.8 5.0 36.4 5.7 4.8 13.6 4.8 31.3 6.8

25.1 37.6 47.1 42.9

19.5 14.3 10.0 9.1

5.85 3.07 2.06 1.38

Plot 15a. Glade. Strongly leached medium-deep chernozem 24.0 12.1 3.5 16.6 9.1 6.0 31.7 23.7 8.9 3.3 14.4 21.0 5.7 41.1 15.5 12.2 3.9 11.0 31.6 4.9 47.5 21.8 6.3 4.3 6.6 32.1 4.3 43.0

14.2 12.5 11.7 11.3

Plot 15. Applewood, 80-year old (density 0.8). Strongly leached medium-deep chernozem 6.44 23.6 10.5 7.9 16.1 7.1 5.2 28.4 16.3 3.79 25.7 9.6 6.2 12.7 17.8 6.2 36.7 11.7 2.36 22.4 7.2 4.9 8.1 31.9 5.3 45.3 9.6 1.08 24.0 8.4 4.7 4.7 32.8 – 35.7 10.8

7.31 3.30 1.99 1.24

Plot 5. Applewood, 80-year old (density 0.5). Strongly leached deep chernozem 9.19 23.2 14.4 11.2 7.0 10.4 4.8 22.2 16.3 3.75 20.4 11.3 8.5 6.7 18.4 6.8 31.9 11.9 2.00 16.9 8.2 8.7 3.5 34.3 6.9 44.7 8.9 1.27 12.5 6.3 11.7 – 43.0 5.2 48.2 7.4

Initial soil C (%)

8.3 7.2 7.4 11.0

9.3 7.0 5.6 9.9

1.5 5.2 5.3 5.5

5.6 9.2 7.6 9.4

2nd

4.8 6.5 3.4 4.8

5.1 6.1 4.9 4.5

6.2 5.8 5.0 4.7

7.0 6.5 5.2 4.8

3rd

Fraction

Fulvoacids

27.3 26.2 22.5 27.1

30.7 24.8 20.1 25.2

27.2 25.3 20.3 19.3

28.9 27.6 21.7 21.6

Total

1.17 1.56 2.10 1.60

0.93 1.48 2.26 1.42

0.92 1.49 2.31 2.24

0.76 1.15 2.05 2.22

Ch. a: Cf=a

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Zailijskei Alatau, 1963

Sample depth (cm)

Humic acids

Percent of total organic soil carbon

Group and fractional humus compositions in strongly leached mountain chernozems.

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Location

Table 2.15.

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Fig. 2.17. The influence of different-aged apple woods on the total composition of mountain leached chernozems.

is enriched by silica. The sesquioxide content is not high and their movement in the soil profile is very slow. The profile homogeneity is confirmed by the molecular ratio of silica: alumina (SiO2:Al2O3) and by the silica:sesquioxides (SiO2:R2O3) ratio in the parent rock. Ratios are somewhat wider in the parent rock than they are in soil layers, except in the uppermost horizons, which have been enriched by SiO2 in the process of biological accumulation. Great differences in comparison

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with treeless plot soils are also not observed in the distribution of sesquioxides Fe2O3 and Al2O3. Apple stands exerted some influence on the calcium distribution. CaO in treeless plot soil is comparatively evenly distributed along the profile. Under apple stands, the Ca content decreases in the upper meter of thickness but increases in the second meter (Dzhangaliev 1973a). Thus, though mountain chernozems under the influence of apple forests do not suffer great changes, when they were analyzed in terms of the redistribution of total oxide content, especially CaO, a podzolization process did not occur. Our observations are in good agreement with the data of S. V. Zonn and A. K. Aleshina (1953), S. V. Zonn (1954), I. M. Rozanova (1960) and D. F. Sokolov (1962), who did not find any change in the podzolization process under the influence of broad-leaved forests. An investigation of 35-year-old apple orchards (M. sieversii rootstock) on dark chestnut leached chernozem soils agreed with the regularity mentioned above (Table 2.14). In the upper horizon, the greater part of dark chestnut leached soils under grass and potatoes are characterized by a low humus content (2.9–3.1%) and low N (0.14–0.17%), which gradually decreases with the soil depth. The upper sodden layer of similar soils, which develop under long-term apple forests (Table 2.14, Plot 1), is considerably different with a higher humus content (4.5%) and a higher N content (0.24%). In regard to the distribution of humus and N as we go down along the soil profile, special characteristics in relation to our investigative objects are not noted. The absorbing complex of dark-chestnut leached soils is fully base-saturated. In composition, the main elements are Ca (80%) and Mg (18%). The sum of the saturated bases in the upper horizons is between 15.4 and 20.4 meq. in 100 g soil. The greatest sum of exchange bases is in dark-chestnut soils under fruit forests, and the least, under potato. Thus, the above-mentioned tendency is also noted here, that is, the upper soil layers under apples, because of the exchange of Ca and Mg owing to apple litter fall decomposition, are very rich in CaO. The reaction of the soil solution is alkaline that intensifies with increased soil depth. Some comparatively high alkalinity is found in the surface horizons of grassland soils (Table 2.14, Plot 2). The amount of hydrolyzable N in dark chestnut soils is between 42 and 106 mg/kg, and these soils have the greatest amount of N when they are under fruit forest canopies. The amount of mobile P2O5 in the surface layer of dark chestnut soils does not exceed 20 mg/kg. No significant differences in the P2O5 content among the investigated objects were noted. The amount of exchangable K2O in the dark chestnut leached

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soils is quite adequate. The greatest amount of K2O was found in grassland plots (400 mg/kg), with somewhat less in the soils under apple forests (309 mg/kg). The lowest quantity of K2O was noted in soil under potato (260 mg/kg), which is related to a greater requirement and output of K by potato (Dzhangaliev 1973a). The total content of dark-chestnut leached soils is rather homogeneous. However, in soils under fruit forest canopies, a slight increase in SiO2 can be seen within the humus horizon (especially in horizon B) and also an increase in K2O and P2O5. Increased K2O and P2O5 are apparently connected with a biological accumulation of these elements. The amount of sesquioxides is almost unchanged. In lea (grassland) soils, sesquioxides, CaO, SO3, and P2O5 increased. In apple stands, because of a highly intensive ash exchange which is caused by the neutral character of humus substances formed by litter, and also as a result of the root system influence of wood stands, a soil structure improvement takes place. According to agronomic concepts, a good soil structure is finely crumbled and granular with aggregates of 0.25 to 10 mm in diameter and has a spongy mechanically strongly resilient quality. This soil structure study showed that in chernozems under apple stands, a cloddy-fine-grained structure is well developed. In the horizon A, crumbly-fine-grained elements constitute 91.7 percent and cloddy (>10 mm) and silty (<0.25 mm) elements, 5.2 and 2.6 percent respectively (Table 2.16). At lower depths, the structural composition deteriorates. This is explained because the structure-forming processes are closely connected with the accumulation and distribution of organic substances in the soil. It was found that water holding is a function of the quantity and quality of humus in the soil. Maximum water holding coincides with a large humus content and humin compounds in the soil. The sizes of soil aggregates are an index of one or another of the physical soil types when the aggregates are holding water. The coefficients of soil structure (potential ability of the soil to become structured) and the water holding capacity of horizon A are 11.0 and 94.0 percent respectively. The removal of apple stands and soil plowing lead to a sharp reduction of structure water-resistance (water holding capacity) not only in tillaged but in undertillaged horizons. The structure coefficient decreases on an average by 4.4 units, while water-resistance decreases by 1.1 units. With constant tillage of mountain chernozems, the plowed horizon loses its natural structure and acquires a cloddy character. Treeless plots are characterized by a better structure and by a greater water holding capacity of aggregates than plowed lands, but they are still inferior to apple stand soils. The improvement of mountain chernozems depends on the age and density of phytocoenosis (plant

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communities), which, in turn, influence the depth of the most structured chernozems and the aggregate water holding capacity (Table 2.16). The structure, intensity of humus color, and some other morphological characters of dark-chestnut leached soils are strongly dependent on the vegetation growing on them. In this respect, perennial fruit plants considerably influence soils. Dark-chestnut soils developing under them are characterized by a somewhat greater humus horizon depth (A + B = 90 cm), intensively dark-gray color, cloddy-grained structure, and a compact texture of the transitional humus in horizon B. These soils also have visible perforations caused by humus horizon earthworms. The morphological peculiarities of these dark-chestnut soils under potatoes and also lea (grassland) plots differ from similar soils under perennial plant canopies by having less depth of the humus horizon (A + B = 70 cm); by a less intensive humus color; by the presence of a strongly packed under-plow horizon, in almost all cases; by appreciably less activity of earthworms, and consequently by less granularity; and by a crumbly structure of the arable layer. For a more detailed characterization of the structural state of mountain chernozems and dark-chestnut soils, their mechanical texture was determined because it considerably influences the development and the character of soil aggregate formation and characterizes a soil’s potential ability to form agronomically valuable structures. The analysis showed that the soils of mountain leached chernozems consist of heavy chernozem soils and are transformed into loess-like chernozems (Fig. 2.18). Sufficient humus (14.0–16.6%) and a high content of clayey particles (16.0–20.4%) condition the formation of a water-holding structure. On the basis of mechanical and microaggregate analyses (Table 2.17), the dispersion coefficient was calculated. The higher the dispersion factor, the finer the microstructure is and consequently the better the soil structure is. These data indicate a rather high microaggregation character of the mountain leached chernozems. Under apple stands, the dispersion coefficient was 5.3 percent and under glades, 7.5 percent. At deeper depths, this coefficient increases, that is, in lower horizons the soil is considerably less aggregated than it is in the upper humusaccumulated horizons. Relative to the mechanical structure, dark-chestnut leached soils are heavy chernozem, and at deeper levels, they change, first into medium and then into light soils. A sufficient amount of medium and fine sand and a considerable amount of dust (mainly coarse dust) and clayey particles characterize them. Moreover, if the sand fraction exhibits a general tendency to increase with increased soil depth, the clayey particles significantly enrich the humus part of the profile, where the clay content is

146

Zailijskei Alatau, 1963

Wet

Dry

Wet

Structure

11.0 7.7 6.6 3.7 2.5 3.0 2.0

Dry

Plot 4. Apple stand, 80-year old (density 0.7). Deep strongly leached chernozem 2–12 5.2 6.5 87.8 82.5 3.9 8.4 3.1 2.6 20–30 7.4 6.5 79.2 73.0 9.3 13.7 4.1 6.8 40–50 9.4 1.6 79.8 59.0 7.1 23.1 3.7 16.3 60–70 18.7 – 67.1 49.9 11.8 29.4 2.4 20.7 80–90 22.0 – 58.5 42.0 12.6 33.5 6.9 24.5 100–110 23.8 – 65.0 40.8 10.2 31.6 1.0 27.6 130–140 31.3 – 63.5 36.2 3.3 22.4 1.9 41.4

Wet

9.2 6.9 5.7 5.1 2.9 3.2

Dry

94.0 92.2 73.9 74.4 71.8 62.8 57.0

94.0 83.6 78.0 52.7 50.3 35.4

82.3 81.1 77.3 70.1 55.4 38.4

Water resistance

Coefficient

Plot 5. Apple stand, 80-year old (density 0.5). Deep strongly leached chernozem 1–10 4.7 8.0 83.4 78.3 6.8 4.2 5.1 9.5 20–30 11.1 6.3 79.0 66.0 8.4 11.1 1.5 16.6 40–50 14.3 – 78.7 61.2 6.4 18.5 0.6 20.3 60–70 15.7 – 80.1 42.2 3.6 36.0 0.6 21.8 80–90 24.6 – 71.0 35.7 3.4 38.1 1.0 26.2 130–140 23.0 – 73.5 26.0 2.8 36.0 0.7 38.0

Wet

<0.25 (mm)

6.1 5.3 4.3 2.9 2.4 3.4

Dry

1.00–0.25 (mm)

Plot 3. Apple stand, 80-year old (density 0.3). Deep strongly leached chernozem 1–10 12.2 12.8 82.0 67.5 3.9 2.8 1.9 16.9 20–30 13.8 12.8 80.5 65.3 3.5 10.9 2.2 11.0 35–45 16.5 6.2 77.6 60.0 3.7 14.2 2.2 19.6 60–70 21.9 – 68.0 47.7 6.7 28.2 3.4 24.1 90–100 27.4 – 66.4 36.8 4.6 30.5 1.6 32.7 130–140 17.0 – 68.4 26.3 8.8 26.9 5.8 46.8

Sample depth (cm)

10–1.00 (mm)

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Location

10 (mm)

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Fraction size

Screening (%)

Table 2.16. Structured and aggregated compounds of leached mountain chernozems under apple stands and in treeless plots (according to Savvinov’s method).

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29.7 24.0 46.5 46.1 51.1

4.8 3.2 2.6 1.5 0.8

0–10 20–30 40–50 60–70 90–100

Plot 15a. Glade. Medium-deep strongly leached chernozem 5.7 1.5 72.1 58.3 10.6 10.5 11.6 11.0 2.7 68.7 55.0 7.7 18.7 12.6 16.7 1.0 65.1 29.4 7.4 23.1 10.8 33.6 – 58.3 20.3 2.4 33.6 5.7 52.5 – 42.8 11.1 1.5 37.8 3.2

2.5 2.7 1.8 1.5 1.7 1.4 1.7 5.6 3.5 2.1 1.8 1.9

19.7 22.7 26.3 45.4 33.5 32.0 44.3

Plot 11. Arable land. Deep strongly leached chernozem 8.1 69.0 58.5 2.4 13.7 19.6 6.8 67.5 55.0 5.4 15.5 19.3 6.3 61.5 45.5 2.3 21.9 8.4 – 57.0 42.0 4.2 12.6 3.4 – 61.0 34.5 2.5 32.0 2.6 – 55.4 32.5 2.2 35.5 2.0 – 59.4 29.5 3.2 26.2 0.9

5.6 4.4 1.8 1.8 1.5 1.5 1.4

80.7 80.0 45.2 35.4 25.9

91.0 85.5 65.5 48.5 35.0

84.8 81.5 74.0 73.6 56.6 58.6 49.7

82.1 81.0 73.2 47.6 48.2 40.9 40.3

4:19 PM

Plot 15. Apple stand, 80-year old (density 0.8). Medium-deep strongly leached chernozem 2–12 2.1 1.9 74.6 68.0 10.4 11.2 13.9 18.9 20–30 14.0 5.3 68.9 54.6 11.7 13.5 9.4 26.6 40–50 30.2 6.7 57.6 32.7 10.4 10.0 1.8 50.6 60–70 36.3 2.7 55.5 27.0 8.7 25.3 2.5 45.0 90–100 47.0 – 48.3 16.9 2.9 25.8 1.8 57.3

9.0 7.8 27.8 35.4 33.9 40.4 36.5

0–10 20–30 40–50 60–70 80–90 100–110 130–140

17.0 22.7 37.4 36.5 44.1 44.6 40.0

Plot 10. Glade. Deep strongly leached chernozem 8.2 75.0 61.6 9.9 13.2 12.0 6.5 74.0 60.0 7.5 10.8 1.7 – 61.5 45.0 3.3 17.6 1.8 – 62.0 29.5 2.9 34.0 4.5 – 57.0 27.5 2.9 28.4 3.6 – 58.0 23.7 1.9 31.7 2.3 – 57.0 23.0 1.0 37.0 3.0

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3.1 17.2 33.4 30.6 36.5 37.8 39.0

0–10 20–30 40–50 60–70 80–90 100–110 130–140

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Fig. 2.18. Soil mechanical structure under apple forests and on forestless plots: 1 = fine sand (0.25–0.05 mm); 2 = coarse dust (0.05–0.01 mm); 3 = medium dust (0.010–0.005 mm); 4 = fine dust (0.005–0.001 mm); 5 = clay (<0.001 mm).

0.05–0.01 (mm)

Dust 0.01–0.005 (mm) 0.005–0.001 (mm)

Clay <0.001 (mm)

90–100

60–70

40–50

20–30

0–10

– 25.3 – 14.2 – 2.4 – 1.2 – 1.1

Plot 15a. Glade. Medium-deep strongly leached chernozem 7.4 41.7 12.5 22.4 39.5 27.6 3.8 2.6 17.0 32.2 11.6 21.3 39.5 27.6 3.8 2.6 11.4 40.2 11.1 18.7 46.4 36.4 10.7 2.6 13.1 36.2 10.7 18.0 43.3 38.5 6.8 8.2 10.8 38.1 10.0 14.7 49.7 34.7 6.2 5.6

16.0 1.2 17.9 1.2 18.6 1.7 22.0 2.2 25.4 2.7

50.9 – 50.8 – 48.4 – 50.7 – 50.1 –

48.4 – 55.1 – 55.3 – 51.3 – 50.4 –

Fraction sum <0.01

Plot 15. Applewood, 80-year-old (density 0.8). Medium-deep strongly leached chernozem 2–12 – 9.6 46.0 12.8 18.7 16.9 36.2 39.4 20.4 1.5 1.6 0.9 20–30 – 4.0 40.9 14.0 21.3 19.8 24.9 32.8 32.5 5.4 3.3 1.1 40–50 – 5.8 38.9 13.9 19.8 21.6 12.2 37.0 34.7 9.1 5.3 1.5 60–70 – 9.9 38.8 11.3 17.5 22.5 3.2 38.6 40.6 7.2 8.4 1.8 90–100 – 8.1 41.5 11.8 15.3 23.3 2.0 38.4 42.6 8.8 6.6 1.6

Sand 1–0.25 0.25–0.05 (mm) (mm)

(continued)

10.7

10.0

9.2

9.0

7.5

6.9

8.0

6.9

5.5

5.3

Dispersion coefficient by Kachinsky

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Dzhungarskei Alatau, 1962

Sample depth (cm)

Fraction size (%)

Apple wood influence on mechanical (numerator) and microaggregate (denominator) soil structure, percent to absolute

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Location

Table 2.17. dry weight.

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149

150

140–150

100–110

72–77

55–65

35–45

20–30

0–10

140–150

105–115

75–85

55–65

35–44

7.4 4.6 4.9 3.6 3.8 3.8 5.4 5.6 20.0 22.3 0.8 0.9 2.6 2.7

6.4 13.6 5.3 6.8 5.6 5.7 2.9 2.9 0.9 0.7 0.7 1.0 0.5 0.6 4.8 4.1 9.5 5.3 7.5 5.0 8.3 3.5 5.3 3.4 3.4 4.7 5.6 3.7

30.2 0.8 22.6 0.6 21.8 0.6 20.3 1.1 14.3 2.6 14.8 0.8 12.5 0.5

0.2 37.2 3.8 38.9 7.5 43.0 11.6 45.7 11.4 26.7 14.6 42.7 17.1 47.2

Plot 2. Lea. Dark-chestnut leached soil 45.5 13.9 43.9 9.4 39.7 11.1 50.7 0.9 38.6 9.6 40.3 7.3 35.2 7.9 37.6 6.5 25.9 7.3 40.1 4.9 38.4 5.9 45.1 5.8 39.2 6.1 41.4 4.5

0.005–0.001 (mm)

Clay <0.001 (mm)

23.3 1.2 23.5 0.9 21.6 1.4 21.6 0.7 20.1 1.2 20.9 0.8 21.5 1.4

0.05–0.01 (mm)

Dust 0.01–0.005 (mm)

Plot 1. Apple orchard. Dark-chestnut leached soil 6.8 36.0 9.9 11.5 41.6 32.5 6.7 4.4 8.9 34.7 11.6 10.2 43.4 37.2 7.6 4.1 8.4 37.7 12.7 8.5 45.2 37.9 5.9 3.9 6.7 42.5 10.2 10.8 43.1 42.9 5.7 4.7 2.3 40.5 10.7 9.4 44.3 45.2 5.4 3.4 3.0 37.0 10.5 9.8 28.2 57.0 8.5 4.5 1.1 41.0 10.2 6.4 34.3 52.9 6.3 4.5

Sand 1–0.25 0.25–0.05 (mm) (mm)

Fraction size (%)

38.9 – 43.2 – 38.9 – 36.5 – 26.9 – 24.1 – 24.5 –

44.7 – 45.3 – 42.8 – 42.6 – 40.2 – 41.2 – 38.1 –

Fraction sum <0.01

Dispersion coefficient by Kachinsky

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15–25

0–10

Sample depth (cm)

(continued)

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Location

Table 2.17.

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8 to 10 percent higher than it is in the soil-forming rock. The great amount of clayey particles in the profile under fruit forests promotes soil structure improvement. These soils possess a significant amount of microaggregation. The content of water-holding microaggregates in the humus horizons is 35 to 42 percent of the soil weight or 75 to 90 percent of the content of physical clay. Under the influence of apple stands, the air, heat, water, and soil characteristics are significantly improved. On average, the field water capacity of the soil under them, in the upper horizon, reaches 54.1 percent. A meter layer of these soils may contain 415 mm of water. Possessing a large water-holding capacity, the upper soil horizons accumulate all thawing water and summer precipitation, which in turn, determines the existence of apple phytocoenosis in a period of summer drought. Because of insignificant surface outflow, even on sloping plots, soil erosion is hardly ever observed. Mountain chernozems of the apple coenosis belt have high water permeability (about 500 mm for the first hour). In highdensity apple woods, the soil permeability increases (Fig. 2.19) due to better structural composition and porosity. Thus, during the first 10 minutes under all apple woods, soil permeability was, on the average, 1.5 times higher than on arable lands. By the second hour, in spite of a reduction of water absorption speed, its level under apple stands was still 17.0, while on arable lands, it was 9.2 mm. Using the field water capacity and the sesquioxide maximum hygroscopicity, we calculated the active humidity diapason. According to N. A. Kachinsky (1947) “the wider the active humidity diapason and the higher its mobility, the more valuable is the soil.” The calculated diapason of active humidity in the upper soil horizons under apple woods is between 44.9 to 52.5 percent and decreases with depth. However, in the layer 130 to 140 cm deep, its value is rather high (15.7–20.9%), which points to the possibility that apple vertical roots may supply the above-ground parts with water in a summer drought period for a long time and constantly. 2. Accumulation and Ash Composition of Leaf Fall and Litter in Apple Forests. Soils under forests are characterized by a continuous change in their composition and special features over time. Depending on the environmental complex, these changes are able to proceed at different speeds, that is, when the combination of these speeds and the direction of the soil-forming processes are stipulated. The stimulus to begin changes is a biological cycle between forest and soil. The capacity and intensity of the biological cycle are quite unequal for different biocoenosis.

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Fig. 2.19. Apple-wood influence on water permeability of mountain chernozems: 1 = arable land; 2 = apple wood, 80 to 100 year-old (density 0.3); 3 = apple wood, 80 to 100 year-old (density 0.5); 4 = apple wood, 80 to 100 year-old (density 0.7).

The annual amount of ash and nitrogen penetration into the soil is one of the most important parts of the intensity stage of the biological substance cycle. Many authors (Letunov 1953; Ignatieva 1970; Orlovsky et al. 1970) pointed to the important role of grass cover in ash and nitrogen exchanges of high-bonitet forests. As our investigations showed, the amount and composition of the annual above-ground leaf fall under apple woods depend on the forest-vegetative conditions of biocoenosis. In the regions investigated by us, fruit forests are of a park character in which the mean density is 0.4 to 0.5. In apple woods, this density promotes grass cover development. Depending on forest stand density, the leaf fall composition varies. Thus, in high-density apple woods (0.7),

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the amount of woody plant and grass debris are approximately equal, while in thin forests, grass prevails, and on glades, there is only grass fall. Nevertheless, in high density apple woods, a thick grass stand can produce as much as 1200 kg/ha. Mesophyte plants of mountain meadows mainly produce the grass cover in apple woods with fresh growing conditions (humid northern slopes). These include the porous-soddy species Brachypodium silvaticum that dominates here, and broad-leaved grasses, such as the rhizogenous species Urtica dioica and Aegopodium podagraria. In dry growth conditions of apple stands (dry south-western slopes), the cereal grasses Dactylis glomerata, Poa nemoralis, and Milium effusum predominate. The more favorable water-nutrient regime of the strongly leached deep chernozems produces a considerably greater development of phytocoenosis vegetative masses. Annually produced leaf fall in highdensity apple woods delayed the mineralization of organic residues under comparatively moderate temperatures and sufficient air and soil humidity during the vegetative period. This leads to a higher accumulation of humus, N, P, Ca, and other nutrient substances in comparison to thin forests and glades. The composition of the leaf fall differs in the different density apple stands and on treeless plots, which influences the ash composition of the leaf fall and thus, the ash accumulation. It is natural that the ash accumulation and its composition in apple stands with different crown closures and on virgin lands depend not only on the leaf fall mass but also on its composition. The greatest ash amount, 422.5 kg/ha, developed on the soil surface of high-density apple forests (0.7). A lesser amount developed under thin forests (0.3), 313.6 kg/ha and on treeless plots, 262.6 kg/ha. In the high-density coenosis apple (0.7), the ash was 50.4 percent, aspen 12.8 percent, and grasses 33.8 percent of the total amount of 422.5 kg/ha. In apple woods with 0.3 density, the ash content of apple leaf fall decreased by 15.5 percent, while the ash content of aspen increased by 27.4 percent, and grass by 57.1 percent. The organic residues of woods and grass vegetation lying on the soil, relative to their content of N and ash elements, differ from each other. Higher N is in the leaves of Crataegus altaica (2.35–2.50%), Padus racemosa (2.32%), Rhamnus cathartica (2.40%), and Ribus meyeri (2.63%). The leaves of Crataegus altaica (0.46%), Abies sibirica (0.45%), and Rubus idaeus (0.45%) are rich in P2O5. The highest content of K2O is in the leaves of Padus racemosa (2.16%), Crataegus altaica (2.06%), Rubus idaeus (3.07%), and especially in Ribes meyeri (4.44%). A considerable amount of CaO and MgO

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accumulates in the leaves of Ribes meyeri (6.70 and 0.44% respectively) and Populus tremula (4.10 and 0.25%). A great accumulation of SiO2 occurs in the conifer Abies sibirica (1.77%), in leaves of Populus tremula (1.88%), Rhamnus cathartica (2.10%), and Ribes meyeri (2.23%). The greatest amount of N and ash elements is in apple leaves, which are growing on strongly leached deep chernozems, and somewhat less in apple leaves growing on medium-deep leached chernozems. The content of these elements in leaves of high-density wood stands exceeds those in leaves of thin wood stands. Thus, the leaves of wild apples which possess a high ash content, CaO (3.7%), K2O (2.8%), N (2.8%), and P2O5 (0.6%) promote the biogenic accumulation of these elements in the soil. An admixture of pure apple woods along with adjacent species, especially Crataegus altaica and Ribes meyeri, which possess a high ash content, increases the ash element accumulation in the soil. In litter-falls of apple phytocoenosis, the content of N (1.4%), P2O5 (0.3%), and K2O (0.4%) is lower, but that of CaO (5.2%), MgO (0.4%), SiO2 (2.1%), Fe2O3 (0.3%), and Al2O3 (0.3%) is higher than in green leaves. Grass fall contains N (1.2%), which is approximately the same as in apple leaf falls, but it contains less CaO (2.1%) and P2O5 (0.2%). The K2O (0.6%) content in grass fall is richer than that in apple leaf falls. Consequently, fruit forests return to the soil by way of leaf-falls, much more CaO, MgO, and P2O5 than treeless plots (Fig. 2.20). Therefore, in the soil under fruit forests, these elements are accumulated in greater amounts. The great mass of organic substances, which is annually deposited on the soil surface, provides nutrients for an abundant insect population in the apple forest litter and soil. Feeding on this dying vegetative mass, kg/ha 160 140 120 100 80 60 40 20 0

1

2

3

4

5

6

7

8

Plot 4. Applewood (density 0.7)

Plot 3. Applewood (density 0.3)

Plot 10. Glade

Strongly leached deep chernozem

Fig. 2.20. The amount of ash elements and nitrogen, returning with leaf fall under apple woods and on treeless plots: 1 = SiO2; 2 = CaO; 3 = MgO; 4 = Fe2H3; 5 = Al2O3; 6 = N; 7 = K2O; 8 = P2O5.

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soil organisms grind it and promote the first decomposition stage of the vegetative residues. The earthworm migration under apple woods involves an intensive mixing of the soil mass. It also results in a transfer of a considerable amount of vegetative residues and humus from the upper layers to the lower layers. This process provides a loosening of the whole soil profile and the formation of a grained-cloddy structure, which leads to a leveling of the physical and chemical properties of the separate layers of the soil profile and to an increase in the horizon thickness. A widespread distribution of microorganisms also promotes a rapid decomposition of leaf fall and litter formation, which enriches the soil with organic substances and ash elements. This soil-forming process under forests depends on the quantity and quality of the forest litter, which should be considered as a unique source of forest soil fertility (Remesov and Pogrebnyak 1965). On strongly leached deep chernozems, litter resources are greater than they are on weakly leached ones; in dense apple stands the litter is greater than it is in thin forests (Fig. 2.21). A comparison of leaf fall ash compounds with those of the litter shows a reduction of 1.4 times for CaO, but an increase of two times for SiO2 and four times for Fe2O3 and Al2O3. CaO, K2O, and MgO from the litter neutralize the acid products, which form as a result of microorganism activity. This promotes a mild accumulation of humus and the fixation of Fe2O3 and Al2O3 in the biological cycle in upper soil horizons. P. S. Pogrebnyak (1970) recommends that, while studying the role of vegetation in soil-formation, one should take into consideration the ratio index of the element amount in leaf fall compared to the element quantity fixed in the phytomass over an extended period. In the apple forests of Zailijskei and Dzhungarskei Alatau, the ratio of litter resources kg/ha

1

2

3

4

5

6000 5000 4000 3000 2000 1000 0 Plot 4. Applewood (density 0.7)

Plot 3. Applewood (density 0.3)

Plot 10. Virgin land

Strongly leached deep chernozem

Fig. 2.21. The influence of apple wood on leaf falls and litter: 1 = litter; 2 = the total mass of fall; 3 = apple fall; 4 = grass cover; 5 = aspen fall.

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compared to leaf fall is less than 1, which reveals the intensive process of leaf fall decomposition under these conditions. All of the annually produced organic substances are fully decomposed by the end of the next vegetative period, which is the basis of the energy cycle of ash elements under forest canopies. This is one of the main reasons for such high fertility in the mountain chernozems of the apple phytocoenosis belt. The favorable characteristics of these mountain chernozems for apple woods were formed under the influence of a whole complex of biological processes unique to these coenosis. Under their canopy, not only are the soils changed, but also the climatic conditions as well. The influence of climate on forest soils is changed by the interaction with the forest coenosis complex growing in them. The forest coenosis of apple forests significantly change the hydrothermal regime of the air and soil and form their microclimate. Consequently, soil and climatic conditions, to a great extent, are regulated by the forest coenosis and they depend on the biological peculiarities of woody plants, the density and age of stands, understory, grass cover and the forest litter components. Thus, in the soils of these investigated regions, while developing on a single parent rock (loess) under apple stands, the podzolization process was absent, but under aspen stands of the same belt, this process proceeded especially distinctly (Dzhangaliev and Zener 1969). The physiological uniqueness of forest-forming species, and their coenosis and in particular their selective consumption of mineral substances from the soil are decisive factors, which are closely related to other quantity and quality features of leaf fall compounds. This is the litter of forest vegetation. The results of these investigations revealed the interdependence of the processes, which occur under an apple forest canopy, and thus establish a regularity of microclimate improvement in the territory and thereby an increase in mountain chernozem fertility. V. CHARACTERISTICS OF WILD APPLE GROWTH AND DEVELOPMENT A. Natural Renewal For ecological adaptability to mountain conditions, M. sieversii is dependent on a natural renewal ability to grow in multiple forms. There are no generalized data described in the literature on the natural renewal of apple woods, though this process predetermines the present and the future of apple forests.

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1. Vegetative Renewal. Vegetative renewal of apple in natural forests is widespread. After visiting and inspecting our station and apple root excavations (Zailijskei Alatau, Gorge Kamenskoe Plateau), P. G. Shitt (1952) stated that The most demonstrative and convincing examples of the ability of woody fruit-plants to propagate vegetatively by root shoots are the data from the biological investigations of wild apple overgrowth and . . . numerous root-system excavations conducted under the guidance of A. D. Dzhangaliev. . . . (p. 64–77)

M. sieversii possesses the ability for vegetative renewal and for spreading, mainly due to the formation of root suckers, to some extent stem shoots, and also by adventitious root formation and self-forming growth. Root-sucker Propagation. Different opinions have been given regarding the root-sucker propagation of woody plants. Most investigators are inclined to think that root-sucker growth may be expected only in cases where the plant trunk, shoots, or roots are damaged. Consequently, general weakness, sick growth, or crown deformation may be observed (Arnold 1891; Turskij 1912; Pravdin 1938; Tkachenko 1952). A. Kerner (1908) explained that “crown saps intended for upper wood stems” could not find another way out and so form “slender shoots, grown in great numbers under the soil.” Regarding this, S. S. Pyatnitskij et al. (1963, p. 282) wrote: For species that are not inclined to form root-suckers under normal conditions, but which can form them on damaged trunks and roots, a rootsuckering ability is a response for the organism’s regeneration.

Yet, many trees and shrubs produce root-suckers when their upper and subsurface parts are not damaged. E. E. Kern (1934) and I. F. Gritsenko (1949) listed trees and shrubs that propagate by suckers when roots are not injured. S. S. Pyatnitskij (1963) considers the ability of trees to produce rootsuckers in the process of their normal development as a useful adaptation, which provides them with resistance in the struggle with other plants and makes it possible for them to steadily retain and occupy new grounds. Our study site investigations showed that M. sieversii (Shitt 1952; Dzhangaliev 1973a) propagated by root-suckers without preliminary damage to its upper and surface parts (Fig. 2.22a). P. G. Krasilnikov (1949) who studied the vegetative renewal of M. kirghisorum in southern Kirghizia

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Fig. 2.22. Root-sucker propagation of Malus sieversii in the Kazakhstan mountains. (a) general view; (b) detail. Surface skeleton roots of wild apple that have young suckers.

also noted this phenomenon. The basis of root-sucker formation, which provides for a natural renewal of M. sieversii in autovegetative propagation, is the horizontal development of surface skeleton roots (Fig. 2.22b) on which new foliated suckers are formed. New roots on the lower parts of the mother root or at the base of the foliated suckers then follow. Rootsuckers are formed from adventitious buds on horizontal roots. E. A. Baranova (1951) ascertained that root rudiments of trees appeared from

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cambium cells or from the cells that produce cambium as ray and phloem wood. For normal growth and development of suckers on roots, suitable environmental conditions of aeration and humidity are needed. Such conditions (Dzhangaliev 1972) are in the upper soil horizons, where roots of the mother plant are located near the surface. Under natural conditions, the success of root-sucker renewal for M. sieversii also depends on the shoot arrangement, the time of appearing and the vitality of the mother root system. As a result of normal stand thinning under natural conditions, several large trees may grow from suckers. They develop at the base of horizontal root branches of mother plants and thereby they are better provided with growth substances. As excavations showed, if a mature tree has several main root branches, the same number of stems differentially develops on the mother root. These trees are equal in their growth and are arranged within a circle of the initial tree. The mother root system separates as if divided into parts, and mutually exchanges assimilators with the nearest sucker stem. Among the numerous root-suckers, the first one along the ascending current is the first to develop and the rest are delayed in their growth and are sometimes suppressed by the first sucker. A similar statement was made by V. P. Sushchenko (quote from Pyatnitskij et al. 1963) for some forest trees. Numerous root-suckers, formed on small secondary roots, fall off in the first years because of light and growth substance shortages. Spreading out while growing, young tree crowns create additional shading, interrupting the growth of the second story specimens, which finally leads to their dying off. Under shading conditions, the first flush of young suckers dies off, then the low branches and later the stems, but root necks and roots stay alive. Then, this dying off of young seed trees passes through another succession: branches, flush and stem-roots. Weak suckers cannot take an active physiological role in any interrelational activities in the root system or in the aboveground parts of a plant newly forming and therefore they do not play any active role in the renewal processes of biogroups. Under natural conditions when mother trees fall in root-sucker apple woods, only the aboveground parts die off, while the roots on surviving trees continue to function. Growth and development of suckers mainly depend on forest-vegetative conditions, soil fertility, light and especially on human influences: felling, grafting, hay-mowing, cattle grazing, and others (Table 2.18). Human influences, such as apple tree cultivation, cause a weakness in root-suckers. Suckers are created that make poor use of the mother root system and do not receive assimilation products. Human influences are associated with a falling off of young suckers and a gradual dying off of the root system.

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Table 2.18. The influence of different methods of cultivation and development on root-sucker renewal in apple forests. Quantity of root suckers, 6 to 10 years Character of tree-stand use

No. trial plots

Slope exposition

(No/ha)

(%)

7 5 11 9

NE NE NE NE

531 102 242 347

100.0 19.2 45.5 65.4

Natural apple stands Haying + pasture Re-grafting Felling-management (thinning, sanitary)

In normal natural conditions, the suckers are fed both by the mother plant (in the first two to four years) and by their own roots, which begin to form in three to four years, producing vigorous growth in the first years. Only their own roots feed eight- to ten-year-old suckers, though the connection with the mother plant does not cease for a long time. Numerous excavations of root systems including adult trees (20 to 25 years old) showed that a sucker root connection with the mother plant continues for a long time (Fig. 2.23). With progress in aboveground development and in the formation of new roots, suckers move toward a relative independence in autotrophic nutrition and in this respect they simulate an apple of seed origin. The intensity of root-sucker formation and stool-shoots of apples after removing mother trees was observed in our 1964 to 1972 trial plots (Table 2.19). The felling of mother plants is accompanied by vigorous growth of shoots on trunks and by new root-suckers in the first year. However, the numbers in next years fell off considerably for stool-shoots and even Table 2.19. thinning.

The dynamics of vegetative renewal status of M. sieversii after woodstand

No. shoots or suckers/ha 1964 Plot number 16 10 13 1

1971

1972

Slope exposure

No. apple trunks

1

2

1

2

1

2

NW NE SE SW

162 200 149 147

4212 7200 10682 3969

2690 2672 3372 2156

843 634 542 448

91 122 107 78

326 244 301 154

88 116 103 72

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Fig. 2.23.

Young daughter tree emerging from the parent tree’s own roots.

161

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more so for root-suckers. The number of root-suckers in nine years after felling decreased on an average by 97 percent, which is a result of severe competition between a tremendous number of stool-shoots and rootsuckers for light, moisture, and nutrient substances. Simultaneously, a negative influence in this struggle for water and nutrient elements is caused by an increasing competition from grass vegetation, the thickness and cover level of which increases after wood stand thinning. Propagation by adventitious root formation with aboveground shoots is one of the means of vegetative renewal of M. sieversii. Adventitious roots from layers are formed in the case where horizontal low apple branches adjoin the soil and are covered by litter (Fig. 2.24). But in natural populations of M. sieversii, there are some cases of forced adventitious root formation, not with branches, but at the base of trunks from root initials exposed during floods and soil erosion. In such cases, new root formation is an adaptive apple response to worsening growing conditions. Root initials growth is a very useful biological adaptation of M. sieversii to extreme growth conditions. Most authors (Komarov 1938; Pyatnitskij et al. 1963) refer to adventitious root formation of plants as a more primitive character than the underground root system. The first psylophites, not having real roots, had rhizoids on underground rootstocks. Plants of the coal period, grow-

Fig. 2.24.

Wild apple propagation by adventitious root formation.

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ing during periodical floods, existed by adventitious roots. This statement, once more proves that M. sieversii is an ancient species in the Kazakhstan mountains. Self-grafting. The accretion (self-grafting) of roots, branches, and stems of some tree and bush plants is a widespread phenomenon (Krenke 1926, 1950; Shishko 1948; Rubtsov 1950; Yunovidov 1951; Ogievsky 1954; Beskaravajnyj 1956, 1958; Kovalev and Saburov 1957; Nikitin 1959; Pyatnitskij et al. 1963). These investigators give a list of the deciduous and coniferous species that self-graft by vegetative organs. M. sieversii is not mentioned among them. Data on the accretion (self-grafting) of wild apple root systems under natural conditions are not cited in studies devoted to the investigations on their roots (Al. A. Fedorov and M. A. Fedorov 1949; Krasilnikov 1949; Shitt 1952). While studying the apple forests of southeastern Kazakhstan, we observed that the roots of close-growing trees, which come in contact with each other, can selfgraft (Fig. 2.25). The tree root systems of root-sucker origin that self-graft are united by the common mother root. Thus, trees that have accreted by roots form apple colonies growing over wide areas.

Fig. 2.25.

Root accretion (self-grafting) between wild apple trees growing close together.

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N. P. Krenke (1950) gives a theoretical explanation of the tree selfgrafting (accretion) technique. According to his concept, tree roots are accreted mainly by the “scope” method, in which due to growth pressure, the cambium accretes and becomes united with both components of the self-graft. External differences of accreted apple roots are similar to that which was given by N. P. Krenke for other trees. At some distance from the point of contact, both accreted components form a typical callus. In this accretion process as a result of natural grafting, trees form a common cambium and make general annual rings, that is, they physiologically develop into an entirely united organism. It is evident that apple proliferation by means of root system accretion is another convincing argument for the survival of this species. The self-grafting of aboveground parts in the forests is seen much more rarely than that of the normal root systems and the role of selfgrafting for apple renewal is less important. For the accretion of stems or branches of different trees, their close contact is needed and this is seen very seldom. With light-demanding apples and under conditions of low relative humidity in an open habitat, the aboveground parts cannot contact each other and therefore they accrete very seldom. For self-grafting of root systems, there are more favorable conditions in the forest. Our excavations showed that the main roots of adult apples spread in a horizontal direction for more than 10 m and then penetrated into the root zone of another tree. In the process of parallel growth, roots can come into direct contact. Thus, a biogroup develops into a common vegetative canopy and evolves into a united colony organism, which preserves and penetrates into new areas. The pressure of the upper soil horizon also promotes root self-grafting, under the weight of which the root arrangement is fixed and where, in addition to that, there are the best conditions of humidity and aeration. Apple trees that are accreted by roots exchange nutrient substances. Nutrient exchange is demonstrated by the continual vitality of attached stumps and the continual accumulation of annual rings even after the aboveground parts have been discarded. Some cases in relatively thick stands of apple (density 0.5–0.6) are found in which stumps continue to live and are gradually overgrown by wood. In these cases, the stumps are connected with the roots of other living trees which support their vitality by an assimilation apparatus (Fig. 2.26). Stump age greatly influences its ability to regrow. After felling, in a stump area of 20-year-old trees, 32.6 percent of the area usually grows again. The felling of trees older than 40 years practically leads to no regrowth of the maternal stumps and they rot. This negatively influences the longevity of a root-sucker colony. In the case of stump regrowth with

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Fig. 2.26. A wild apple stump, functioning at the expense of a connection with the roots of living trees.

new tissues, the mechanism, as described by P. I. Molotkov (1956) during his investigation on beech nutrient substance translocation, apparently operates. Moisture deficiency in the aboveground parts of a tree with damaged roots is compensated by an adjacent tree, which has a common root system. Because of a lower negative water pressure potential, a healthy maternal tree transfers part of its water resources and nutrient substances to the suffering tree. On the other hand, in the case of the defoliation of one of the trees in a biogroup, a negative pressure is formed in stems of adjacent healthy trees, which are connected by common roots, and a part of the water resources and nutrient substances translocates from the suffering tree to the healthy ones. Perhaps, attached trees withstand snowfall and erosion in the mountains better than unattached trees. S. S. Pyatnitskij et al. (1963) suggest that self-grafting of forest trees plays a positive role in their vitality, otherwise this innate characteristic would have been blocked by natural selection. 2. Renewal by Seed. Heavy bearing and the production of quite viable apple seeds testify to renewal by seeds. Seed output from 1 ton of fruits is considerably higher in all forms of M. sieversii (21.4–26.7 kg) than in cultivated apple (e.g., ‘Aport’, 2.2 kg; ‘Renet Burkhardt’, 6.3 kg) (Dzhangaliev and Tasymov 1971). The weight and size of fruit seeds are

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important indexes of their potential ability for survival. Our investigations point to heavy seed weights (seeds were taken from 200 fruits) in all ecotypes of M. sieversii (23.7–37.6 g) in comparison with M. prunifolia (19.4 g). Some of them (ecotypes III and IV) had larger seeds (35.7–37.6 g) than ‘Renet Burkhardt’ (32.0 g) and ‘Aport’ (35.2 g). An analysis of long-term investigations (Dzhangaliev and Tasymov 1971) showed that the best seed germination came from ecotypes of M. sieversii (33.5%), M. niedzwetzkyana (20.2%), M. domestica (23.5%), and M. prunifolia (16.2%). We investigated renewal of apple forests by seed in the mountains of Dzhungarskei Alatau in phytocoenosis of different ages, density and composition of wood species on slopes of different exposure, and steepness. In order to measure germination and seedling growth, 40 experimental plots were laid out, each of 100 m2 in size. Apple seedling growth was measured at the ages of 1 to 2, 3 to 5, and 6 to 10 years. The amount of M. sieversii self-seeding under a canopy of its own stands is insufficient (1 specimen on each 5–20 m2) and after the age of 5 years, it regularly decreases (Table 2.20). Under the unfavorable conditions of mountain regions, renewal by seed is insignificant. Thus, according to V. I. Infantiev’s data (1957), renewal makes up 10 to 15 percent of the total amount of seedling growth in Dzhungarskei Alatau. Limited renewal by seed of wild apples was pointed out by M. G. Popov (1929), M. G. Popov et al. (1935), A. A. Fedorov et al. (1945), Al. A. Fedorov and M. A. Fedorov (1949), A. D. Dzhangaliev (quote in Shitt 1952), and I. T. Vasilichenko (1963). The predominance of stands of vegetative origin in apple forests is apparently the reason for the statement that M. sieversii lost its ability for renewal and propagation by seed. However, all this may be the result of a disruption of interrelations in modern apple biocoenosis, both with its biotic components and with its abiotic environment. In V. T. Langenfeld’s opinion (1970), the apple in the process of evolution may have adapted to spread its seeds by animals, which eat its fruits. The influence of gastric juices considerably promotes the breakdown of long seed dormancy and stimulates the rapid germination of seeds. Without the influence of acid, organic seed dormancy of apples is broken only after a long period (about 6 months) of low temperatures. During this period of time, insects, birds, and mice eat great quantities of apple seeds. Of no less importance, the other cause limiting seed germination of M. sieversii is low soil humidity. Insufficient precipitation and dry summer periods unfavorably influence seed germination and further limit the growth of wild apple seedlings. V. I. Zapryagaeva (1949, p. 14) considers, that under arid conditions, all moisture-loving species (maple, walnut, apple, and others) are prac-

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Table 2.20. The amount of M. sieversii self-seeding under stand canopies on slopes of different exposure and steepness. Location Species characteristics of stands

No. self seedings/ha

Exposure

Slope steepness, grade

1–2 yearold

3–5 yearold

6–10 yearold

Total

8 apples (30–40 year-old), 2 hawthorns. Density 0.4. Bonitet II

NW

15–20

1300

800

–

2100

8 apples (20–30, 50 yearold), 2 aspens (10–15, 30 year-old), 1 hawthorn. Density 0.5. Bonitet II

SW

5–10

200

200

100

500

10 apples (20–30, 50 yearold). Density 0.5. Bonitet III

SW

10

400

200

–

600

7 apples (40–50, 70 yearold), 1 hawthorn, 2 aspens. Density 0.8. Bonitet II

SW

15–20

300

200

300

800

8 apples (30–40 year-old, 60 year-old), 1 hawthorn, 1 aspen. Density 0.7. Bonitet II

SE

30

300

300

200

800

4 apples (50–60 year-old), 2 hawthorn (50–60 year-old), 4 aspens. Density 0.5. Bonitet III

NE

20–30

200

200

200

600

4 aspens (40–50 year-old, 20 year-old), 1 hawthorn (40 year-old), 5 aspens (20–30 year-old). Density 0.4. Bonitet III

SE

15–20

–

600

300

900

tically deprived of a normal renewal by seeds. Therefore, it is natural and even necessary to use every vegetative means for their propagation or to essentially change ecological conditions of their growing. Thinning of apple woods and mass tree grafting cause microclimate changes to dryer conditions in which the soil temperature increases and water evaporation from the soil surface increases. Our investigations (Dzhangaliev 1972, 1973a) indicate that the soil humidity on the surface layer, where apple self-seeding takes root under an apple stand canopy, is always greater than the soil humidity in open glades. Stepping is accompanied

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by a change in the grass cover composition in which elements of xerophytic steppe vegetation begin to dominate. These conditions unfavorably influence self-seeding and apple regrowth. The data in Table 2.21 show that greater germination occurs with apple seeds from slopes with a southern exposure, because they are more humid than northern slopes. Also, the shading of the young apple seedlings by the mother canopy and by grass cover competition play a definite role in self-seeding success. The self-seeding, which is concentrated under the mother tree crowns dies off to a considerable extent without ever reaching the stage of regrowth. On slopes with northern exposures, the shading formed by the mother canopy is accentuated to some extent by the shading of the slope steepness and this plays a more important role in self-seeding development than it does on southern slopes, where enough light is provided. On slopes of southern exposure, almost all apple trees, which grew from seeds, reached an age of 6 to 10 years. Insufficient renewal by seeds of M. sieversii is to a great extent determined by the grass cover. On southern slopes in comparison with northern slopes, grass vegetation in the apple forest is more weakly developed and only 52 percent of the soil is covered by grass on the average. Satisfactory apple renewal by seed occurs in comparatively inaccessible places which have a slope of 20 to 30° or higher, where the apple woods are preserved in a natural condition (Table 2.21). Fruit harvesting to collect seed for apple renewal in favorable environments greatly decreases the need to plant apple plots, because it is practically impossible to harvest the whole yield and the unharvested fruit would provide self-seeding. 3. The Vitality of Forest Stands from Vegetative Origin. We wondered if the long, natural vegetative propagation process of M. sieversii leads to apple forest degeneration. We have not found any data in the literaTable 2.21.

Influence of steepness and direction of exposure on M. sieversii renewal. Slope location Wood stand origin

Wood stand characteristic Mixed apple wood. density 0.5 Pure apple wood. density 0.6 Pure apple wood. density 0.4 Mixed apple wood. density 0.4

Exposure

Steepness (°)

Seed

Vegetative

Trees from seed (%)

NW NE SE SW

19–22 20–25 20–25 25–30

42 63 72 73

308 357 228 197

12 15 24 27

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ture that reports on the influence of vegetative apple tree propagation on the stand’s vitality and progeny. The data on other forest trees are of a contradictory nature. Thus, E. Romeder and G. Shenbach (1962, p. 60) wrote: The old disputable question, repeated for many centuries that vegetative propagation leads to degeneration or at least to aging and decreasing tree resistances, is still not elucidated. Many botanists do not admit to clone aging, while others, especially some craftsmen and fruit-growers take it for granted. (1962, p. 60)

Furthermore, they advocate methods and recommend taking the youngest parts of a tree (stool shoots, etc.) for plant material production by vegetative propagation. D. I. Prutenskij, on the basis of the analysis of research by G. F. Morosov, A. S. Yablokov, V. T. Petrov, I. M. Naumenko, N. K. Vekhov, M. P. Iljin, I. V. Michurin and other investigators, draws this rather contradictory conclusion: As we see, two points of view on sprout renewal exist. The first is that the negative characters of sprout renewal are unimportant, and the second, on the contrary, recommends the substitution of seed plantings with sprout plantings. (1958, p. 42)

Then he states that there are no data on growth, productivity, longevity, or resistance of sprout or seed plantations of Persian walnut. He is against Persian walnut sprout propagation as a method, pointing to its natural renewal. S. S. Pyatnitskij et al. (1963) believe that vegetative sprouts of woody species and their regeneration do not experience premature aging, do not differ from seed plantings, and that this sprout renewal ability was gained during the evolutionary process. The differences between seed and vegetative origin of woody species “are mainly conditioned by the age and status of those vegetative organs on the mother plant which initiated their growth” (p. 382). M. M. Beskaravajnyj (1956), A. P. Slyadnev (1953), and G. R. Eitingen (1949) noted that the self-grafting of pine tree root systems does not lead to planting degradation. N. I. Vavilov (1931, p. 103) pointed out that the possibility of finding the reason for isolated species proliferation is in sprout propagation which is peculiar to wild fruits. Wide distribution of sprout and root-sucker propagation of some fruit trees should be taken into account and investigated in detail. For a study on the biological role of vegetative renewal in the process of natural formation and development of M. sieversii tree stands, we

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chose a plot in Gorge Soldatskoe of Zailijskei Alatau, where there was a pure apple stand with a density of 0.6. It is situated in the middle of a northern slope (steepness 19°) at an altitude of 1300 m. The soil is a strongly leached medium-deep chernozem, with a humus horizon having a thickness 70 cm and a carbonate layer 120 cm deep. The main mass of apple roots is arranged in a soil layer of 10 to 15 cm deep. The soil cover consists of Urtica dioica (nettle), Milium effusum (carpet grass), and Aegopodium alpestre (Goatweed). An experimental plot (100 × 100 m) was chosen in the apple stand. In preparations of the aboveground horizontal skeletal roots of trees, their origin (vegetative or seed), height, and diameters of their trunks and crowns were determined. There are 323 trees of M. sieversii on the plot, of which 23 (7.1%) are of seed origin and 300 (92.9%) of vegetative origin. The apple stand has a clearly expressed structure (Fig. 2.27), where closely growing stands of different aged specimens alternate with open

Fig. 2.27. Plot scheme of Malus sieversii biogroups (Zailijskei Alatau): 1 = stumps; 2 = apples of root-sucker origin (40–60 years); 3 = apples of root-sucker origin (15–20 years); 4 = apples of seed origin (40–60 years); 5 = apples of seed origin (15–20 years).

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(free) glades. As a rule, different aged young-generation trees are concentrated around the mother trees, but the suckers in many stands are at a considerable distance (7–8 m) from the mother plants. There is a young re-growth among the trees of the older generation. The common root system promotes the vigorous growth of root-suckers. In the case of one of the trees of the group dying, the remaining trees use its root system, which also increases their current growth rate. Somewhat apart from this group of trees grow trees of seed origin, which are inferior in size to the size of trees of vegetative origin. Taxonomic indexes of medium-sized trees of vegetative origin at an age of 15 to 20 years are larger than of those of trees of seed origin at the same age. A medium-sized vegetative tree exceeds a seed-grown tree in height by 17.7 percent, in trunk circumference by 24.1 percent, and in crown diameter by 7.6 percent. This difference in these indexes decreases with tree age and after 20 years they are 14.6 percent, 13.2 percent, and 2.7 percent respectively (Dzhangaliev 1973a). These data agree with the results of S. S. Pyatnitskij et al. (1963), who found that forest species of vegetative origin (layers, rootstocks, or root-suckers) grew more from seeded plants in height and diameter at 5 to 10 and even at 20 to 25 years of age. Thus, the growth processes of vegetative apple woods are not dwarfed and signs of tree aging do not appear. The contact between root systems of M. sieversii of a vegetative origin forms a complex organism, which provides the tree biogroup with great vitality and adaptation to mountain ecological conditions. B. The Influence of Tree Thinning and Apple Cultivation on Apple Renewal and Stand Preservation Judging from the appearances of modern apple stands, tree thinning was used long ago. N. A. Severtsev (1873) noted that wild fruit trees in the foothills of Zailijskei Alatau were relentlessly cut down by local populations for economic needs and agricultural crops were sown in their place. M. G. Popov et al. reported: Premountain belts were subjected to various influences. Man’s influences essentially changed the original vegetative cover. Such influences refer to: 1. Fruit forest thinning. 2. Grass and bush burning. 3. Cereal cultivation. (1935, p. 10)

M. G. Popov in connection with the influence of thinning apple writes: Many trees (50%) have such disfigured crowns that it is impossible to define their types. (p. 25)

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A. G. Klabukov states that in apple thickets of Zailijskei Alatau, where tree thinning and cattle grazing take place, forest renewal has ceased. In the Kamenka and Bogdanovskaya gorges, tree clearing and cattle grazing do not take place and therefore the number of young apple trees constitutes 50% in comparison to bearing apples. (quote of Popov et al. 1935, p. 86)

According to our observations in the territory of Talasskei Alatau (Aksu-Dzhabagly reserve), the number of young apple trees is three to four times greater than it is in neighboring apple woods where farmers practice cattle grazing and hay mowing. In the Talgar gorge of Zailijskei Alatau, older apple generations, which originated by seed renewal, coincide with the time of organization of the Alma-Ata reserve (1951). Thus, in apple forest territory, which is used intensively as pastureland, the soil cover changes. Young seedlings are damaged and this has an adverse effect on the rootsuckering ability of horizontal roots, which are located close to the soil surface. The grubbing of eglantine, barberry, honeysuckle, and almond thickets and other shrubs during improvement cutting and cultivating in apple woods deprived the apples of biological protection, which had been created by these prickly shrubs because they are inedible to cattle. This cutting drastically changes the growing conditions around the apples and reduces their natural renewal. From 1930 to 1936, in fruit forests, some sanitary and thinning improvement cuttings, directed at apple forest reconstruction were made. In mixed forests, fruit trees with weakly-developed crowns, low productivity and bad fruit quality and also nonfruiting wood stands were cut down. In circumstances of a considerable amount of undesirable species and an admixture of nonfruit trees and shrubs in an area receiving improvement cutting, it usually involved clear cutting. The roots of the remaining apple trees, which had a common root system with felled trees were broken and the upper fertile soil layer was taken away. Any plant damage stimulated its sprout-producing ability. With respect to tree development, the sprouting capacity is reduced for cut limbs and trunks. According to D. F. Protsenko’s data (1958), on cutting trees, the ratio between the root-system and aboveground parts is sharply disturbed because their dormant buds begin growing in large numbers. The same response occurs when trees are damaged naturally by frost, aridity, pests, and so on. Thus, bud-breaking is a response of an apple tree to unfavorable growing conditions and development by attempting to replace the destroyed leaf canopy. This ability of the apple characterizes it as a vital resistant species by possessing such an adaptable organ of regeneration such as the long-lived and productive dormant buds.

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Our investigations (Table 2.22) showed that trunks and skeletal apple branches, two to three years after felling, are entirely covered with numerous regenerative sprouts from dormant buds, especially if roots and aboveground parts of the mother tree had been damaged. In the following years, their growth gradually ceases and then they die. This applies mainly to the sprouts in their first year of life, apparently, having had no time for adaptation to the new conditions because they are strongly overshadowed and are formed in the lower parts of trunks. Though these sprouts do not live long, they negatively influence applestand growth and productivity and lead to weakness and dying of young apple trees of vegetative origin. Because of this weak development of generative organs, such trees rarely bear fruits. From Table 2.22, it is evident that sprout formation is directly dependent on the degree of crown damage. Through long-term experiments by fruit growers and foresters, it has been found that sprouts abundantly spring up and absorb mineral nutrition elements, which rise from the root systems to the crowns and this leads to desiccation of the tops. With respect to the biological effects, these sprouts rejuvenate the drying crowns, because crown dryness without dormant buds would lead to full death of the trees. S. S. Pyatnitskij et al. (1963) wrote, “Trees with watersprouts are distinguished by potential waterlogging, low cambium productivity and weak activity of apical meristems.” Improvement felling in apple woods leads to canopy thinning and improvement of light conditions for water shoots, on which leaves of fragile structure are formed. These are distinguished by stronger transpiration, leading to a disturbance of the whole stand status. Mass shoot appearance and the beginning of tree-top desiccation without human influence, and not provoked by sudden changes in the environment, simultaneously occur with a biological aging period, which, under natural habitat conditions, begins after 70 to 80 years of tree life. According to V. V. Smirnov’s data (1953), cut and exposed aspen roots produced suckers even at a depth of 1 m, but according to V. Z. Gulisashvili’s studies (1928), aspen root-suckers, forming on healthy Table 2.22. thinning.

Sprout stimulation of M. sieversii depending on intensity of apple stand

Thinning degree of woodstands Control (without thinning) Moderate thinning Vigorous thinning

Density

No. measured trees

No. sprouts/tree

Sprouts relative to control (%)

0.7 0.5 0.3

122 138 140

6 16 20

100 266 334

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roots, appear from a depth of 2 to 3 cm below the soil surface. The output of suckers by aspen with healthy root systems is considerably higher than the output from aspen with injured root systems. Because of the shallow position of apple roots, developing suckers are easily damaged during tree felling, cattle grazing, or hay mowing, which cause changes in natural forest vegetation. In apple forests, especially in those which are ripe and overripe, it is not expedient to make improvement felling in full volume, because this destroys their status and negatively influences natural renewal, especially root-sucker renewal. Here, it is enough to move away brushwood, dead trunks, and drying trees and to cut damaged branches. The following have written about apple forest cultivation: A. G. Klabukov (Popov et al. 1935), A. G. Araratyan (1940), G. I. Monashev (1947), Yu. E. Pukhtinskij (1948), N. F. Batygin (1950), E. I. Kharchenko (1952), Kh. Z. Gubajdullin and V. I. Dyachkova (1953), B. Z. Sabirov (1958), S. N. Krajnov (1961), A. G. Gusejnov et al. (1964), A. I. Pisarenko and M. N. Drozhalov (1966), and A. C. Abeev and N. T. Bondarev (1967). They consider grafting to be the main method of wild apple wood stand cultivation, that is, using them as rootstocks for grafting apple varieties directly in the forest. Grafting accelerates fruit bearing of forest orchard trees and enhances its longevity, yield, resistance to pests, diseases and unfavorable environmental conditions, and above all, it increases fruit quality, because cultivated fruits considerably exceed wild fruits in flavor and chemical composition. Grafting was mainly done between 1932 and 1967 in Kazakhstan, Kirghizia, Uzbekistan, Tajikistan, Azerbaijan, Daghestan, Armenia, the Krasnodar Territory, the Voronezh Region, the Crimea, and the Ukraine. The most extensive mass apple-stand grafting was done in Zailijskei Alatau (A. G. Klabukov, quote of Popov et al. 1935). Between 1932 and 1935, 120 thousand apple trees were cultivated. M. I. Gajdin (1962) cited data showing that in Sovkhoz “Gornyi Gigant” (near Alma-Ata) in 1957 to 1960, six thousand trees were grafted, and by 1965, it was planned to graft 100 thousand of such trees. Apple cultivation has continued. The authors of publications on forest orchards do not cite data on grafted apple stand areas, or on their preservation and economic efficiency, though millions of trees were cultivated. Until now, studies on forest orchards have not been summarized and no analysis of the practical use of forest orchards has been made. In order to elucidate the status of forest orchards, we inspected grafted apple trees in Zailijskei Alatau during 1970 to 1972. Scions of apple varieties were grafted into trunks and crowns in the greater parts of apple forests during 1931 to 1937 and 1959 to 1967. The height of the grafts was usually 80 to 110

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175

Effect of sapling origin on the development of roots.

Sapling origin

Measured

Rooted

Rooting of saplings (%)

30–50

Seed Root-sucker

53 127

24 34

46 27

50–70

Seed Root-sucker

38 142

14 26

36 18

70–90

Seed Root-sucker

64 123

18 15

28 12

No. saplings

cm, however sometimes grafts were as high as 2 to 3 m or as low as 40 cm. The main grafting methods are: whip-grafting, bark-grafting, cleftgrafting and semi-cleft grafting, that is, the usual means of grafting, which are used in horticultural practice. During these investigations on tree status, a record was made of successful grafts in relation to rootstock origin (seed or root-sucker), cultivar, and rootstock age, and means and time of grafting. From Table 2.23, it is evident that rooting by saplings from trees of seed origin, was much higher than from root-sucker trees. Rooting was apparently induced by a greater regenerative capacity of seedling trees. This is expressed by increased root-suckers and watersprouts from selfpropagating trees connected by a common root-system. The dying off of topworked cultivated variety components, which were grafted on young root-suckers of 4-to-5-year-old trees may be explained as follows. These young trees, up to that time had not yet formed their own roots and the stump on which the grafting was done did not provide the scions with nutrient substances. We observed mass dying of grafts on trees of root-sucker origin. Grafting of adult apple trees gave negative results because of overgrowths at the graft union and a sharp disturbance of the system due to an unbalanced ratio of the aboveground parts versus the root system. Overgrowth was greater if their diameter did not exceed 30 mm and only partially, with a diameter of 30 to 50 mm. They did not overgrow with a diameter of 50 to 70 mm or more, which led to the dying of a great number (48%) of grafts (Table 2.24). The best self-grafting was on rootstocks of seed origin. Large grafted trees appeared suppressed and blight-injured, which may be explained by a sharp change of growing conditions after grafting. Table 2.25 shows an unsatisfactory status 10 to 12 years after grafting. Rootstock types, grafting methods, and periods all had little

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176 Table 2.24.

A. DZHANGALIEV Breakage of grafts on wild apple tree cuts of different diameters. No. grafts

Cut diameter (mm)

Rootstock

Measured

Broken

Broken (%)

30–50

Seed Root-sucker

47 111

8 32

18 29

50–70

Seed Root-sucker

28 105

7 32

26 31

70–90

Seed Root-sucker

58 129

23 62

39 48

influence on the development of grafted trees, but rootstock origin had a much greater influence. The whole grafting status on rootstocks of seed origin was evaluated at 3.0 to 3.9 on the Kapper-Formozov 6 point scale and with root-sucker origin at 2.3 to 2.8. The increase in height on seedling rootstocks was 2.0 to 2.5 m and on rootstocks of root-sucker origin, 1.8 to 2.1 m, and the crown size was 1.8 to 4.0 and 1.5 to 2.8 cm, respectively. The total take of all grafts measured by us was 20 to 30 percent, but only 10 to 15 percent of them remained as fully formed trees. These trees growing in apple woods are considerably separated from each other, and this excludes the possibility of mechanical management and crop harvesting of these trees, which are in almost inaccessible mountain relief conditions. Those supporting the transformation of wild apple woods into cultivated forest orchards, assumed that the production of this type of Table 2.25.

The status of apple grafting components, depending on rootstock origin.

Height (m) Scion cultivar

Rootstock origin

Graft Tree union

Trunk diameter (cm) Rootstock

Crown size (cm)

Graft Scion union NS

EW

Whole tree status, point scale

Aport

Seed Root-sucker

3.6 2.6

1.1 0.7

7.2 5.2

5.2 4.3

8.7 5.3

2.7 1.7

2.7 1.5

3.5 2.3

Renet Burhardt

Seed Root-sucker

2.7 2.8

0.7 0.7

5.7 5.7

4.5 3.8

5.8 7.3

1.9 1.6

1.8 1.5

3.9 2.7

Zailijskoe

Seed Root-sucker

3.6 3.0

1.0 0.9

8.5 7.3

7.4 4.8

10.4 9.3

2.9 1.8

2.8 1.9

3.0 2.3

Pestrushka

Seed Root-sucker

3.5 2.7

1.3 0.9

8.1 7.1

7.2 7.9

9.8 10.0

3.6 2.8

4.0 2.5

3.5 2.8

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orchard agrosystem would be of a higher quantity and quality and more useful for people, than would the natural ecosystem of fruit forests. They supposed that cultivated varieties, when grafted straight into forests onto resistant rootstocks of wild apple adult trees (in contrast to methods of usual orchard growing—nursery to fruit-bearing orchard— which requires eight to ten years) would begin bearing in two or three years and produce high fruit quality. They envisioned the possibility of forest orchards, with high economic efficiency without any great energy or expense. Today’s point of view is that cultivated orchard production, especially with an intensive palmette orchard on dwarfing rootstocks, has considerably higher production than a wild apple wood ecosystem. The disadvantage of the orchard agrosystem is that production requires considerable labor and operations expense. The transition from complex ecological systems to monoculture of cultivated forest orchards brought short-term economic advantage as judged by such traditional factors as apple size, appearance, and flavor. However, the natural forest orchard systems lost the natural dynamic features of apple populations, which in their vital activity are tightly associated with other plant communities. The sustainability of ecosystems (Fedorenko and Rejmers 1974) is based on the common cybernetic principle of numerous duplications. The elements of this duplication in ecosystems are many systemic blockspecies, ecological populations, synusias, and so on. These numerous interspecific duplications, characteristic of natural ecosystems, are expressed by natural hybridizations, especially introgressive between M. sieversii and M. kirghisorum species and their numerous hybrids. These interspecific duplications, because they are members of the crosspollinated genus, Malus, were replaced in monoculture by intravarietal crossing within an impoverished genetic base, which negatively influenced apple species populations. The practice of cultivated variety grafting onto wild apples directly within the forest for developing forest orchards is not well-founded on biological principles. By this utilitarian method, it is not only possible to permanently destroy existing wood stands of apple, but to deprive them of their ability for natural renewal, and once and forever to lose the wealth of present apple forests of Kazakhstan with their unique gene pool. C. Growth and Development of Wild Apple Trees in Relation to Natural Renewal During the development of M. sieversii tree populations, regular changes in the growth characteristics of individual specimens occur. These changes allow us to divide the vital cycle of populations into various age

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periods, beginning with seed germination or autovegetative suckers, through regrowth formation into fruit bearing trees, and then to the dying of aged trees. While defining these regular occurrences, we took into consideration the theoretical preconditions of age changes by fruit plants discovered by P. G. Shitt (1952). The process of simultaneous and interconnected renewal and dying off is peculiar to all fruit trees. But its content and concrete forms of expression, which were elaborated by P. G. Shitt for grafted components of cultivated fruit trees, differ from the processes of natural renewal, growth and development, which are unique to natural M. sieversii populations or to wood stands formed by it. I period is the growth of vegetative tree parts from seed germination or the appearance of autovegetative specimens from suckers until the beginning of fruit bearing (to age 6 to 8 years). In this period, the active growth of stems and skeletal primary crown branches with overgrowing branches take place. Seedling and sucker numbers reach 3500 to 4000 pieces/ha, but a considerable portion of them dies off in the first years. In two to three years, 1000 to 1500 pieces/ha are left and they are arranged into group order. II period is growth and fruit bearing—from the beginning of the first to regular fruit bearing (to age 10 to 12 years). During this period, fruit bearing increases and the number of skeletal secondary branches and regrowing spurs also increase. The growth rate of vegetative tree parts becomes more moderate and the character of their growth is qualitatively different in that numerous, but smaller branch arrays and fruit spurs are formed. During this period, species differentiation of young regrowth, population formation and an increase in the dying of the rest of the trees are characteristic. Five hundred to six hundred of them per hectare continue to thrive. III period is fruit bearing and growth—from the beginning of regular cropping to the maximum of fruit bearing (age 10 to 12 until 25 to 30 years). In connection with active fruit organ formation, productivity increases, the growth of new skeletal branches is still more delayed, especially in the crown periphery, and small regrowing spurs die off. In connection with a progressive growth cessation of vegetative parts, the developmental processes and the dying of species populations are delayed; about 350 to 400 trees per hectare continue to live. Among the autovegetative young trees (suckers) the first ones near root nutrition are of primary development, which in their turn, at the end of the period, initiate new suckers. Young self-sown seed plants appear. IV period is full bearing—beginning at 25 to 30 years of age and continuing approximately until 60 to 70 years of age. The growth of skeleton parts fully ceases, and many short fruit spurs arise in the middle and

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upper parts of the crown with their simultaneous dying off in the interior shaded areas of the crown. The main skeletal branches gradually become bare from the base out to the periphery, and semi-skeletal branches in the lower crown visibly dry up. The surviving young regrowth gets a chance for vigorous development, and new trees arise and are formed. After 60 to 70 years, a period of rapid desiccation of overgrowing twigs, desiccation begins in the semi-skeletal branches and fully skeletal branch. In connection with the ceasing of their growth, all developing apical points form only short internodes. This results in vigorous new fruiting branch formation and a high yield with low market value. Later on, a progressive dying of overgrowing branches begins, then the dying of semi-skeletal and skeletal branches continues, initially in the shaded areas of the crown, and later water shoots appear. At that time, the spreading crowns of surviving trees of the second generation occupy developing gaps. A gradual dying of old trees begins, and this leads to the formation of gaps in the canopy. The surviving tree crowns because of their delayed growth processes cannot fill these gaps. These gaps, as a result of growth cessation and natural dying of the older generation of the population, gradually increase and new self-sown plant groups and natural suckers appear in them. The duration for species development depends on the genetic composition of the species and the environmental conditions affecting growth. For M. sieversii populations, the division of individual tree development into age periods is of great importance for understanding the natural renewal processes that take place during woodstand selfthinning. The sharp increase of population species differentiation in the first and second age periods is explained by a growth process involving the activation of skeletal and overgrowing tree parts, then subsequent crown closing. This crown closure acts as a regulating mechanism for light conditions in the population. During this period, trees begin to die because of a light shortage inside the canopy of the population. This delays individual growth and development. As a result of light shortage, growth under the canopy of the dominating trees is suppressed. Dominant trees at the periphery of the stand, being under optimal light conditions, increase in volume because the first and secondary skeletal branches spread. Vertical growth is limited and crowns develop oval forms. Counter to this: trees dominating in the center of the population grow more vertically; trunks are more clearly expressed; the highest skeletal branches are laid at sharper angles; and their crowns are narrow. Well-developed forests by M. sieversii are of the thin park type. Apple stands of these forests are divided into more or less isolated and rather

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thick groups of trees, regarded as populations. Populations may be considered as communities of apple trees that are relatively independent, strictly corresponding with each other in their development cycles, and occupying a definite territory. Depending on environmental conditions, the quantity, age composition and ecological–morphological characteristics of apple populations may differ. None the less, natural renewal is obligatory and common for them all, and natural hybridization also occurs. The population composition includes both seed and vegetative trees. In addition to taking part in cross-pollination and territory seeding, these trees promote an optimal density for apple stands. Together with other species of the community, they prevent the establishment of competing species. The root systems of seed trees frequently interlace with the roots of vegetative trees but, as a rule, their contact is not of a physiological but of a mechanical character. Within the territory occupied by a population, young seedlings and suckers grow. Dead apple trees and stumps with dead roots are also present and they sometimes form the whole underground accumulation. Thus, the process of self-thinning of trees in a population proceeds from seedlings or suckers to adult trees. A regulating mechanism for thinning, which differs depending on stand density, is influenced by different development periods. The primary species is provided with nutrition and this prevents the invasion of the territory by other species. The inhibition of the self-thinning process in mature apple populations creates the necessary preconditions for fruit bearing of surviving trees, for germination and growth of seeds, and for vegetative propagation. Thus, vegetative renewal in natural populations does not lead M. sieversii to reduced vigor. This species inherited an ability to adapt and has been provided with a very reliable potential to repropagate. As a result of natural selection, the following forms of vegetative apple renewal are preserved: mainly root sucker renewal, renewal by stool shoots, adventitious root-formation and self-grafting. These renewal forms promote the vegetative auto-propagation of new apple trees. In natural apple populations, the root system, besides its main function of nutrition, carries out an additional role as a propagation organ. A lack of knowledge of these natural renewal processes, especially concerning the many forms of vegetative apple propagation, led man to disrupt the established interdependence between the apple tree and its environment. This disturbance resulted in irrational economic management practices in apple stands: apple stand reconstruction aimed at forest orchard formation, regrafting, improvement felling, cattle grazing, hay mowing, and others. The application of these utilitarian and exhausting practices in natural apple populations resulted in reducing

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total tree numbers and in population disruption. It decreased the ability for natural seed and vegetative renewal and destroyed several age groups of vegetative propagated young trees. Thinning also harmed roots of mature trees. A population, as a self-regulating tree community, consists of seedlings and specimens of vegetative origin of different age groups. When trees within a population or tree groups at the extremes of adjacent populations are closely associated, root self-grafting takes place. This leads to the formation of large colonies. In regard to active growth of trees within a population and their intensive differentiation according to their age and developmental period, the process of tree self-thinning proceeds and is regulated by the density of their stand. D. Structural Features of Apple Root Systems in Relation to Growth Conditions Energy exchange as controlled by root nutrition and photosynthesis responds to an interconnection between roots and the aboveground systems and this contributes to the status of the plant’s vitality and productivity. Understanding the biological features of fruit tree aboveground parts and their productivity is impossible without first studying their root systems. For a long time, there was an opinion in science that the root is a simple organ, serving to attach a plant to its substratum and receiving water and nutrient substances from it. Subsequently, data obtained by domestic and foreign investigators showed that roots are able to absorb and assimilate not only minerals but also organic substances, which sometimes have complex structures. Vitamins produced by soil microorganisms and organic nitrogen in the form of amino acids are also absorbed (Thomas 1926; Litvinov 1927; Ratner and Dobrokhotova 1956; Ratner et al. 1956). The conversion of mineral nitrogen into organic nitrogen occurs in roots as does alkaloid synthesis (Shmuk 1940; Rubin and Artsikhovskaya 1948; Kursanov 1954; Turchin et al. 1955; Petersburgskij 1957). V. V. Tserling (1970) pointed out that roots of rosaceous fruit trees have a strongly developed renewal capacity. At the same time, roots that are growing in a soil substrate considerably influence the soil quality. They not only absorb the elements needed for plant life, but also draw out a great number of mineral and organic substances, including ferments. They actively influence the soil substrate and especially the biorhizosphere (Rovia and Harris 1961). Malus sieversii actively transports substances from roots, litter-fall and outwash from crowns, which are low in toxicity. Thus, this species is distinguished by its low allelopathy and its low soil depletion (Popivshij 1974). Root extractions by apple are more abundant than extractions by other

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species. However, a microorganism influence on roots indirectly exerts an effect on species interrelations, which in turn determines their competitive ability. Malus sieversii’s forest-forming function is to generate multiple species communities. Close contacts and root-interlacing among different species within the ecosystem are indirect indications of apple’s weak allelopathy and its compatibility with other species within the plant communities of the mountains of Kazakhstan. At this time, considerable factual data on aboveground plant parts have accumulated. There are many fewer data on roots of perennial wood plants, because apparently, great difficulties exist when studying them. Excavations conducted by a number of investigators revealed that root activity and habitat are determined by a plant’s hereditary nature. The soil complex also has some influence on root formation. Connon (1925) noted a dependence of root arrangement on the thickness and physical properties of soil horizons. T. K. Kvaratskhelia (1947) found that the layers of growth and the distribution of roots in soil are determined by differences in physical properties of some created horizons, by air-soil and water-soil regimes and by species litters. These distribution characteristics of woody-plant roots, which depend on soil conditions, was studied by P. G. Shitt (1936); P. G. Shitt et al. (1937), V. S. Shumakov (1946), M. P. Levina and A. Z. Tsivindo (1969) and E. M. Kovalenko (1972). They ascertained a close and direct connection between root formation and growth with the availability of nutrient substances in soil. Rogers (1939), P. G. Shitt and Z. A. Metlitsky (1940), V. A. Kolesnikov (1954), and A. K. Prijmak (1959) and other investigators showed that if nutrient substances in the soil are unevenly distributed, the best growth and branching of roots occur in horizons that contain the greatest amount of nutritional elements. The work by V. S. Zakotin and A. P. Atanasov (1972) elucidated an interconnection between the separate organ-forming processes in the aboveground parts or root systems and the annual cycle of apple growth and development. Wild apple root systems were studied by A. A. Fedorov et al. (1945), P. K. Krasilnikov (1949), Al. A. Fedorov, and M. A. Fedorov (1949), A. P. Dragavtsev (1956), V. I. Zapryagaeva (1964), and A. D. Dzhangaliev (1973b), who described the surface arrangement of the main root mass and noted their exceptional capacity to form suckers. Less studied than many other ecological and biological problems of Malus sieversii are the growing conditions of root systems. The subjects of our investigations were 30-to-60-year-old trees of M. sieversii growing in selected habitat types. Root system organization was studied by skeleton and cutting methods, and nutrient absorbing of root parts by profile methods (Krasilnikov 1950, 1957; Kolesnikov 1972). For typical

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trees, soil profiles were made at a distance of 2, 4, and 6 m from the trunk, but sometimes closer. In the second cutting from the trunk, the morphology cut method was used to describe the soil profile and soil samples were taken for analyses. The lateral cutting walls served for studying the root system by the cutting method. The soil horizon borders and the arrangement and diameter of root cuttings were drawn on hatched paper. Soil profiles were taken to a depth of 150 cm in 5 replications by means of a steel cube (10 × 10 × 10 cm). During processing, roots were washed, dried, and sorted into three fractions: diameters 1 mm and smaller (the first fraction); 1 to 3 mm (the second fraction); 3 mm and thicker (the third fraction). Table 2.26 shows that the more productive apple stands (density 0.6 to 0.7) are formed under humid growth conditions, for which deep and medium-deep mountain chernozems of different leachings are characteristic. The chernozems of Zailijskei Alatau are distinguished by their rich stores of nutrient substances (Table 2.27). Under humid growing conditions, apple trees normally develop to heights of 7 to 8 m and their yield is 51 to 63 kg per tree. Periodical over-saturating of meadowchernozem soils (especially in the spring) unfavorably influences apple productivity, so that bearing, in comparison with humid growing conditions, decreases by 20 to 25 percent. Weakly-leached chernozems under dry growing conditions are characterized by fewer stores of nutrient substances. Here, trees reach a height of 5 to 6 m and they produce

Table 2.26. (1972).

Characteristics of M. sieversii stands according to growing condition types

Mountain system and growing conditions

Diameter Predominant age (years)

Tree height (m)

Trunk (cm)

Crown (m)

Mean yield/tree (kg)

Very dry Dry Humid on slopes Humid in valleys Wet

20–40 30–60 50–80 50–70 50–60

Zailijskei Alatau 2.7 5.0 8.8 7.4 7.5

17 26 41 38 40

3.2 5.0 8.4 7.0 6.9

4 19 63 51 45

Humid in valleys

40–60

Talasskei Alatau 5.0

23

4.3

36

Very dry

20–40

Karatau 3.9

16

4.0

6

184

Upper plot, southern slope, steepness 35°

Watershed plot, western slope, steepness 20°

Northeastern slope, steepness 16°

Foot of northern slope

Northeastern slope, steepness 12°

Eastern slope, steepness 12°

Dry

Humid on slopes

Wet

Humid in valleys

Very dry

67

32

Karatau Thin mountain-steppe soils

70

95

57

35

Talasskei Alatau Mountain brown leached soils

Deep meadow-chernozem soils

Deep strongly leached chernozem

Medium deep, weakly leached chernozem

Zailijskei Alatau Thin mountainsteppe soils

Soil

200

240

300

496

351

132

Humus

14.8

18.2

16.0

22.4

16.1

10.8

N

15.2

15.6

16.6

16.0

14.0

8.7

P

Nutrient substance reserves in a meter of soil layer (t/ha)

230.0

270.0

255.0

284.0

212.0

109.4

K

4:19 PM

Very dry

Relief

Humus horizon thickness (cm)

Nutrient substance reserves in soils according to growth condition types of woodstands.

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Mountain system and growth conditions

Table 2.27.

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small yields of 17 to 20 kg. Mountain-steppe soils with very dry growing conditions possess the least stores of nutrient substances. Under these conditions, apple tree growth is usually severely suppressed, tree longevity is considerably shortened and cropping is very light (4 kg per tree). In regard to nutrient reserves, mountain-brown soils of Talasskei Alatau are also good for apple stand growth and development, but fewer nutrient substances are contained in the mountain-steppe soils of Karatau (Tables 2.26 and 2.27). The first clearly perceptible outward index, which reflects plant adaptation to a soil substrate condition is the thickness and character of the root system arrangement of fruit trees. Root system excavations of M. sieversii, aimed at studying the features of their distribution in the mountain chernozems of Zailijskei Alatau, in the mountain-brown soils of Talasskei Alatau and in the mountain-steppe soils of Eastern Karatau, showed the near-surface arrangement of the primary skeleton of tree roots (Fig. 2.28). Apple forms well-developed root systems from a great number of vigorously branching horizontal roots, growing directly from the root neck. The main mass of skeletal roots with diameters to 5 cm,

Fig. 2.28. Alatau.

Malus sieversii root system in a deep strongly leached chernozem in Zailijskei

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in deep and medium-deep mountain leached chernozems (humid growth type) is in horizons A1 and A2 (at a depth of 10 to 60 cm). A thicker network of skeletal roots develops in the lower part of the horizon A2 (at a depth of 20 to 50 cm), that is, mainly in the layers with stores of rich nutrient substance. In the lower soil horizons of this type of growing condition of apple, a few skeletal roots are formed. They are formed near the trunk in a radius of not more than 2 to 3 m and are directed downward and are weakly developed. In the alluvial horizon and mother rock, a skeletal root network is mainly formed at the expense of the downward branching of roots. These skeletal roots are distributed in the soil layer down to 20 to 50 cm. An effervescence line often restricts the intensity of the penetration of skeletal roots deep into the soil. Thus, the primary roots, branching into secondary roots, form a thick network and occupy a considerable volume of soil. In the horizontal direction, M. sieversii roots stretch far beyond the boundaries of the tree crown. In the mountain chernozems of Zailijskei Alatau, apple root systems extend rather evenly through all root layers, except the roots in the peripheral zones, which are mainly concentrated in the upper three basic soil horizons. The penetration depth of vertical roots into thick mountain chernozem reaches 3 to 4 m. On mountain slopes, apple root systems spread unevenly. The roots stretching downward on a slope are longer and are concentrated in the shallower soil horizons than are the roots stretching upwards on a slope (Fig. 2.29). Apple is readily able to remodel the morphological structure of its root system in response to environmental change. For example, the formation of new roots from root rudiments in the upper clay layers occurs during the spring flooding period when the main root systems have temporary difficulties. The morphological characteristics of M. sieversii root systems in relation to their growing conditions should be stressed. Under humid growing conditions, apple forms well-developed skeletal roots in deep and medium-deep leached chernozems. However, under very dry conditions in mountain-steppe soils with tight bedding of the mountain rocks, it mainly forms root filaments at the expense of smaller skeletal parts. The structural features of root systems have an adaptive significance for the species and are closely related to the water-nutrient regime of apple growing conditions. A study of the growth and distribution of root systems in basic soil horizons was of great importance for a fuller understanding of the characteristics of fruit plant nutrition. The simultaneous application of root cutting and soil profile methods revealed structural features of apple root systems under different growing conditions and

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Fig. 2.29. Alatau.

187

Asymmetric structure of a wild apple root system on a steep slope of Zailijskei

formulated quantitative indexes of root distribution of soil profiles in relation to their total volumes. The excavation of apple root systems by the root cutting method at the distance of 2, 4, and 6 m from the trunk showed that the main mass of horizontal roots is arranged in the humus horizon: under humid growing conditions (subtype A), 76.7 percent, in humid soils (subtype B), 64.6 percent, in dry growing conditions, 51.9 percent, in very dry conditions, 66.0 percent, in wet growth conditions, 74.8 percent, and also in the mountain-brown soils of Talasskei Alatau (Fig. 2.30a), 72.4 percent, and in the mountain-steppe soils of Karatau, 73.9 percent (Fig. 2.30b). That portion of the root system growing in the 0 to 20 cm soil layer under humid growing conditions of subtype A was 18 percent, subtype B, 23 percent, dry, 18 percent, very dry, 32 percent, and wet, 34 percent. The portion in the 100 to 150 cm layer in humid growing conditions of subtype A was 17 percent, subtype B, 15 percent, while under dry growing conditions, 14 percent, very dry conditions, 4 percent, and under wet growing conditions, 7.3 percent. The largest mass of large roots was concentrated in the uppermost (0–20 cm) layer of the humus horizon, which is explained by a favorable nutrient regime and the good waterphysical properties of this soil layer. Here, the roots grow very quickly and are intensively branched out, with their ends thickly divided.

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Fig. 2.30. Character of a developing wild apple root system: (a) in mountain-brown soils of Talasskei Alatau; (b) in mountain-steppe soils of Karatau.

Under humid growing conditions, the presence of fewer very large roots in the soil profile is characteristic, which undoubtedly is related to a very thick humus horizon and an availability of substantial stores of nutrients. With worsening growing conditions (wet and very dry) the number of very large roots in the upper (0–50 cm) soil horizon increases. One variable that plays a big role is the level of ground water, and another is the exposure to hard mountain rock. These two factors unfavorably influence the development of perennial skeletal parts of apple root systems. Thus, the arrangement of very large apple roots is closely related to a clearly expressed soil differentiation in the basic horizons. Weights of roots taken within 2 m intervals of the trunk showed that the greatest concentration of roots is within a radius of 2 to 5 m from the trunk. However, under very dry growing conditions, most of the very large roots are arranged within a radius of 1 to 3 m, that is, under the tree crown projection (Table 2.28). The coefficient of root branching (the ratio between the weights of the first and the second fractions, expressed as percentage) depends on the nutrients available in the soil. The highest branching coefficient is found in the upper (0–20 cm) soil layer. Correlation between the thickness and the length of roots with diameters up to 1 mm were mainly positive (r = +0.86±0.05). The soil moisture content greatly influences the root distribution of M. sieversii. The selected growing condition types of apple communities clearly differ in relation to this index during the dry second half of summer. During this period under very dry growing conditions, at a depth of 0 to 20 (40) cm, moisture is unavailable for plants, and at a

Medium-deep strongly leached chernozem

Deep strongly leached chernozem

Deep meadow chernozem

Mountain-brown

Mountain-steppe

Humid on slopes

Fresh along valleys

Wet

Humid along valleys

Very dry

Root weight and length based on 1000 cm3 soil.

Medium-deep weakly leached chernozem

Dry

a

Shallow mountain-steppe

Karatau 200 400 600

0.15 0.06 0.06

52 30 45

85 49 32

Talasskei Alatau 200 0.28 400 0.18 600 0.07

361 117 66

245 109 67

116 50 38

161 65 43

0.45 0.22 0.12

0.42 0.26 0.11

0.34 0.11 0.08

76 44 –

Roota length (cm)

0.32 0.13 0.08

200 400 600

200 400 600

200 400 600

200 400 600

Zailijskei Alatau 200 0.16 400 0.07 600 –

Roota weight (g)

0–20 (cm)

0.13 0.09 0.05

0.28 0.19 0.06

0.16 0.08 0.04

0.27 0.18 0.17

0.24 0.11 0.15

0.18 0.10 0.06

0.13 0.09 –

Root weight (g)

54 35 36

94 76 29

65 42 30

102 79 47

95 56 42

63 52 33

51 32 –

Root length (cm)

20–50 (cm)

0.09 0.06 0.05

0.21 0.17 0.07

0.14 0.06 0.04

0.21 0.15 0.19

0.23 0.12 0.10

0.12 0.09 0.06

0.08 0.08 –

Root weight (g)

41 34 27

86 37 32

46 39 24

68 40 40

87 52 31

38 34 33

37 33 –

Root length (cm)

50–100 (cm)

4:19 PM

Very dry

Soils

Distance from trunk (cm)

Depth of profile

Weight and length of M. sieversii roots with diameters up to 1 mm, depending on the growing conditions.

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Mountain system and growing conditions

Table 2.28.

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depth of 20 (40) to 220 cm is barely available. Under arid conditions, the supplies of available moisture in a soil layer of 0 to 200 cm reach 54 mm, but in humid soil, 218 mm, and in wet soil, 320 mm. The most favorable supply of water for apple under humid growing conditions is where active growth of suckering rootlets is observed, especially in the lower horizons of the soil at a depth of 50 to 100 cm. Growth intensity gradually falls under conditions of less humidity. Surface formation of root systems, mainly consisting of a great number of branching, adventitious roots, promotes an effective use of atmospheric water. Such roots instantly take up and absorb water as soon as it penetrates into the soil. At the same time, less atmospheric moisture is evaporated from the warm surface of the soil horizon, and also the deep roots take up less moisture. The more drought-resistant apple stands with their unique root structure, conserve water both in the skeletal roots, though the amount is almost insignificant, as well as in the aboveground parts. They discharge water more economically than their vigorous parent roots which have developed skeletons and more taproots. These penetrate into the soil to depths up to 4 m, reaching the zone of capillary moisture, which ascends up from ground waters. In many waterproviding soil horizons, the skeletal taproot is covered with filaments, which promote a relatively even supply of tree water. The arrangement of the skeletal roots with their adventitious buds in the surface (20–30 cm) layers of the soil has important biological significance for the process of apple natural renewal. These buds, from which trees are formed in due time, are in dormant state during drought periods and differentiate under more favorable conditions of soil moisture. It is natural that among drought-resistant apple trees in steppe zones, the forms with shorter vegetative periods and early maturing fruits prevail. Thus, seasonal changes of water and mineral nutrition are reflected in the root structure and distribution of M. sieversii. A layer of descending roots uses nutrient substances from decomposing litter on the surface mainly in spring, when rains maintain constant moisture in the soil. Later on, when surface soil layers dry, the succeeding root layers join in the process. Deep root interlacing, especially on trees with self-grafting roots, helps M. sieversii to withstand droughty periods and supports its competitive ability against other species. Higher nutrient content distinguishes physiologically active plant parts, such as leaves and small roots. The content of nutrient elements noticeably changes in the different organs and tissues of apple. Of the aboveground organs, leaves are distinguished by a higher concentration of nutrient elements. Here, a direct positive connection between the sup-

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plies of soil nutrient substances and their accumulation in leaves is seen. Greater quantities of nutrient substances result in heightened concentrations in the leaves. So, apple leaves, growing on deep leached chernozems of Zailijskei Alatau contain N (2.6%), P2O5 (0.4%), and K2O (1.8%). On depleted mountain-steppe soils of Karatau, apple leaves contain N (1.5%), P2O5 (0.2%), and K2O (0.8%). We found a correlation between the nutrient content and diameters of very large roots. Thinner roots have higher concentrations of N, P2O5, and K2O. The highest concentrations of elements were found in roots with diameters up to 1 mm (Table 2.29). Thus, M. sieversii is well adapted to mountain leached chernozems and under these conditions it forms a root system, distinguished by extraordinary thickness, considerable depth and by numerous horizontal branchings. Deterioration of growing conditions (very dry type) suppresses root system development, which affects the whole tree appearance, its growth, and its productivity. A direct correlation exists between growing conditions (availability of nutrient substances and moisture reserves) and root system development, and, in turn, between apple growth and apple productivity. The root system structure of an apple tree makes it biologically well adapted to different growing conditions in the mountainous regions of Kazakhstan. E. Growth, Productivity, and Fruit Bearing of Wild Apple During our field studies between 1949 and 1951, we compiled an inventory of wild fruit plant communities, taxonomic descriptions of the woody stands in gorges, and tables listing the species composition of plants growing in apple stands in the mountain regions of Kazakhstan (Dzhangaliev 1951). The course of growth of normal woody stands was determined by the point stand method. For that, 64 experimental areas (1 ha each) were selected in forest stands, which had not been affected by fellings, fires, or other forest-disturbance factors. In order to compare woody stands with each other in a natural developmental series, growth in heights by the largest trees (I–II growth classes by Kraft) were used as models. The trees in the study areas were classified into two thickness categories. The height of each tenth tree was measured. The species were determined by the usual methods, accepted by forest taxonomy. In trial areas, the following species comprised the main canopy: apple, 5,907 or 93 trees per hectare; aspen and other forest species, 1,567 or 26 trees per hectare; in all, 10,107 trees or 158 trees per hectare. The average age of trees, modeled from trees of the main forest canopy, was 55 years. The average stand composition (number of individuals of each

192 200 400 600 200 400 600

Humid along valleys

Dry

Wet

Humid along valleys

Humid on slopes

Dry

0.78 0.56 0.67

0.78 0.76 0.50

0.78 0.73 – 0.93 0.70 0.74 0.98 0.87 0.90 0.99 0.87 0.78 0.56 0.45 0.53

0.27 0.34 0.26

0.35 0.31 0.32

0.24 0.17 – 0.31 0.32 0.35 0.22 0.25 0.24 0.24 0.25 0.21 0.12 0.10 0.11

P2 O5

Karatau 0.64 0.98 0.70

0.58 0.45 0.45

0.10 0.11 0.12

0.31 0.36 0.30

Talasskei Alatau 0.64 0.62 0.82 0.82 0.86 0.86

P2O5

0.28 0.24 – 0.18 0.10 0.15 0.21 0.31 0.25 0.09 0.10 0.14 0.06 0.06 0.08

N

Zailijskei Alatau 0.62 0.67 0.58 0.50 – – 0.68 0.62 0.62 0.68 0.62 0.60 0.70 0.50 0.82 0.63 0.82 0.64 0.80 0.59 0.96 0.48 0.96 0.44 0.88 0.22 0.68 0.28 0.70 0.39

K2 O

Root diameter 1 mm +

0.48 0.62 0.54

0.58 0.77 0.88

0.26 0.32 – 0.32 0.54 0.64 0.36 0.54 0.56 0.48 0.46 0.42 0.84 0.80 0.58

K2O

0.40 0.39 0.37

0.39 0.37 0.37

0.50 0.50 – 0.62 0.58 0.54 0.48 0.54 0.40 0.42 0.26 0.36 0.17 0.28 0.36

N

0.06 0.06 0.06

0.36 0.32 0.35

0.19 0.16 – 0.15 0.11 0.14 0.10 0.08 0.14 0.10 0.10 0.15 0.11 0.12 0.14

P2O5

Large

0.24 0.28 0.32

0.44 0.56 0.77

0.42 0.44 – 0.36 0.34 0.32 0.20 0.25 0.32 0.50 0.33 0.48 0.86 0.78 0.54

K2O

4:19 PM

200 400 600 200 400 600 200 400 600 200 400 600 200 400 600

N

<1 mm

8/8/02

Very dry

Distance from trunk (cm)

Content (% dry weight)

Effect of diameter and distance from trunk on the nutrient element content in M. sieversii roots.

Mountain system and growing conditions

Table 2.29.

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species) is characterized by the following formula: 6 apples, 2.5 hawthorns, 1.5 aspen. The data on tree numbers per hectare and their ages testify to the fact that the thickness observed makes up approximately 60 percent of their normal density relative to the stages of crown closure. If woody stands with a predominance of wild apple are considered as fruit orchards, then the index for the average number of trees per hectare is similar or superior to all orchards cultivated for fruit on vigorous rootstocks in the southeast of Kazakhstan. Apple woods, from the point of view of forest structure, have a density which is approximately 20 to 25 percent that of the normal forest density. However, it is necessary to take into account that the light-demanding apple, engaged in the formation of its fruit organs and normal fruit-bearing, prefers a thinner tree arrangement over that of the more shade-tolerant forest trees which produce wood. Apple stands at a long distance from large populated areas on plots without fellings or cattle grazing form rather dense woody stands with the characteristic structure of real forests (Table 2.30). Based on the distribution of forest-covered areas by density and by a common measuring scale of forest tree closure, in Kazakhstan, apple forests with a thin average density of 0.1 to 0.2 constitute 9.3 percent of the total apple forest area, with a low density of 0.3 to 0.4, 40.2 percent, with an moderate density of 0.5 to 0.6, 32.2 percent, and with a high density of 0.8 to 0.9, 6.8 percent. Depending on biotic and abiotic conditions, apple stands have great variation in both dominant canopy composition and productivity. On the slopes with northern exposures where deep chernozem soils prevail and where human influence is less noticeable, apple forms rather closed thickets with densities between 0.5 to 0.7 and 0.8. A determination of tree-stand density according to the degree of crown closure showed that the density in some virgin apple groves in the middle mountains of Dzhungarskei, Zailijskei, and Talasskei Alatau Table 2.30.

Percent area of stand density classes within mountain apple forests. Area (%)

Mountain system Tarbagatai Dzhungarskei Alatau Zailijskei Alatau Talasskei Alatau Karatau

0.1–0.2

0.3–0.4

– 29.8 11.1 5.4 –

43.8 36.6 38.6 44.6 37.6

Density class 0.5–0.6 42.6 20.3 28.5 21.2 48.2

0.7

0.8–0.9

13.6 6.5 10.3 16.7 10.3

– 6.8 11.5 12.1 3.9

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reaches 0.8 to 0.9 and even 1.0. Densities of tree stands such as these are never seen in other mountain regions of Kazakhstan or in low and high mountain populations, which will be explained later by naturalhistorical conditions. On slopes of southern exposures, where soils are generally gravelly rubble, and on steep slopes and on stone mounds, apple tree stands are thin. Their density according to the common bonitet scale is between 0.2 and 0.4. Data on distribution of forest covered areas according to bonitets show that apple woods of V and Va bonitets constitute 0.8 percent, III and IV bonitets, 60.3 percent, and I and II bonitets, 39.9 percent (Table 2.31). Because of the lack of approved bonitet tables, fruit-tree bonitet is often determined according to a common scale, which naturally adversely affects the exactness of the cited data. Perhaps, due to the error of the bonitet, M. sieversii communities on rich chernozem soils in the gorges of Osinovoe, Dalnee, Mikushino in Zailijskei Alatau and the Chernaya Rechka, Soldatskoe in Dzhungarskei Alatau are mainly attributed to III and IV bonitets. Because of the growth character and the incremental rate of increase in the wood mass, M. sieversii may be referred to as a forest tree of the third size. By studying the rate of growth in height, and barring any traces of fire or fellings, the following features of this process are evident: in closed stands, trees are rather slender, while in thin overgrowth, tree stems are branching. Growth culmination and maturity begin at 45 years of age. The total amount of vegetative mass produced by apple stands at 65 years of age, in better growing conditions reaches 134 m3/ha, and in the poorest growing conditions, 31.3 m3/ha. The central shoot shows that, independent of abiotic conditions, the intensity of apple stand growth regularly decreases after the age of 10 to 15 years (Table 2.32). The table data characterize the apple as a slowly growing and longlived species. At the age of 65 years, it ceases its great growth cycle, but Table 2.31. Distribution of areas covered by apple forests according to bonitet classes. Area (%) Mountain system

I–II

III–IV

V–Va

Tarbagatai Dzhungarskei Alatau Zailijskei Alatau Talasskei Alatau Karatau

7.0 69.2 84.3 5.1 29.1

90.1 30.8 15.1 94.4 70.8

2.9 – 0.6 0.5 0.1

5 10 15 20 25 30 35 40 45 50 55 60 65

2.0 3.0 4.0 4.5 5.0 6.0 7.0 8.0 8.5 9.0 9.5 10.0 10.0

Average height (m) 2 4 6 8 11 15 19 23 27 31 35 39 44

Average diameter (cm) 0.32 0.51 0.54 0.53 0.51 0.50 0.46 0.40 0.38 0.36 0.34 0.33 0.32

Visible no. – 0.002 0.006 0.013 0.024 0.046 0.081 0.123 0.170 0.215 0.292 0.378 0.500

With bark – 0.002 0.006 0.012 0.023 0.044 0.074 0.112 0.158 0.210 0.277 0.360 0.475

Without bark

Average tree vol. (m3)

0.25 0.38 0.50 0.63 0.68 0.64 0.61 0.59 0.54 0.52 0.47 0.45 0.41

Coefficient of trunk form 6.7 10.0 10.0 8.3 5.7 3.8 2.4 1.7 1.0 0.7 0.3 0.1 0.1

Current increment (%)

2.08 2.08 2.18 2.24 2.27 2.31 2.26 2.17 2.09 1.93 1.84 1.57 1.38

Current

2.08 2.05 2.07 2.16 2.19 2.24 2.22 2.20 2.15 2.12 2.07 2.03 1.96

Average

Increment (m3)

4:19 PM

Age (years)

Growth rate of normal M. sieversii stands in Zailijskei and Dzhungarskei Alatau.

8/8/02

Table 2.32.

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continues to bear fruits. For example, during this investigation of growth rates, an exemplar tree was cut down in the Pikhtovka gorge of the Topolevskei forest (Dzhungarskei Alatau) at the age of 136 years without any heartwood rot and from which 1100 apples were picked up. Hundred-year-old trees with similar features are not exceptions. Longevity of the wild apple is especially apparent in comparison to cultivated orchards, in which the life cycle is completed within 45 to 50 years. The coefficient of trunk form, which characterizes tree tapering and depends on tree age, was between 0.25 and 0.68 for apple. The largest value for this coefficient was observed between the ages of 20 and 30 years, for trees growing within a group under conditions of side shading. The M. sieversii trees of Zailijskei and Talasskei Alatau are mainly medium-sized, most populations having trees 5 to 8 m in height. Rather few populations have trees that are very high (8 to 10 m or higher) or low (to 3.5 m) in stature. In the middle mountains of Dzhungarskei Alatau, vigorous trees (on average, 7.7 m) prevail, while in Karatau and Tarbagatai, apple trees are dwarfish. On chernozem soils of the northern slopes, adult trees reach 8 to 9 m in height (Fig. 2.31). Individual trees in the middle mountains of Dzhungarskei Alatau are gigantic, 15 to 16 m in height (Fig. 2.32), but on stone debris of the southern slopes, trees are small-sized, 4 to 5 m in height and tapery (Fig. 2.33). On the best soils, free-growing apple trees in full light form welldeveloped plane-spreading, oval-spherical, pyramidal crowns, with adjustments in their shape depending on individual tree characteristics. In closely spaced apple stands, where tree interdependence is prominent, tree shapes are differentiated according to developmental classes.

Tree height (m) Southern slope

9 8 Northern slope

7 6 5 4 1000

1200

1400

1600

1800

2000

Site altitude (m) Fig. 2.31. Influence of site altitude and slope exposure on wild apple tree height (75–100 years old).

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Fig. 2.32.

197

Wild apple giant (Dzhungarskei Alatau, Pikhtovka gorge, 1230 m above sea level).

The trees of Class I have well-developed crowns, which rise somewhat above the common forest canopy. Trees of Class II also have welldeveloped crowns, but do not rise above the forest canopy. The trees of Class III and other classes are variously deformed depending on their interdependence characteristics. Here, one may find trunks with compressed crowns on each side or with lopsided or partially dead crowns. The typical character of apple growth is in thickets (height, 1.0–1.5 m), mainly with an inclined arrangement of skeletal branches. In the upper parts of spreading apple trees, the stems develop in a bushy arrangement. Fruit-bearing is the main index of M. sieversii vitality. A. N. Formozov (1964) showed that the yield of fruits and seeds of forest trees, which are fundamental forage for animals, are of great importance for

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Fig. 2.33. Dwarfish wild-apple selection (Dzhungarskei Alatau, Pikhtovka gorge, 1540 m above sea level).

ecosystems, especially for communities with many species. Fruits of different woody plants create the conditions under which animals in mixed forests, such as fruit forests, are provided with food from year to year. By foraging on fruit and seed, animals promote the distribution of plant seeds throughout their territories. When considering the fruit-bearing role of M. sieversii, it is necessary to emphasize the importance of the duration and preservation of normal fruits and seeds under natural conditions. On the ground of apple stands, many times we found apples from the last year’s crop and seeds from the last year in the litter. Judging from their appearance, seeds were of good quality, leading to the possibility that year-old seeds can participate in natural renewal, even in a year of light harvest. V. T. Langenfeld (1970) believes that in fruit evolution, a great role was played by the increase of apple size by the expansion of juicy flesh. Apparently, the best forms of wild apple fruits attract animals because of their high content of nutrient and biologically active substances (Dzhangaliev 1974a). Tree age composition characterizes not only the status of modern apple stands but also helps to understand in which direction natural renewal will develop, either toward plant community regression or new

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199

Age composition of apple trees in different mountain system areas. Composition (%)

Mountain system Tarbagatai Dzhungarskei Alatau Zailijskei Alatau Talasskei Alatau Karatau

Young I – II

Middle-aged III

Becoming mature IV

Mature and overmature V

– 33.9 41.2 14.8 18.4

68.8 51.7 25.1 45.3 27.0

17.1 13.5 30.5 33.6 11.8

14.1 0.9 3.2 6.3 42.8

generation initiation. Table 2.33 lists apple stands as young, middleaged, becoming mature, and partially overmature. Apple age composition is well-distributed. Young trees in age Classes I and II (up to 10–12 years) are more numerous than old dying trees (over 40 years). Trees of Classes III and IV are growing and are in full fruit-bearing (from 10–12 to 60–70 years), and they, on the average, include 32.4 percent of all trees. Thus, the fruit-bearing portion of the stand is constantly being replenished by young trees that enter into a growth period of fruitbearing. They become a seed source for natural renewal. Because of the different aged trees in apple stands, years of an absolutely bad harvest are never seen. New trees of wild apple, both from seed and vegetative origin, enter into bearing at the age of eight to ten years, and they produce heavy yields beginning from 12 to 20 years of age. In relation to tree age composition, the tendency toward renewal and multiplication prevails at the expense of young trees. These investigations showed that apple fruit-bearing depends on the environment and on the individual hereditary properties of trees. In view of the great biological and practical importance of fruit-bearing and also due to the lack of data and the difficulty of getting reliable yield information from wild fruit communities, the determination of apple stand fruit-bearing was calculated by a complex method. This was the Kapper-Formozov scale, which measures the crops on trees in experimental plots. It also includes biological tree observations and a study of fruit organs (Shitt 1952). Estimates of apple fruit yields during routine travels were made using a 6-rank scale. Five trees in each age group at an altitude of 1100 to 1900 m (Table 2.34) were under observation. Fruit bearing periodicity characterizes apple stands. Heavy yields regularly alternate with average and very low yields. Apple fruit bearing is also related to the site of growing (Table 2.35).

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Table 2.34. Fruit bearing index of M. sieversii populations according to the six-point scale of Kapper-Formozov. Fruit bearing index Mountain system Tarbagatai Dzhungarskei Alatau Zailijskei Alatau Talasskei Alatau Karatau

1971

1972

1973

not determined 3.7 3.4 not determined –

1.9 2.4 2.0 2.9 1.7

not determined 3.5 3.3 3.0 1.9

Apple fruit cropping is the heaviest in the belt of mixed broad-leaved forests and the least in the steppe zone. Light cropping is caused by a moisture deficit and frequent below-zero temperatures in the steppe zone during the apple blooming period (Dzhangaliev 1972). These observations showed that pure apple stands seldom occur in nature, but in high-yield years, they produce higher crops (3.6) than mixed apple stands (3.2). However, yields in mixed apple stands are even less in years of unfavorable weather conditions (1.8 versus 2.6, respectively). A determination of the fruit productivity of selected apple forms, according to flower-bud set and an estimation of the current-year yields on representative branches of fruit-bearing trees, showed the following dynamics of biological yield in 1969 to 1970. Fruit yield predictions according to flower bud set for these years are considerably higher than they are on Kapper-Formozov scale. This lower expected productivity is explained by local frosts. In 1969 in Zailijskei Alatau, frosts damaged a significant portion of the flower clusters on the trees, while in 1970 Table 2.35. Estimates of the average apple cropping in Dzhungarskei Alatau according to the type of growing conditions (Kapper-Formozov scale). Kapper-Formozov scale Growing conditions Very dry Dry Humid on slopes Humid in valleys Wet

1971

1972

1973

1.9 2.1 3.7 3.9 2.7

1.4 1.8 2.9 3.2 1.6

2.0 2.3 3.5 4.1 3.1

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cold, pouring rains during the period of pollination and fertilization prevented insect flying and thus prevented apple cross-pollination. A yield decrease in 1972 is also explained by the heavy cropping of apple stands in 1971, which led to a reduction of the nutrient stores of trees and slowed down the initiation of generative organs for the next one to two years. Also, a negative influence of insects (leaf roller, apple moth, codling moth) and scab occurred during this period. In apple stands intensively used for cattle grazing (Stepnoi leskhoz of the Semipalatinsk region), weak fruit-bearing was caused by poor soil conditions and an unsatisfactory tree status. Cattle ate their annual shoots at the lower crown levels, their leaf canopies, and their overgrowing branches. Here, apple trees bore fruits when they were closely surrounded by prickly overgrowths of Amygdalus ledebouriana or Rosa spinosissima. These not only protect the trees from cattle but also favorably influence the microclimatic and the soil conditions in the ecosystem, which is evident from the data in a comparative study of these conditions in apple stand communities (Dzhangaliev 1973a,b). It was also discovered that apple buds at their phase of bud-break, die at a temperature of –8°C, flower buds die at –6°C, flowers die at –3 to 5°C, and ovaries at –3 to 4°C. In the premountain and mountain regions of Zailijskei Alatau at an altitude of 1000 to 1500 m (following a stable transition of average daily temperatures to 10°C in the spring), these cited temperature levels may cause damage to emerging plants in 10 to 30 percent of the cases (Dzhangaliev 1972). Generally, hail rarely influences apple stand yield, but, in June 1973 in the belt below 1200 m, hail damaged a considerable portion of the very young fruit. V. D. Utekhin (1964) refers to M. sieversii in Talasskei Alatau as a medium-productive species. Our investigations showed that a complete lack of fruit yield in apple stands is never seen within the area as a whole. In years of very light harvest, some plots with fruit-bearing levels of only 1 to 2 points are encountered, which side by side with vegetative renewal, provide adequate seed propagation. Productive and nonproductive apple trees are unevenly scattered through the territory. Productivity is influenced both by the variation of natural conditions among wood stands and by the genetic properties of the trees. Fruitbearing by the whole stand as a unit may depend mainly on ecological conditions, but in individual trees, fruit bearing can be related to the hereditary peculiarities of that specific tree. In various parts of their range, forest trees produce different amounts of fruit. Under conditions of mass crop failure, individual trees or even whole populations bear fruits well. However, sometimes when apple stands on the whole produce heavy fruit yields, some trees do not any bear fruits. Studies of

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apple branches in winter and at the beginning of spring showed individual variability for potential blossoming, expressed as flower bud set, that depends on the genetic characteristics of the tree. Groups of fruit-bearing apple woods are spread over the middlemountain regions at an altitude of 1100 to 1500 m, on chernozem soils, and under conditions of favorable moisture-temperature regimes for their growth and development. In such plant communities, high-yielding forms mainly grow up to 1400 m, but above that, the percentage of lowyielding forms gradually increases, associated with exceptionally unfavorable heat resources for any apple variety. In the mountain regions between altitudes of 1400 to 1450 m and 1700 m, only specific early maturing forms become ripe. At altitudes of 1800 to 1900 m, under the most severe of apple fruit-bearing conditions, the number of average and low-yielding forms increases. Interdependence was found to exist between apple stand yield and soil moisture relative to the site altitude. The optimal level of soil moisture is above 70 percent of field water capacity for fruit bearing, which corresponds to the humidity of capillary breakage (Dzhangaliev 1972). The yields in the mountain regions increase with a rise in the altitude of the site, but only to a definite level, above which low temperature is a limiting factor (Fig. 2.34). The fruit-bearing characteristics of apple were studied on 37 experimental areas (1 ha each), laid out on various slopes and exposures, between the altitudes of 1000 and 2000 m, in stands of various ages (Table 2.36). Here, out of a total of 7658 apple trees, 4933 were fruitbearing. They were divided into four groups according to the amount of fruit they bore. Trees with single fruits were in the first group; trees with 1 a light crop (fruit covering less than ⁄2 crown) were in the second; trees

Yield (kg/tree) 250 200 150

Southern slope

Northern slope

100 50 0 1000

1200

1400

1600

1800

2000

Site altitude, m

Fig. 2.34. The influence of site altitude and slope exposure on wild apple productivity in Zailijskei Alatau.

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2. THE WILD APPLE TREE OF KAZAKHSTAN Table 2.36.

Tree number

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43

203

Morphological characteristics and fruit-bearing of representative trees.

Tree Tree Slope Slope age height exposure (% grade) (years) (m)

SW “ “ “ “ NW W SW SE “ “ “ NE “ SW W “ NW “ “ “ SW “ “ “ “ E “ NW “ “ “ “ SE S “ “ “ “ “ “ W “

15–20 15–20 15–20 15–20 15–20 5 5 6–8 5 5 15–17 5–7 5–10 5–10 15–20 3 3 15–20 10–15 10–15 10–15 5–10 10–15 10–15 10–15 10–15 10–15 10–15 5–10 5–10 5–10 5–10 5–10 10–15 5–10 5–10 5–10 5–10 5–10 5–10 5–10 10–15 10–15

55 51 60 55 55 46 65 65 65 58 50 35 65 55 50 50 15 30 32 34 36 65 101 93 85 68 46 72 50 65 45 70 60 72 62 60 55 75 80 65 70 76 65

8.8 8.9 9.0 8.8 8.8 8.0 8.5 8.5 8.5 7.5 7.5 8.0 9.2 8.5 7.0 8.2 4.0 7.5 7.0 7.2 7.5 7.0 8.5 8.0 8.6 8.5 6.5 7.0 8.0 9.0 8.1 7.2 9.1 8.4 6.7 9.4 7.6 8.4 8.0 7.2 6.3 8.2 7.8

Diameter near Crown stump length (cm) (m)

36 37 38 36 44 46 65 39 34 30 35 23 52 32 24 50 10 19 29 37 32 40 35 33 33 31 53 42 36 45 31 34 95 70 58 64 80 53 70 55 57 39 32

8.0 7.0 8.0 7.4 7.5 7.0 7.0 7.7 7.9 6.1 6.0 8.0 8.4 8.0 5.0 8.0 3.2 6.0 6.0 4.0 4.0 6.0 4.5 4.0 6.0 5.0 4.0 4.5 4.5 6.0 6.0 6.0 6.5 5.5 6.0 5.0 5.2 6.0 5.8 6.0 5.0 6.7 6.2

Crown diam. (m)

No. skeletal branches

Yield (kg)

8.0 5.0 7.0 8.0 9.0 13.0 13.6 8.5 9.0 4.0 4.0 7.5 9.0 8.0 4.0 4.0 2.0 6.0 4.0 4.5 4.5 7.0 6.5 6.5 5.5 7.5 8.5 8.0 8.0 8.0 6.0 7.7 9.0 7.0 10.0 10.7 11.0 10.5 11.2 11.0 8.2 7.0 7.0

3 2 1 1 3 4 7 5 4 3 4 3 6 4 3 4 2 3 1 3 2 3 2 2 2 3 3 2 4 3 2 2 3 3 4 6 4 4 4 4 4 3 2

55 80 63 49 98 426 850 300 140 99 21 66 262 200 51 152 3 71 16 34 34 105 55 40 35 113 80 73 71 130 103 53 142 193 117 256 220 116 102 156 148 169 116

(continued)

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Table 2.36.

Tree number

44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86

(continued)

Tree Tree Slope Slope age height exposure (% grade) (years) (m)

“ S “ “ “ “ “ “ “ NW SW “ NW “ “ “ “ N “ E “ SW E “ “ NE “ “ “ “ SE “ “ “ “ SW “ “ W “ “ “ SE

10–15 5–10 5–10 5–10 5–10 5–10 5–10 5–10 5–10 15–20 15–20 15–20 15–25 15–25 15–25 15–25 15–25 5–15 5–15 10 10 15–25 10–20 10–20 10–20 10–15 10–15 10–15 10–15 10–15 10–20 10–20 10–20 10–20 10–20 5–10 5–10 5–10 15–20 15–20 15–20 15–20 20

40 30 20 30 45 15 45 55 45 50 45 40 25 35 23 30 65 45 25 30 30 30 45 48 65 45 40 30 40 25 40 45 50 25 35 65 40 25 50 25 18 18 62

6.0 6.5 4.5 7.2 7.1 5.0 8.1 11.7 8.0 8.5 8.1 7.6 6.3 7.2 6.1 6.4 9.4 7.6 5.7 6.4 6.5 7.5 8.1 8.6 9.4 7.0 6.5 5.0 6.2 6.3 8.0 9.2 7.6 4.5 6.5 8.4 7.9 7.5 8.1 6.0 5.0 5.0 7.5

Diameter near Crown stump length (cm) (m)

24 16 11 24 39 14 36 44 32 32 28 46 16 30 22 8 33 30 22 23 20 22 26 36 40 31 25 24 32 19 26 36 31 14 22 58 22 17 47 18 16 16 32

5.0 6.5 3.6 4.0 5.0 3.0 7.0 8.2 6.5 7.9 7.9 6.4 5.0 6.0 6.0 5.4 8.0 6.6 4.0 4.0 4.0 4.5 7.0 7.4 8.2 4.0 5.0 3.5 4.2 3.3 4.0 6.0 3.0 3.0 3.0 6.0 4.5 3.5 7.0 3.0 4.0 3.0 6.2

Crown diam. (m)

No. skeletal branches

Yield (kg)

6.0 5.0 4.0 5.0 7.5 3.0 6.0 7.8 8.0 6.2 7.2 4.8 5.7 6.0 5.6 5.5 7.5 7.5 4.0 5.0 4.0 5.6 6.0 7.0 9.0 5.0 4.0 3.5 6.0 3.5 4.0 7.0 5.5 3.0 4.0 8.5 5.8 3.5 9.0 4.0 2.0 1.5 9.0

2 1 1 2 2 2 2 2 1 2 2 1 1 2 2 1 5 2 3 2 2 3 2 2 2 1 1 1 2 1 2 2 1 1 1 4 1 1 4 2 2 2 2

94 7 1 74 65 2 44 135 93 33 121 98 34 98 61 50 103 121 30 27 19 33 46 78 144 170 66 39 141 17 50 92 64 23 29 325 37 17 70 18 9 2 178

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Tree number

87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130

Tree Tree Slope Slope age height exposure (% grade) (years) (m)

“ “ SW “ “ “ “ “ NW “ “ SE “ “ E “ “ NE “ “ SW “ SE “ “ “ “ “ “ “ NE “ E “ “ “ “ “ S “ E W “ E

20 20 5–15 5–15 5–15 5–15 5–15 5–15 10–15 10–15 10–15 10–15 10–15 10–15 10–20 10–20 10–20 10–20 10–20 10–20 10 10 5–10 5–10 5–10 5–10 5–10 5–10 15 10 15–20 5–10 15–20 15–20 15–20 15–25 15–25 15–25 5 5 10–20 15–20 15–20 15–20

55 45 72 60 15 55 45 25 55 60 50 45 20 50 45 40 50 60 65 75 75 95 65 65 65 90 85 60 25 50 25 75 60 55 45 45 40 35 75 65 60 45 65 45

8.2 7.5 7.9 8.6 4.0 9.4 9.6 7.2 9.6 10.4 10.0 8.0 5.0 8.0 8.1 7.6 8.5 9.0 8.0 7.0 9.1 9.8 7.2 7.8 9.0 10.3 8.8 9.0 4.6 8.0 5.0 8.8 9.2 8.8 8.1 8.0 8.0 8.4 8.2 7.8 9.2 8.8 9.4 7.2

205

Diameter near Crown stump length (cm) (m)

38 38 63 60 6 42 36 22 36 47 39 36 14 39 29 26 30 49 42 46 66 66 40 40 48 74 52 46 10 40 14 44 52 55 25 38 34 35 76 52 36 28 40 34

7.3 6.0 6.5 6.6 3.0 9.0 9.0 6.0 8.2 9.0 9.0 7.6 4.5 6.0 7.3 7.0 7.5 8.0 6.8 6.1 7.1 8.0 6.0 6.5 7.8 8.1 6.5 8.0 2.6 7.0 4.0 7.3 8.0 8.0 5.4 7.0 7.0 6.0 7.0 7.0 8.0 7.0 8.2 5.5

Crown diam. (m)

No. skeletal branches

Yield (kg)

10.0 7.5 7.0 10.2 4.0 7.0 8.0 6.0 8.0 9.0 6.0 6.0 4.0 6.0 7.0 5.0 1.0 7.0 7.0 6.0 11.0 10.0 6.0 5.0 8.0 8.0 8.0 8.0 3.0 8.0 4.0 8.0 8.0 8.0 4.0 6.0 6.0 6.0 8.0 9.0 9.0 7.0 7.0 5.0

4 2 – – 3 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

305 60 155 475 8 136 212 25 130 475 80 140 300 250 139 50 250 370 60 150 730 620 120 80 230 350 50 250 10 180 10 220 160 210 40 266 150 50 650 740 80 120 80 22

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1 with medium crop (fruit covering ⁄2 crown) in the third; and trees with heavy crops (fruit over the whole crown) in the fourth. From the total number of fruit-bearing trees, 130 representative trees were selected on which yields per tree were measured. Among these representative trees, 13.2 percent bore heavy crops (40–80 kg per tree); 31.4 percent bore average crops (20–40 kg), 28.4 percent bore poor crops (to 20 kg), and 27.0 percent bore very little crop. On the whole, 44.6 percent of the trees bore heavy or average crops. 17,341 kg of fruit were collected from the 130 representative trees, which is an average of 133 kg per tree. On individual, well-developed trees, yields reached record figures. From tree No. 7, growing on a mediumdeep leached chernozem on a western slope in the gorge of Chernaya Rechka in Lepsinskaya woodland, 850 kg of apples were harvested. The tree height was 8.5 m; the trunk diameter was 65 cm; the average crown diameter was 13.6 m; the number of skeletal branches was 7; and the foliage was good. The rate of fruit-bearing increases until 45 years of age, then production evens out. Full fruit-bearing continues until 65 years of age:

Tree age (years) 10 15 20 25 30 35 40 45 50 55 60 65 Fruit bearing (kg per tree) 4 11 22 33 43 51 59 65 63 54 45 36 Fruit bearing gradually falls from its highest point down to 100 years of tree life. At the age of 100 years, on average fruit bearing is 21 percent of maximum. Thus, M. sieversii populations are usually described as rather longlived, slow growing, low-stemmed, and medium productive trees. Moderately vigorous trees constitute the population base, but in high-bonitet apple groves of the middle mountains (in the zone of optimum ecological conditions) giant trees are seen. Under extreme conditions in the high mountains, bushy tree populations are formed. In tight thickets, apple trees are differentiated by developmental classes. As forest builders, they take part in the composition of development classes I and II, when their well-formed crowns rise above or are part of the whole forest canopy. Tree crowns of other classes, including class III, are badly developed, and their role in forest formation is secondary. The principal growth cycle of an average tree ceases at the age of 65 years, but the fruitbearing period continues until 100 years, and for some trees, it is longer. A comparative analysis of the growth stage index and apple fruitbearing testifies to the economic utility of the apple stands for fruit products and for seed production. This utility also points to the necessity of stopping tree felling for wood. Apples should be regularly harvested

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from high- and medium-density apple stands, but from low-density apple stands, only in years of heavy yields. VI. INTRASPECIFIC POLYMORPHISM OF WILD APPLE Polymorphism within one species refers to several clear morphologically and pomologically different forms; the formation and steady maintenance in the population of two or more genetically and phenotypically different forms, conditioned by mixed habitats and adaptation of new types to the environment. As a result of our long-term field and experimental investigations of the gene pool of wild apple species growing in the natural forests of Kazakhstan, we are convinced that each species differentiates into numerous intraspecific taxa (phenotypes), which possess many qualities with unique biological and economic values. We selected tree phenotypes of wild apple, which can be directly introduced into cultivation to establish orchards or may be used as selections to breed stress-resistant cultivars. Nature itself, by spontaneously creating polyploid individuals with colossal intraspecific variety, promoted the success of apple domestication for many millennia. Many breeders, and especially I. V. Michurin (1948), when crossing cultivated apple with a number of these accidental wild types, noticed a dominance of negative characters in the hybrids, such as small fruits, numerous spines on trees, and so on. These observations, a result of insufficient scientific data on individual genotypes and intraspecific taxa of Malus, resulted in a negative view of wild polymorphic species for the improvement of cultivated varieties, now suffering from genetic erosion. Up to now, breeders have selected a pair of cultivars for crossing from the so-called world collections, not knowing about nor obtaining wild genes of phenotypes with established physical features which we revealed in the wild apple of Kazakhstan. Our studies on the polymorphism of the wild apple were based on N. I. Vavilov’s (1927) idea that resistant seed plants for breeding should be obtained from their original geographical native land. This proposal of N. I. Vavilov is directly connected with his theory about gene centers leading to the origin of cultivated plants. Accordingly, after his visit to the apple forests around the town of Alma-Ata in September 1, 1929, he came to the conclusion that “It was possible to see with our own eyes that here we were in a wonderful center of cultivated apple origin.” The primary Kazakhstan gene center has a decisive role in world agriculture, because the structural gene pool of species and intraspecific populations were historically established here. In this connection, nowadays it is impossible to identify a species only on the grounds of classical comparative plant morphology as previously done.

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Investigations in intraspecific polymorphism of the wild apples of Kazakhstan were conducted using our elaborate method for genetic inventory, selection and classification of apple forests according to the phenotypic, and pomological characteristics of individuals in plant stands and populations. These studies showed that species have been differentiated into local phenotypes as a result of an interaction between genetic recombination and natural selection. A decisive role in the stratification of populations has been played by the spontaneous hybridization of individuals in heterogeneous apple populations and by natural evolution. These processes led to the accumulation of favorable adaptations, because unfavorable ones are selected against. By this means, intraspecific polymorphism appeared, that is, by the formation and steady maintenance in the population of more than two genetically and phenotypically different forms. The intraspecific genetic resources of Malus are large. Proper consideration is not given to their stable preservation and rational use. There is much more biological genetic variation in wild phenotypes than there is in cultivated types of M. domestica, which number more than 10 thousand varieties in the world today. The natural hybrids of wild apple are of great interest, because in nature practically unlimited possibilities exist for selecting valuable forms among individuals with new properties and characteristics, which are not yet introduced into cultivated plants. Studying the polymorphism with populations of Malus species was first conducted in the period of botanical field investigations by identifying elementary phenotypical features in natural populations, expressed in different areas and vertical zones of their range. Intraspecific taxa of Malus from our collection of phenotypes were described according to the duration of flowering period, age composition, height, branching characteristics, crown form, length of vegetative period, winter-hardiness, resistance to disease, yield, and pomological characteristics of fruits. All forms were heterogeneous in each of the properties and characteristics studied. The number of trees with clearly differentiated, distinctive properties and characteristics is so great that each apple tree can be described as an independent form differing from neighboring ones. For instance, trees grown in one location have early, late, or intermediate times of blooming and fruit ripening; various vegetative periods; early and late initial bearing; small, large, and middle sizes of fruits; fruits with white, yellow, or pink color; high-stemmed and low-bushed with different branch and crown structures; and different levels of cold-resistance, immunities, and productivity. The wide range of variability of most morphological characteristics can appear within a single apple stand or population growing on a sin-

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gle plot. In spite of the generally accepted division of variability into phenotypic and genotypic traits, it is very difficult to classify the enormous phenomenon of individual apple differences in nature using such terminology. In this connection, we conducted an investigation on the variability of the characteristics of some important biological and economic features of different forms growing under different conditions. In order to determine hereditary characteristics, a hybrid analysis of some hybrid lineages, bred by the crossing of wild forms was also made. These investigations are of particular significance concerning the theory and practice of genetic assimilation of species characteristics, which are especially important in studies of selection and introduction. Before discussing the analysis of Malus polymorphism by means of our specimens, it should be noted that up to the present time, the general system of evolutionary phenomena and factors underlying the formative process, has been described by many scientists (Dubinin 1967; Dubinin and Glembotsky 1976; Lobashov 1967; Timopheev-Resovsky et al. 1973). Proceeding from the general concept of evolution, the vast intraspecific polymorphism of the apple is the result of the genetic richness within populations, the appearance of mutations during the developmental process, and natural selection under the influence of complicated mountain environments. Each apple population differs from the others by specific features of variability, because for a long time the individuals were intercrossing on limited territory and under definite soil-climatic conditions. Populations develop under the influence of the concrete conditions of existence, as a result of the interaction of hereditary factors, variability and selection. The initial cause for numerous intraspecific apple variants was the genetic variability from mutation, influenced by external and internal factors. This results in new traits in the population and also in many kinds of crossings among trees. Individual trees cross with each other within a population and also between different populations of a species, and under close proximity, between kindred species, such as M. sieversii, M. kirghisorum, and M. niedzwetzkyana. In some cases, individuals from different genera cross (Popov 1929; Dubinin and Glembotsky 1967; and Zhukovsky 1971). This wide distribution of hybridization processes gives evidence of their evolutionary role. Cross-pollination of apples among and between populations establishes genetic connections across wide areas. Isolation, when due to various geographical and biological barriers, can disturb the entire pollination scheme, because borders divide the neighboring populations or a single population internally into separate groups. Some essential properties of apple populations are their territory of distribution, the

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possibility of mutual crossing of individuals, the number of individuals and age composition, and the influence of different environmental factors on the level of selection pressure and on such biological apple features as “the radius of individual activity” (Timopheev-Resovsky et al. 1973, p. 66). All of these properties influence the isolation or unification of the population. Sharply expressed horizontal and vertical boundaries create a landscape mosaic that is characteristic of the apple forests of Kazakhstan. The isolation of species and apple populations in different parts of the species habitat in the mountains of Tarbagatai, Dzhungarskei, Zailijskei, and Talasskei Alatau is absolute, because desert spaces and large territory breaks between the mountain chains rise up in front of populations as insurmountable barriers. Undoubtedly, isolation of parts of the Malus area is related to the species’ history and to the long-term history of the landscape within this geographical territory. As A. I. Tolmachev (1954) showed and then as N. V. TimopheevResovsky et al. (1973) confirmed from the genetic point of view, a species’ fate is inseparably linked with the physical-geographical and regional conditions in which it grows, and with the community of other species of plants, animals and microorganisms. Consequently, any further evolutionary fate of the species, in particular its character and distribution trends, depends on the fate and distribution of the whole community. Judging from the characteristics of the species’ territory (Fig. 2.1) and the number of individuals within, it is possible to come to some conclusions about the history of Malus species in Kazakhstan. Dominating and being the most widely spread species, numerous population groups of M. sieversii are seen in all parts of its territory, occupying a wide vertical belt and various types of habitats. In other words, M. sieversii is the main component of fruit forests. In contrast, M. kirghisorum has a limited range in Dzhungarskei and Zailijskei Alatau, growing under more mesophilous conditions in the middle mountains. The differences between these two species are explained by the characteristics of their distribution after the period of maximum glaciation during the pleistocene period. M. G. Popov (1940) and other investigators consider that the modern distribution of such species are the residues of earlier species that existed in continuous areas, which were broken during the glacial period. Malus kirghisorum, growing together with remnants of primitive walnut forests and more closely related to the initial tertiary apple than M. sieversii (Al. A. Fedorov and M. A. Fedorov 1949), found its refuge in the favorable habitat of the mountains during the period of maximum glaciation. Later on, during the process of changing to the desert types of vegetation (Popov 1940), M. kirghisorum did not widely distribute or develop in the severe and

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cold conditions of Tarbagatai or in the arid conditions of Karatau, where instead, M. sieversii became the main component of fruit forests. Malus sieversii is also observed in the severe upper borders of the mountain belt in Dzhungarskei and Zailijskei Alatau, where M. kirghisorum is not spread. In all these cases, the species history appears to reflect the character of the area and the species number. The glacial period and subsequent desertification did not weaken the heterozygosity of Malus, which is its main evolutionary base. This base is illustrated by heterogenetic populations of M. sieversii and M. kirghisorum existing in areas of their close contact, where valuable forms and natural hybrids were selected (Dzhangaliev 1968). Human economic activity certainly promoted the formation of mosaic areas. But here, we do not dwell on the species of M. niedzwetzkyana, because its trees are solitary and do not form any significant populations. From the time of the publication of N. I. Vavilov’s theory (1927) about the centers of cultivated plant origin, it became universally recognized that some regions are distinguished by a great variety of habitats, which are especially favorable for the development and preservation of genetic variety. Favorable and variable climatic, soil and biotic environments (Dzhangaliev 1972, 1973c) in the mountains and their distribution in small areas, are reflected in the plurality of apple populations with their intraspecific polymorphisms. Differences between population habitats were distinguished not only by differences in the abiotic and biotic environmental conditions, but also by the intensity of their activity. Some species (M. sieversii, M. kirghisorum) are distinguished by their responses to environmental conditions. The more plastic M. sieversii populates various habitats (dry, wet, humid, or any combinations). The distribution of M. kirghisorum, which can endure only very narrow limits of environmental variation, spreads solely into humid areas. The successful growth of trees in some severe habitats could indicate a possible presence of genotypes with various and wide resistances. Consequently, the separations of populations, which takes place within a species depend mainly on the size and natural conditions of the plot of land occupied by its woody stands. Malus sieversii, which covers a wide area of mountain systems having diverse natural-historical conditions and spreading through many degrees of latitude, is separated by natural selection into genetically different populations which have various levels of polymorphism. In one condition, populations are monomorphic, while in the other, populations preserved all of the characteristics of a polymorphic species. Owing to a common low level of selection in the geographical center of the species in Dzhungarskei and in Zailijskei Alatau at altitudes between 1100–1500 m, and in Tarbagatai, the widest extent of variability

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is preserved. But under the severe conditions of the high mountains and in Tarbagatai, many possible variants of apple populations are excluded, except highly adaptive and highly specialized variants. In the geographical centers, the largest number of available habitat conditions and the largest populations exist. Consequently, there are numerous dense apple populations and a concentration of different mutations. On the other hand, at the periphery of the species areas, a spotty distribution of populations appear within small areas, in small numbers and with low tree densities, which leads to a change in the intrapopulation and interpopulation genetic diversity. These factors, which are different from those in some other parts of the species area, influence the level of selection pressure on them. An analysis of these studies shows that a number of crossed apple trees have different genes that influence intraspecies polymorphism. Apple populations in the middle mountains of Zailijskei and Dzhungarskei Alatau, where M. sieversii and M. kirghisorum grow in a continuous line of contact that extends 80 to 100 km, are characterized by a large variety of characters. This variety of characters is the consequence of allopathic hybridization of these species, which results from a breach in the integrity of geographical isolation barriers between them. Under optimal conditions at an altitude of 1100 to 1500 m, not only isolated specimens, but tens of thousands of trees take part in this developmental process, even in the thickest thickets of individuals per hectare in high-density and pure apple forests. Here, there are favorable conditions for hybridization in nature, including the absence of any geographical isolation, approximately equal rhythm of development of most neighboring growing individuals, and a similar flower structure and physiological compatibility of kindred apple species. All of this promotes their crossing and the development of seed vitality and hybrid progeny. Besides the total number of trees, the age of the individuals exert some influence on the quantity of offspring produced. The system and the complexity of cross breeding change depending on whether one or many different flowering and propagating-age groups of trees take part in natural hybridization. In populations where several age groups of cross-pollinating mature trees take part in the natural hybridization process, the genetic mixture is sharply increased, and this influences the morphological and physiological features of individuals. Apple populations as a genetic and evolutionary unit consist of trees of different ages (Table 2.37). The duration of an individual life and the period of tree flowering during the life cycle are important factors for the well-being of the population. Apple stands are distinguished by their

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213

Age composition of tree populations of wild apple. Distribution (%) Age

Altitude (m) 1101–1300 1301–1500 1501–1800

Up to 10 yr

11–40 yr

41–70 yr

71+ yr

5.8 4.8 50.0

44.5 42.2 16.6

44.9 50.1 33.4

4.8 2.9 –

complex age composition, which includes possible variations of combinations in tree populations descending from different seedings and different age groups. Other factors are the considerable duration of an individual life (up to more than 100 years) and an ability to bloom and to cross from 10 to 70 years of age. Consequently, apple populations consist of individuals of different seedings and of various age groups born from one set of parents. Not taking into consideration the periodicity of “flowering–bearing,” apples are able to pollinate over a considerable territory and for a long period, and thus create tree populations of different seedings and ages. We have characterized the vegetatively propagating apples, which have a longer reproductive period than seed propagated trees. The reproductive period is longer because the formation of autovegetative trees is not related to the phase of cropping periodicity. Both seed and vegetative trees can be pollinated by individuals of any age, which further complicates the picture of natural hybridization. Age composition of populations is determined not only by the long life duration and the periodicity of blooming and of pollination, but by the life duration of the progeny and, later, mortality, which are sharply different in the various age groups. Besides the effect of species, there are other populational differences among apple trees. These differences are seen in relation to the index of the age structure of apple stands, which vary within different areas (Table 2.37). Here, in the process of natural hybridization, young trees (age class I) and old tree (age class V) take part. Added to other population differences are differences in the age of first cropping, depending on vertical zone (Table 2.38) and differences in their reproductive periods (Table 2.39). They are not identical for the different populations. In apple populations, both in different areas and within one area, a harmonious and balanced multiage composition of trees is observed. The apple forests of Kazakhstan do not suffer sharp fluctuations in number

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A. DZHANGALIEV Table 2.38. Distribution of wild apple populations in relation to age of first cropping. Distribution (%)

Altitude (m) 1101–1300 1301–1500 1501–1800

5–7 yr

Beginning of bearing 8–10 yr

10+ yr

– – 60.0

43.6 39.8 40.0

56.4 60.2 –

of trees as do other forests, wherein young individuals are a major component. Young trees are characteristic of populations that are suffering a stage of depression. The age structure of populations reflects a common type of tree, which is particular to the species. At the same time, tree age is a highly unstable characteristic of populations. The age composition of apple trees varies considerably, not only in different populations, but also within a single population during various years. These effects are dependent on a number of features caused by fluctuating environmental conditions. For example, the beginning period and the duration of spring frosts during flowering have a definite influence on the pollination and fertilization of reproductive apple groups, which, according to the time of blooming are different, not only in different populations, but also in one and the same population (Table 2.39). The same could be said about spring precipitation, which, depending on intensity, duration and regional scope, may have a negative influence on pollination and fertilization, both directly and indirectly. The indirect influence of precipitation is due to barriers to numerous flying insects that promote cross-pollination, especially wild bees. Slope exposure, type of relief, and plant community may influence the pollination rate, reproductive features and distribution of different apple populations, either by accelerating or delaying any action that is taking place. There are pure biological and ecological types of genetic isolation of apple populations. This means that trees differ from other individuals by the time they reach their reproductive age, and also there are different reproductive territories. In natural populations, side by side in a plot with equal soil and biotic growing conditions, apple trees can be of different ages and they can be early or late flowering (Table 2.39). Genetic drift occurs during the flowering period. This drift during the blooming period can lead to intrapopulation reproductive isolation, which may be

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partial, but not absolute. Many populations can carry both late flowering variants and intermediate flowering variants. The extension of flower timing of all groups allows them to cross-pollinate with individuals within a population, and also, when in close contact, with individuals of other populations. Mid-season bloomers, which demonstrate population adaptability, are shown in Table 2.39. Superiority in resistance to unfavorable environments among the progeny in nature occurs in heterozygous populations of Malus. This genetic structure of populations, under the influence of natural selection, may change in allele frequencies, promoting a high level of heterozygosity in populations, and consequently, the adaptability of its individuals. In the present case, neither the early flowering trees nor the late flowering trees can be sustained alone in populations. However, alleles for a range of flowering times are preserved and their frequencies are secured, including alleles for intermediate time of flowering. In apple habitats where the beginning of early spring frosts start both in the early period and in the later period (the second ten-day period of April to the beginning of May), apple populations were selected according to their generative adaptability to climatic conditions. Evenly flowering individuals with long flowering periods were preserved. Analyses (Tables 2.39–2.41) confirmed that the same condition in apple populations, which provide steady polymorphism to trees was observed in all of the other characteristics studied. Under fluctuating conditions in the mountain regions, varied forms of polymorphism provided an opportunity for better survival by apple progenies in severe habitats. As N. N. Turbin (1971) correctly wrote that a balanced polymorphism promotes an optimal level of adaptability for populations. As a result of mutual enrichment between populations of M. sieversii and M. kirghisorum, mixed populations appear, among which, in the process of natural selection, the more adaptive trees are empowered with the privilege of further existence. Depending on tree height and fruiting indexes, populations in the zone where the two species come in contact with each other at elevations of 1100 to 1500 m (in more favorable habitats for apple) have features like M. kirghisorum. At elevations higher than 1300 to 1500 m (under more severe conditions), M. sieversii features dominate the populations in the zone of contact (Tables 2.42–2.46 and Figs. 2.35–2.38). Intrapopulation selection from these two species has gone on as long as interpopulation selection has gone on. This is verified by viable individual crossings between both species and by the fact that the biological isolation between their populations are mostly only partial and not absolute. As Timopheev-Resovsky et al. (1973) showed,

216

1101–1300 1301–1500 1501–1800

Altitude (m)

12.9 12.6 40.0

Low (< 20 kg) 24.9 33.2 60.0

Middle (20–40 kg)

Yield distribution (%)

62.2 54.2 –

High (40–80 kg)

May 7 May 14 –

Early

24.7 15.0 –

May 12 May 21 May 24

Middle

42.4 15.0 –

Insufficient hardiness

32.9 41.3 100.0

Hardy

May 15 May 23 –

Late

Winter-hardiness (%)

May 6 May 10 –

Not hardy

Apr. 20 – –

Very early 7 – –

Very early

11 10 8

Middle

6 5 –

Late

38.3 41.3 100.0

Healthy

61.7 27.9 –

Susceptible

Disease susceptibility (%)

9 9 –

Early

Relation between yield, winter-hardiness, and disease susceptibility of wild apple trees and altitude.

May 2 May 8 May 16

Late

Table 2.40.

Apr. 27 May 1 –

Middle

Flower duration (days)

Apr. 23 – –

Early

Flower ending

4:19 PM

1101–1300 1301–1500 1501–1800

Very early

Flower beginning

Relation between flowering times of wild apple and altitude.

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Altitude (m)

Table 2.39.

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Table 2.41. Relation between habitat altitude and the number of trunks per tree and tree heights, 20 to 40 year-old apple trees. Distribution (%) Trunk Altitude (m) 1101–1300 1301–1500 1501–1800

Height

Onestemmed

2–4 stemmed

Bushystemmed

<3 m

3.1– 5.0 m

5.1– 8.0 m

8.1– 10.0 m

55.8 62.6 –

44.2 30.9 40.0

– 6.5 60.0

– 4.1 60.0

37.9 18.7 20.0

47.4 68.3 20.0

14.7 8.9 –

Table 2.42. Distribution of wild apple populations (age 20 to 40 years) according to form and diameter of tree crowns. Distribution (%) Crown form Altitude (m) 1101–1300 1301–1500 1501–1800

Crown diameter

Plane1.1– 3.1– 5.1– 7.1– Pyramidal Oval branchy <1 m 3.0 m 5.0 m 7.0 m 10.0 m 10.0+ m 7.4 – –

59.4 71.5 60.0

33.2 28.5 40.0

– – 20.0

– 2.4 60.0

18.4 23.6 20.0

49.5 55.3 –

25.8 17.9 –

6.3 0.8 –

Table 2.43. Distribution of wild apple populations (age 20 to 40 years) according to the time of fruit ripening. Distribution (%)

Altitude (m) 1101–1300 1301–1500 1501–1800

Table 2.44.

Very early (before Jul. 10) 22.4 – –

Time of ripening Early Middle (Jul. 10–Aug. 19) (Aug. 19–Aug. 31) 33.8 22.6 –

Late (Sept. 6 and later)

32.2 50.9 100.0

11.6 26.5 –

Distribution of wild apple populations according to form and fruit weight. Distribution (%) Weight Form

Altitude (m)

Round

Conic

Cylinder

<10 g

10– 25 g

25– 50 g

50– 75 g

75– 100 g

100+ g

1101–1300 1301–1500 1501–1800

88.2 84.5 100.0

9.9 12.3 –

1.9 3.2 –

7.9 7.3 6.7

67.7 69.4 90.0

21.1 23.3 3.3

1.5 – –

1.1 – –

0.7 – –

217

218 Green

Sweet

Sourishsweet

1101–1300 1301–1500 1501–1800

Altitude (m)

34.4 30.3 100.0

Smooth

46.9 55.2 –

Ribbed Slightly ribbed 18.7 14.5 –

Weakly ribbed 5.2 – –

81.7 75.1 –

Orange 11.8 18.2 –

3.4 5.2 –

40.0 30.8 –

1.3 6.7 100.0

Green

58.3 63.3 100.0

No color

Distribution (%)

2.2 2.4 100.0

Ground color

1.9 3.8 –

Yellow

27.3 29.2 –

White

68.6 64.6 –

4.3 3.5 –

Crimson

1.3 3.2 –

14.2 13.1 –

Red

Sweetishsour

Distribution of wild apple populations according to ribbed fruits and fruit color.

21.1 21.9 100.0

Pink 2.2 6.2 13.2

Sourishbitter

18.1 9.1 –

Pink

0.4 0.3 –

Scarlet

Cover color

45.6 41.8 60.0

Sour

Flavor

1.7 4.8 –

3.0 5.9 –

Orange

4.3 7.2 –

Sweetishbitter

Purplish red

3.2 5.6 26.8

Bitter

4:19 PM

18.5 12.2 –

Cream

Table 2.46.

White

60.4 65.9 –

Firm

1101–1300 1301–1500 1501–1800

Medium

Flesh color

Soft

Flesh consistency

Distribution (%)

Distribution of wild apple populations according to flesh consistency, color and flavor.

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Altitude (m)

Table 2.45.

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a

b

c

d

Fig. 2.35. Characteristics of stem structure of wild apple: (a) one-stemmed; (b) twostemmed; (c) five-stemmed; (d) bushy.

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a

b

c

d

Fig. 2.36. Features of the crown structure of wild apple: (a) pyramidal crown; (b) vaseformed crown; (c) round crown with uneven edges; (d) flat crown.

competition between these two species and populations is also a factor for selection. Thus, intrapopulation and interpopulation crossings by Malus individuals lead to a leveling of differences among them. The degree of these differences depends on the pressures of intrapopulation and interpopulation isolations. These pressures are determined by tree number and depend on when the trees are ready to cross-breed. Their readiness

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Fig. 2.37.

221

Variability of pomological fruit indexes of Malus species in Zailijskei Alatau.

Fig. 2.38. Variation in the fruit features of natural hybrids between Malus sieversii and M. kirghisorum in the species centers of Dzhungarskei Alatau: fruit color, size, form, stem, and cavity. Photo by V. P. Bochko and L. N. Yupatova.

to breed may be influenced by the density of populations and by the course of microevolutionary processes. Independent of the number of trees, the densities of trees cause changes in local populations. When divided by mountain peaks or by thickets of other tree species, it is difficult to cross-pollinate with trees in nearby apple populations. In such isolated populations, as a result of kindred crossing, the possibility of homozygosity increases. Natural selection of apple tree forms may

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become intrapopulational. This is based on the high degree of isolation between the different spatially close populations, in which different genotypes react to the influence of the local surroundings. As a result of frequent transformations of different genotypes toward greater population adaptability to these environments, local adaptations and the formation of populations of trees with specific genetic make-ups will occur. Interspecific crossing of M. sieversii and M. kirghisorum, along with inherent heterogenesis of their populations, both multiform genetic mixtures and unfavorable adaptation-genetic differences between populations, lead to an obligatory interpopulation selection in the zones where there is contact between them. Thus, the natural interactions of populations within an apple species mainly depend on the nature of their territorial closeness and on the degree of isolation pressure between adjacent populations. Genetic isolation of any type, depending on its duration, disturbs the whole lot among or within population groups. This leads to both the formation and blending of any group differences. In the case of a disturbance to the isolation between adjacent populations of M. sieversii and M. kirghisorum, one- or double-sided penetrations of intensively selected genotypes or mutations become possible. As a result of crossing, new genotypes appear, and under the influence of natural selection, they may either disappear or remain standing in the struggle for life. Interpopulation selection favors complex changes in the frequencies of different genotypes in adjacent populations, while intrapopulation selection leads to gradual local changes of correlated adapted genotypes. Because of the evolution of apple features, a large role is played by intrapopulation polymorphism, which promotes species polymorphism and the formation of intraspecies taxa. For artificial selection and for use in cultivation, populations of high-mountain trees with short vegetative periods are of special interest. These are distinguished by early initial cropping and precocity, and also by apple populations of the middle mountains with high pomological fruit quality (Tables 2.43–2.46, Fig. 2.38). All tree forms were selected according to fruit indexes within these mountain populations (Dzhangaliev 1968). Isolation of apple populations can result in mutational differences affecting morphological and physiological factors, such as rate of pollen growth, flower structure, and generative organs. Further investigation may reveal that these differences are due to karyotypic differences, sterility, and other factors. The degree of geographical isolation is defined not only by the spatial distribution of interpopulation barriers but by apple dispersal or “the radius of individual activity” (Timopheev-Resovsky et al. 1973). In spite

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of the seeming isolation of separated apple stands, dispersal to other places are characteristic of tree populations, which sometimes covers a large territory, promoting the mixture of genes. Spread to other places is conditioned by a number of biological features of apple, such as the distances of pollen dispersal and seed scattering, the shading of tree colonies, and by the extent of root attachment. Apple trees, depending on their development cycle and the place they occupy in the population, and also on their developmental class, have different degrees of gene exchange. Thus, mature trees with high crowns and well-developed reproductive organs, formed according to developmental classes I and II and possessing airborne pollen, disperse the species well. Steep slopes cause ripe fruits with seeds to roll down hill a considerable distance. If apple trees have secondary importance in the community or ecotype, then these processes proceed with more difficulty. Pollen will spread 200 m, similar to that of cultivated varieties. The highly nutritional qualities of apple fruit attract wild animals which promotes seed scattering about the mountain territory. Especially in high-yield years, animals choose the most delicious fruits, which is one element of negative selection. Root self-grafting of trees and colony formation also promote contacts between different genotypes and genetic mixing over a large territory. These factors influence the genetic structure of apple populations, which can be illustrated with the following examples. In Zailijskei and Dzhungarskei Alatau, where populations of M. sieversii and M. kirghisorum come in contact, the combined traits of both species are observed. Trees are short, medium or high (height 3–10 m and higher) (Table 2.41, 2.42), fruits are small, medium or large, sour or sweet (Table 2.44, 2.45), leaves are sparse or dense, and variably hairy. Populations consist of trees that are in different stages of transition. For example (Table 2.39), in apple populations at the altitude of 1100 to 1300 m, where bidirectional cross-pollination is possible, there are early flowering (April 23) and late flowering (May 7) trees. Between these extreme variants, there are numerous transitional trees. The superiority of heterozygosity over homozygosity (flowering simultaneously), in regard to resistance progenies, reproduction offers populations a great selectional advantage. Natural selection picks out heterozygous individuals as more resistant. Variability in the gene pool is large. Periodic changes in environment conditions (mainly the climate), prevent any form from being consistently superior. The drift in the population composition, which takes place during the processes of evolution and during periodical changes in environments, leads to a change in the distribution of heterozygotes and new forms. In these populations, as a result of mutual exchange of genetic

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material, the “purity” of apple species features was wiped out. Thus, the parent species lose their identities because of gradual transitions of characters in their hybrids. This obliterates any interspecies taxonomic subdivision. Introgressive hybridization comes into play, because in apple stand populations, natural hybrids are repeatedly crossed with each of the parents and between themselves. This constantly repeated process leads to the masking of their hybrid nature. Characteristics pass from one species to another, and offspring seem not to be the hybrids, but a very changeable Malus genus. Thus, interpopulation selection, especially in the zone where there is close contact between different populations of Malus species, promotes the formation of new genotypes and this leads to the creation of new directions for selection in a wide hybrid zone. Apparently, interpopulational hybridization created in nature some valuable apple cultivars that were then selected by us (selections 28, 43, 269, and others) (Dzhangaliev 1968). Although they grow in the same mountain region, local apple populations within the high-mountain belt are considerably distinctive in their make-up and differ from populations from the middle-mountains. The high-mountain apple stands of Dzhungarskei and Zailijskei Alatau are very similar to Tarbagatai stands. Natural selection in local populations leads to the stabilization of a number of characteristics as relatively uniform forms. Thus, on a slope in the Soldatskoe gorge at the altitude of 1700 to 1800 m, it is possible to see high-mountain apple populations. They grow on stone soils under severe conditions in the upper part of a watershed plateau. An investigation of the features of apple stands showed that in highmountain habitats the apple populations sharply differ from the middlemountain ones. A comparative analysis revealed that the high-mountain populations are formed from low-growing, bushy individuals, comprising 60 percent of the population and the rest, 40 percent, from low- and middle-growing trees, while single-stemmed trees are absent here (Table 2.41). They bloom in mid-season (Table 2.39), fruits ripen relatively early (Table 2.43), and according to these characteristics, they are not distinguished by as much polymorphism as are the individuals of the middle-mountain populations. The most important population feature is the early-ripening of fruits and seeds. In spite of the short vegetative period, fruits will ripen in August. Precocity of trees is also important (Table 2.38). Most of the trees (60%) began to bear at the age of five to seven years. The trees in the middle-mountain populations are considerably delayed in bearing, but they are more long-lived (Table 2.37). High-mountain trees are shorter-lived, more uniform in fruit

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weight and form (Table 2.44), more uniform in ground and cover color and ribbed fruits (Table 2.46), more uniform in the flavor and texture of the flesh (Table 2.45), more uniform in the structure of fruit carpel and the number of seeds (Table 2.47), in the structure of fruit cavity and calyx (Table 2.48), and in the yield (Table 2.40). They are sharply distinguished by their structural features from the high-mountain populations. Monomorphism of the high-mountain populations occurs not only in tree habit, but also in fruit characteristics. Only small and sour apples that are mainly green are seen. The high-mountain populations appeared to be frost-resistant (Table 2.40) and they were not affected by apple scab (Venturia inaequalis) or apple powdery mildew (Podosphaera leucotricha) (Table 2.40). Here, the trees are not vigorous, but are of bushy form and average yield. Populations of apple are scattered in forest thickets that are connected by narrow, local complexes with certain nutrient and soil environments. In these populations, the character of stand structures differs as forestvegetative conditions change. For example, when moving up a slope (1500 m and higher) or downwards (lower than 850–900 m), a deterioration of the ecological conditions for apple habitat occurs and the high density and pure apple woods are no longer seen. Instead, apple forms only thin forests and stands occupy severe habitats. They, as a rule, are not forest builders but fall into class III or a lower developmental class. Apple grows on the steppe part of the rubble of high-mountain plots or on dark-chestnut soils in the premountains, among mixtures of wormwood. These dense forest centers also include apple populations (Timopheev-Resovsky et al. 1973). Apple populations, as biological systems, possess ability for selfregulation of such vitally important processes as natural regeneration and cross-pollination. However, not every isolated apple group may be Table 2.47. fruits.

Population groups of wild apple according to core form and fruit carpel of

Distribution (%) Core

Fruit carpel

No. seeds

Altitude (m)

Bulb

Turniplike

Cordate

Closed

Semi closed

Open

1–6

7–8

10+

1101–1300 1301–1500 1501–1800

53.1 49.3 90.0

32.7 40.2 6.6

14.2 10.5 3.4

43.8 41.0 56.7

31.4 38.1 36.6

24.8 20.9 6.7

15.4 20.1 –

57.9 53.9 66.7

26.7 26.0 33.3

226

Deep

47.5 54.5 –

1101–1300 1301–1500 1501–1800

36.6 34.6 26.7

Middle

Depth

15.9 10.9 73.3

Shallow 42.4 49.4 –

Wide

Fruit cavity

22.8 13.9 40.0

Middle

Width

34.8 36.7 60.0

Narrow 6.9 7.5 6.8

Russet

93.1 92.5 93.2

No russet

43.9 43.2 –

Open

14.6 17.4 33.3

Semiopen

Calyx

41.5 39.4 66.7

Closed

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Distribution (%)

Population groups of wild apples according to structural features of fruit cavity and calyx.

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Altitude (m)

Table 2.48.

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considered to be a population. For example, short duration single specimens or groups of a few apple trees, in developmental classes III or IV, accidentally sprung up in a thick mixed forest on the steppe premountains. Growing at a considerable distance from other apple stands, these trees are able to exist without interruption for one or two generations. They have no perspective for further evolutionary development and, apparently, represent only residues of larger populations, which grew here in former times. This does not refer to the periphery of a species’ area, where the sizes of apple populations and the number of trees are small. The peripheral areas have increasing isolation pressure among populations that apparently, creates preconditions for an accidental separation and fixation of recessive mutations and polyploids, important for the initial formation processes of species. Besides that, on the periphery of the species’ area, abiotic and biotic conditions surround apple populations that favor the development of trees resistant to these environmental factors, which are at a minimum here. Based on N. I. Vavilov’s theory (1927) about the special character of hereditary variability at the periphery populations, the processes of initial apple development occurred at the most northeastern part of the mountain systems of Central Asia and Kazakhstan (in Tarbagatai, Dzhungarskei, and Zailijskei Alatau). In Zailijskei Alatau (Popov 1927; Vavilov 1931), there exists a great geographical intraspecific variability that is characteristic of Malus. Wide areas of M. sieversii include not only the mountain regions of Kazakhstan, but other ridges of Tien Shan, and also the Pamirs-Alai with their multiple natural conditions. The area spreads through many degrees of latitude and is separated by natural selection into a great number of genetically different populations. Conversely, this species, while necessarily growing within narrow, limited, and severe habitats remains monomorphic. Consequently, adaptive polymorphism within the species may change from one end of the area to the other. Or changes may be limited without developing into new forms and remain as independent populations. All populations of M. sieversii were distinguished by such essential adaptive characters as growth rate, frost-resistance, and early-ripening. These features were associated with each other and with tree height. In humid and wet forests in the middle mountains (Dzhangaliev 1974a), vigorous high-growing trees arise during long periods of vegetation and this distinguishes them by their vitality and freezing susceptibility in winter. The very dry forests of the high mountains and the dry-steppe belt of the low mountains are inhabited by populations of low-growing individuals with bushy forms. These populations are characterized by a long winter dormancy and a high frost-resistance.

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An analysis of the temperature requirements both for passing through the separate phases of apple development (flowering, setting of reproductive organs, maturing of fruits and seeds, autumn leaf coloring) and for the completion of the full developmental cycle was conducted (Dzhangaliev 1972). This investigation showed that in the highmountain populations, only early-maturing, vegetative short-period forms of trees grow, which characterize them as frost-resistant. In such isolated populations of M. sieversii there were no interesting forms with good fruit quality. We did not select a single tree for its good fruits. But good fruits were relatively unimportant, since these trees possessed frost resistance. In the high-mountain populations, all fruits are green, small, sour, and so on (Tables 2.45 and 2.46). The founder of population genetics, S. S. Chetverikov (1926, p. 12) concluded: “Species, like a sponge, absorb heterozygous gene variations, but remain phenotypically homogeneous.” M. E. Lobashov (1967) pointed out that individuals, which are genotypically distinguished, for example, according to one gene, may not differ morphologically, but may possess a different vitality, a different productivity or some other difference. Evolution resulted in early-maturity of fruits and seeds and also, precocity of apple trees. Under cold or other severe conditions in the high-mountains, tree populations with a late period of fruit ripening were selected against because they did not have any progeny. During artificial hybridization of wild maternal forms, 75 percent of the hybrids had precocious indexes, and this also characterizes the high adaptability of wild apple to the severe mountain conditions. Indexes of apple frost-resistance (early-ripening and precocity) are probably a result of physiological reactions, and are less dependent on selection, or other indexes. The variability of M. sieversii individuals, forming numerous populations in the most different ecological conditions, may be explained by the plasticity of its genotypes in their response to the environment. In addition, in populations formed in extreme conditions, natural selection cannot reduce their heterozygous individuals to some homozygous form. It is possible that local highmountain apple populations reflect a strong resistance of M. sieversii genotypes to intensive natural selection. Developed M. sieversii populations, along with their habitats, emerge as a complex natural union. In M. sieversii populations, a more advanced progressive species in comparison to M. kirghisorum, indexes and characters are easily correlated with different external conditions of the habitats. As the result of these correlations, functions of individuals become similar. Adaptation includes immunities, which can protect individuals from the influence of harmful diseases under these new

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conditions. But as a whole, the main task of the adaptation complex is to prevent a conflict between species populations and their environments, to help the populations to adapt themselves to external conditions of the habitats, to preserve the constancy of internal conditions of individuals. All of this is homeostasis, a state of physiological equilibrium when stresses are reduced. The ability of M. sieversii populations to adapt is characteristic of their individual members with their complex of indexes and features, separate organs and also the population itself as a whole. The scale of population adaptation depends on the conditions to which it needs to adapt at any given moment. An example is the formation of air roots of trees during a flood and erosion and the subsequent change of habitat structure, depending on the conditions at the place of growing. Such counter measures help populations to endure unfavorable conditions in the habitat. Consequently, in each specific case, an optimal variant of population existence is put forward. Thus, there are many adaptation mechanisms of Malus, with strong reaction intensities and large scales. Adaptation, as a process of population adaptability to climatic and geographical conditions in broken areas and habitats, takes place for any progeny of individuals that enter into the plant community. During orthogenesis or during several generations of phytogenesis, constant reconstruction is taking place, including self-correction of population functions as a whole and of all individuals and their organs. Before that, they had acquired the ability to live under these new conditions. Thus, the action pattern of adaptive features and indexes of adaptation are rather wide for M. sieversii populations. This pattern varies within definite limits and directions in each population, which is especially clearly demonstrated in interspecies crossing. These populations are well adapted to new and unusual conditions and even to extreme conditions. Populations differ among themselves, just as they differ for fruit morphology, morphological characters of trees, duration of vegetative period, precocity and so on. What do the reserves of population adaptability depend on? Actually, they are determined by their genetic traits. But the adaptation reserves are not something that are predetermined, once and for all or programmed when individuals appear. To a great extent, they are conditioned by the species history, by features of the formation of other community species and by the ecological environment, in which species populations and its ancestors existed and exist to the present day. During the life span of the population and for its individuals in phytogenesis and orthogenesis, a definite dynamic stereotype forms. A regularly repeating irritant can cause a responsive reaction, which becomes

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automatic. A complex of such consolidated repeated reactions is the dynamic stereotype. Like a mirror, it reflects the adaptive possibilities of the population. For example, in populations of M. kirghisorum, growing under mesophilous conditions or in the surroundings of heat-loving plants, the whole system is adapted to relatively optimal conditions, but their mechanisms of adaptation are not sufficiently flexible. This explains why populations of M. kirghisorum do not exist higher than 1100 to 1300 m in Dzhungarskei and Zailijskei Alatau, while its natural hybrids with M. sieversii do exist higher than 1400 to 1500 m. In such populations, “pampered” by optimal conditions of the middlemountains, adaptation to changing influences of the environments occurs with difficulty. A chilling action of the environments is of great importance for the adaptation mechanisms. Systematic cooling at high altitudes in extreme habitats has an effect on the populations over long periods of time, and after some generations, promotes adaptation to frost. An analogous conclusion about the role of chilling on the adaptation mechanisms can also be made with respect to the drought resistance of individuals growing at high altitudes on rubble soils of the southern slopes or under very dry conditions of Tarbagatai steppe, where water and temperature are at a minimum. Thus, a required precondition for any geographical site is some degree of spatial isolation located between separate parts of an apple species. This provides for their genetic isolation and thereby promotes an acceleration of the formative processes. Factors affecting the development in the mountains, where sharp contrasts in ecological conditions of apple reaches its maximum over a large territory, are fluctuations in light, the hydrothermal regimes in different habitats, and variations of chemicalphysical, biological, and soil properties relative to the vertical zones. Horizontal and vertical mosaic layers also affect the distribution of ecobiomorphologic components of apple forests. Ionizing and long-wave radiation are factors that influence spontaneous mutagenesis in the highmountain regions (Dubinin 1976). An uneven distribution of environmental factors in time and space greatly influences the processes of genetic alteration in apple populations, either accelerating or retarding their rate. The cause of this influence (Timopheev-Resovsky et al. 1973) is in an uninterrupted genetic mixing inside the population. The mixing of genes prevents any fixation onto any specific genetic form. Examples, which confirm a high level of independence of tree organogenesis of wild apple from external factors, follow. Highly polymorphic species, such as M. sieversii and M. kirghisorum, are represented by numerous individuals that sharply differ from each

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other in their biological and economically important characteristics. These attributes can be used for introduction into cultivation and for research on selections. To illustrate this, we described some selected forms of wild apple. Selection 28 (interspecific hybrid) was selected from apple populations in Dzhungarskei Alatau in 1962. It was first described by A. D. Dzhangaliev and N. K. Volkova in 1968 (Dzhangaliev 1968). Trees are well-developed: height is 6.5 m; trunk circumference is 82 cm; crown diameter is 7.0 m. The apple is frost-resistant, resistant to winter withering, powdery mildew, scab, and apple moth. Its fruits are less damaged by codling moth than cultivated apples. The time of fruit ripening is autumn–winter (September 25). Fruits are of conic-ribbed form and are extraordinarily attractive, exceeding ‘Aport’ in fruit beauty. The fruit are large; the average weight is 239 g, the height is 87 mm, the diameter is 90 mm (Fig. 2.39). The pedicel is of medium thickness, short, strongly hairy, and slightly curved. The fruit cavity is wide and deep, rust is thin and light. The basin is deep with large margins, wide, and large-ribbed. The calyx is of medium size and semi-open. Sepals are long, conic, and green. The medium tough skin is soft, glossy, and with slightly waxy bloom. The ground color is pink; the cover color is red with a crimson tint, and flecked. Fruit dots are large, washed out, whitish, few in number. The core is bulb-shaped and well-outlined. The carpels are open and

Fig. 2.39.

Fruits of selection 28 (natural size). Photo by V. P. Bochko and L. N. Yupatova.

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large. Seeds are brown-red with a pinkish tint and plump. The flesh is pink, and on the margins is red to crimson, hard, and juicy. The fruit flavor is sourish-sweet with a pleasant aroma. The yield from a 90-year-old tree is 300 kg. Fruits are highly transportable and they are notable for their long keeping (6–7 months). The tree form is of interest for ornamental horticulture and selection. Selection 35 (M. kirghisorum) was selected from fruit forests in Dzhungarskei Alatau in 1948 (Fig. 2.40). This form was first described by A. D. Dzhangaliev and V. I. Komarova, and later, in 1968 by A. D. Dzhangaliev and N. K. Volkova (Dzhangaliev 1968). Trees are vigorous: tree height at 27 years of age is 5.5 m; trunk diameter is 30 cm; crown diameter is 6.5 m. Trees are resistant to frosts, scab, and powdery mildew. Time of fruit ripening is autumn–winter (September 20). Fruits are round, medium sized; the average weight is 140 g, the height is 63 mm, and the diameter is 53 mm. The pedicel is slender and of an average length, moderately hairy, and slightly curved. The fruit cavity is deep, narrow, and weakly-russeted. The basin is deep, of medium width, and furrowed. The calyx is medium sized and semi-open. Sepals are long, of narrow-conic form, and green. The skin is tough, thick, smooth, opaque, with thick whitish and waxy bloom. The ground color is strawyellow, and cover color is red with dark-red wide stripes. The fruit dots are numerous, inconspicuous, and whitish-greenish. The core is oval and well-outlined; fruit carpels are open and large; seeds are brown, large, and plump. The flesh is white with a carmine tint, medium firm,

Fig. 2.40.

Fruits of selection 35. Photo by V. P. Bochko and L. N. Yupatova.

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medium-grained, medium-juicy. The fruit flavor is sour-sweet and the aroma is pleasant. Fruit bearing is good: the average yield in some years was 103 kg per tree. Good transportability and long storage distinguish fruit. Selection 35 is promising as an ornamental, commercial processing and breeding. Selection 43 (M. kirghisorum) was selected from fruit forests in Dzhungarskei Alatau in 1948. This apple was first described by A. D. Dzhangaliev and N. K. Volkova, and later by E. N. Nikitenko (Dzhangaliev 1968). Trees are of average growth and development: height is 5 m; trunk circumference is 93 cm; crown diameter is 6 m. Trees are resistant to winter frosts, but they are susceptible to powdery mildew. The time of fruit ripening is autumn–winter (September 20). Fruits are ovate and large with an average weight of 180 g. Fruit height is 74 mm and its diameter is 75 mm. The pedicel is thin, short, densely hairy, and slightly curved. The fruit cavity is medium deep, narrow, and slightly russeted. The basin is shallow, narrow, and wrinkled. The calyx is large and closed. Sepals are long, triangular, and green. The skin is tough, smooth, dull, and greasy with a heavy waxy bloom of bluish-violet color. The ground color is pale-green, but during ripening, it turns yellow. The cover color is dark-red, dull, and striped. Fruit dots are absent. The core is bulb-shaped and partially outlined. The fruit carpels are semi-closed and of medium size. Seeds are large, dark-brown, and plump. The flesh is greenish, medium firm, tender, and juicy. The fruit flavor is sour-sweet and rather pleasant. Fruit-bearing is good, with an average long-term yield of 111 kg per tree. The fruits are of medium transportability and good keeping (until February). This form No. 43 is useful for all purposes. The main trend of its fruit use is for the production of juices, jam, and for fresh use. Selection 238 (M. sieversii) was selected from fruit forests in Zailijskei Alatau in 1962 (Dzhangaliev 1968). Trees are of medium stature: height is 5 m; trunk diameter is 74 cm; crown diameter is 5 m. Trees are highly frost-resistant and resistant to winter withering, scab, and powdery mildew. The time of fruit ripening is summer–autumn. Fruits have a flatround shape, medium-sized, with an average weight of 100 g. The fruit height is 57 mm and the diameter is 73 mm. The fruit stem is mediumlong, thick, and slightly curved. The fruit cavity is wide and of medium depth. The basin is deep and wide. The calyx is open. Sepals are medium-sized, triangular, and green. The skin is tough, smooth, has a bluish waxy bloom. The ground color is pale-yellow and the cover color is dark-cherry, striped. Fruit dots are absent. The core is bulb-shaped, and partially outlined. Fruit carpels are semi-open, and medium sized. Seeds are brown, medium, and plump. The flesh is cream-colored and

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sometimes near the skin it is pink and of medium-grained consistency. The flavor is insipid-sweet, pleasant, with a honey taste and aroma. Fruit-bearing is regular. The average long-term yield is 76 kg per tree. The transportability of fruits is good, and fruit can be stored for two to three months. Selection 238 is of interest as an ornamental, commercial fruit processing and the fresh use of fruits. Selection 269 (M. sieversii) was selected from fruit forests in Dzhungarskei Alatau in 1948. It was first described by A. D. Dzhangaliev and N. K. Volkova, and later by E. N. Nikitenko (Dzhangaliev 1968). Trees are of medium size and development: height is 5 m; trunk circumference is 79 cm; crown diameter is 6 m. Trees are frost-resistant, and resistant to scab, and powdery mildew. The time of fruit ripening is late summer (August 25). Fruits are flat-round, of medium size, with an average weight of 80 g. The fruit height is 55 mm and the diameter is 67 mm (Fig. 2.41). The stem is medium-long and thin. The fruit cavity is of medium depth and width, and weakly russeted. The basin is shallow, wide, and wrinkled. The calyx is large and open. Sepals are large, triangular, and hairy. The skin is medium-thick, tender, smooth, glossy and semi-greasy with a waxy whitish-colored bloom. The ground color is waxy-cream and the cover color is pink with stripes. The core is bulb-shaped, and

Fig. 2.41.

Fruits of selection 269. Photo by V. P. Bochko and L. N. Yupatova.

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partially outlined. Fruit carpels are open and of medium size. Seeds are large, light-brown and plump. The flesh is cream-colored, firm, finegrained, very juicy, tender, aromatic, and with an excellent wine-sweet flavor. Fruit-bearing of trees is good with an average yield for some years of 91 kg per tree. Fruits are transportable and they will store until March. Form No. 269 is very promising for research on the breeding of highly winter-resistant and immune varieties. Fruits are suitable for fresh use and commercial processing. Selection 1048 (M. sieversii) was selected in Tarbagatai in 1972. This description is written here for the first time. The tree is single-stemmed with indistinct layers of branches. The height is 5 m, the trunk circumference is 61 cm, and the crown width is 5 m. The tree is frost-resistant, and resistant to scab and powdery mildew. The crown is spreading flat plane. The bark is gray and scaly. Shoots are short and thick. Buds are of medium size and appressed. The predominant types of fruit shape are round and oblate. Fruit bearing is concentrated on 3–6 year-old wood. Leaves are of medium size, long-ovate, dark-green, wrinkled, and slightly-hairy. The leaf margin is finely-serrate. Flowers are whitishpink, and of medium size. Fruits are round, small, with an average weight of 45 g. The fruit height is 43 mm and the diameter is 48 mm. The stem is long and thin. The fruit cavity is narrow, deep, and slightly russeted. The basin is medium-sized, not deep, and slightly ribbed. The calyx is medium-sized and semi-closed. Sepals are medium-sized, triangular, and green. The skin is tough, soft, with a little waxy bloom. The ground color is green, and the cover color is red with dark-red wide stripes. Fruit dots are numerous and whitish. The core is oval, and partially outlined. Seeds are small and brown. The flesh is white, firm, finegrained, and juicy. Fruit flavor is sour. Fruit bearing is satisfactory, with an average yield for some years of 54 kg per tree. The time of fruit ripening is autumn. Fruits are transportable. Form No. 1048 is recommended for afforestation and breeding research. Fruits can be used for commercial processing. Selection 1051 (M. sieversii) was selected in Karatau in 1972. This description is given here for the first time. Trees are single-stemmed, with good layers of branches. The height is 7 m, the trunk circumference is 61 cm, and the crown diameter is 6 m. Trees are winter- and droughtresistant, and resistant to scab and powdery mildew. The crown is round-globular in shape. The bark on the trunk and branches is scaly and gray in color. Shoots are thick, round in cross-section. Buds are mediumsized, appressed, and hairy. The predominant types of fruit-shape are round and oblique. Fruit-bearing emerges from three-to-seven-year-old wood. Leaves are of medium size, ovate, dark-green, wrinkled, and

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hairy. Leaf margins are double-serrated and stipules are narrow. Flowers are whitish-pink and medium size. Fruits are round, of medium size, with an average weight of 57 g. Fruit height is 49 mm, and the diameter is 53 mm. The fruit stem is of medium length and thickness, and is medium-hairy. The fruit cavity is wide, deep, and slightly rusty. The basin is deep, wide, and slightly ribbed. The calyx is of mediumsize and closed. Sepals are short, triangular, and hairy. The skin is tough, smooth, oily, and with a slightly waxy bloom. The ground color is light-yellow, and the cover color is absent or with a light amber blushing. Fruit dots are whitish, but scanty. The core is of medium size, and bulb-shaped. The fruit carpels are semi-open. Seeds are large, lightbrown, and plump. The flesh is white, firm, juicy, and grained. Fruit flavor is sweet with a pleasant wine taste and aroma. Fruit-bearing is good, with an average yield for several years of 69 kg per tree. The time of fruitharvest is summer–autumn (August 20). Fruits are transportable. Selection 1051 is promising for afforestation, for ornamental horticulture, for breeding, and for the fresh use of fruits. Selection 1054 (M. sieversii) was selected in Talasskei Alatau in 1973. This description is given here for the first time. The tree is singlestemmed, the branch layers are weakly expressed, the height is 6 m, the trunk circumference is 71 cm, and the crown width is 5.5 m. Trees are frost- and drought-resistant; they are not damaged by powdery mildew, and are resistant to scab. The crown is widely spreading. The trunk bark is dark-gray and scaly. Shoots are thick, long, and dark-brown. The buds are large, oval, and slightly hairy. The types of fruit shapes are oblate and oblique. Most of the fruit are born on three-to-six-year-old wood. Leaves are of medium size, ovate, wrinkled, green, and on young trees, they have a pink tint. The leaf margin is fine-crenate, and stipules are narrow. Flowers are medium in size, and pink in color. Fruits are oblong (almost cylindrical); the average weight is 64 g, the height 58 mm, and the diameter is 47 mm. The fruit stem is long, thin, and slightly hairy. The cavity is narrow, deep, and slightly russeted. The basin is of medium size, deep, and ribbed. The calyx is of medium size and is semi-closed. Sepals are of medium size, conic, and slightly hairy. The skin is tough, soft, and with a heavy waxy bloom. The ground color is pink, and the cover color is crimson and striped. Fruit dots are scanty and inconspicuous. The core is oval fairly well outlined, and carpels are closed. Seeds are medium-sized, dark-brown, and plump. The flesh is pink, firm, grainy, and juicy. The fruit flavor is sour-sweet. Productivity is good with an average yield is 76 kg per tree. The time of ripening is autumn (September 1). Fruits are transportable. Selection 1054 is very promising for

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ornamental horticulture and selection. Fruits are good raw-material for commercial processing. In summary, the study of Malus polymorphism in Kazakhstan shows a role for wild apple cultivation and breeding. First, a great deal is obtained through the direct use of intraspecific varieties, existing in nature. This seemingly primitive procedure of selection can be used for the initial introduction of new apple forms into cultivation, of which we managed to single out a number of valuable forms. They can be used on a large scale in forestry, in ornamental plantings, in the processing industry and horticulture. The necessity of making selections relates not only to those indexes and characteristics that we studied, but to those concealed recessive features, which can reveal themselves during their reproduction in cultivation. Recessive characters may turn out to be those genotypes, which appeared in cultivation as a result of a mutational process, but because of being incongruous with the habitat conditions, were eliminated by natural selection. They can reveal themselves in culture as a result of less severe environmental conditions. Tree forms of apple can distinguish themselves by their requirements for ecological conditions, by their duration of seasonal development, winter-hardiness, resistance to diseases, productivity, common physical characteristics and also by their pomological, chemical, and morphological fruit traits. The application of this great intraspecific variety may become an effective method of increasing productivity, qualitative composition, and resistances in natural forests. These valuable apple forms may also be effectively used in introduction work for increasing the forms of cultivated plants. Introduction and reintroduction of these wild apples may be especially successful in combination with research on selections. This could be done by means of direct selection of highly productive and vitally resistant stands in natural apple woods and from them, some trees could be selected for producing seeds and as planting material (Dzhangaliev 1976a). Until now, in establishing forest-fruit nurseries and seed sowing at permanent sites by forest-cultural and gardening establishment projects, attention has not been paid to the quality of the maternal plantations and trees. Seeds have been collected in the most easily accessible apple woods, severely disrupted by long human activity. As a result of the century-old selection by people of the most valuable wood for economic needs in these few afforested regions, the hereditary variety of apple woods has been greatly impoverished. Humans have cut all but the worthless and weak apple trees that have unsatisfactory crown forms.

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These fellings, as a factor of negative selection, have led to apple-wood degeneration, and the retention of less-fertile and faulty trees, especially in lower borders of the forest belt. This systematic felling of the trees deteriorated the succeeding forest generations, because they were removed from the process of natural hybridization, and any renewal continued from the remaining worse forms. A no less negative influence on the development of these apple woods and a worsening of their habitat conditions has been the century-old systematic damage by cattle, hay-making, and plowing of the land for the sowing of annual crops. An even more negative influence on the preservation of this forest gene pool was the mass regrafting of wild apple trees with cultivated varieties to create orchards in the mountains. The healthiest and the best trees were grafted, and after grafting they were no longer able to take part in cross-pollination and natural renewal; their gene pool was lost forever. The apple-wood degeneration on these plots is visibly evident today, trees are leafless and poor yielding. Thus, the degeneration of the forest is the result of a reduction in the number of the best and an increasing of the worst trees, and the deterioration of woodstand habitat conditions. In the remote unpopulated regions and nearer to the upper borders of the applewood areas, and in association with less human influence, woodstand degeneration is almost never seen. Here, more of the best trees have been preserved. Because apple gene pool preservation is dependent on the hereditary characters of maternal trees, the preparation of seeds and planting material from the worst trees is a great tragedy. It is equivalent to the propagation of the less valuable apple stands. Our work led to the use of the best stands and individual apple trees for the propagation of succeeding seed and vegetative generations. As a result of expeditionary investigations relative to the forest tree classifications (Lindquist 1958; Pravdin 1963; Nekrasov 1973), three apple wood groups were categorized for propagation: plus, normal, and minus. The plus apple group includes long-lived highly productive trees with wide spreading crowns, well-developed skeletal branches, that regularly and annually bear fruit on twigs that are 12 to 15 years old. The preparation of seeds and planting material of only the best plants is necessary from this apple wood group. The normal apple wood group includes medium yielding trees with strongly developed crowns, that bear fruit on branches 8 to 10 years old, and that bear fruit no less than one to two years after a year of bad harvest. Here, high-yielding and longlived trees constitute no less than 15 to 20 percent of the total. These apple woods are the main base for the mass preparation of seeds and plant material for propagation.

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The minus apple wood group mainly consists of narrow-crowned, leafless trees with badly developed skeletal higher branches, bearing only a few fruit on twigs 2 to 3 years old, and producing fruit only every two to tree years. In such stands, which are mainly situated near populated regions, harvesting seeds and plant material for future propagation is absolutely unacceptable. In plus and normal apple wood groups, the best trees were selected, especially those that distinguished themselves by productivity, ornamental value, and resistance to unfavorable environments, including pests and diseases. The importance of individual selection was noted by B. Lindquist (1958): “The discovery of one natural elite tree may be as important as all the long-term work on selection” (p. 15). As a result of these investigations, 15 forest-seed plots were selected, and these may fully provide forest-fruit nurseries of southeastern Kazakhstan with apple seeds (Dzhangaliev 1976a). Taking into consideration the inadvisable transport of seeds from the regions of their origin to other natural geographical conditions, we weighed their adaptability to corresponding natural zones of the mountain regions during apple wood selection (Fig. 2.42). However, at present, seed apple woods are used very rarely. The rules of seed picking in the best stands and from the best trees are badly kept. Grafting plantations of pole trees does not exist. Seed sections of nurseries, as well as sowing forest crops in an established place, are created without taking into account the hereditary and the seed features of mother trees. Instead, they are mainly from the seeds of random pickings. In order to increase the fruit bearing of seed apple plots, thinning of stands and training of apple tree crowns by severe cutting of branches are practiced. Removing forest trees, bushes, and fully mature apple trees from the plot causes a disturbance of the stable conditions in a thicket community and damage to natural vegetative renewal. The training of free-growing wild trees, depending on the type of cultivation, involves pruning, which often results in severe removal of six to eight year increments from the upper crown. The upper bearing crown part is removed, which decreases not only the fruit yield, but also the class of tree development and the tree’s competitive ability in the forest, in particular in the struggle for upper light. Until now, there has been a strong opinion that wild apple is homogeneous in its composition and thus, it is simply called a wildling. These materials show that the forms of Malus species sharply differ among themselves in their characteristics and peculiarities as do cultivated varieties of the hybrid species, M. domestica. These differences include the economically important features of the different forms.

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Fig. 2.42. Seed plot locations of wild apple: 1 to 15 = numbers indicate seed plots (seed genetic reservations of wild apples, selected by the author in all mountain systems of Kazakhstan).

Apple polymorphism may be caused by many different hereditary and nonhereditary forces, creating populations with different genotypic and phenotypic characteristics. Sometimes intraspecific taxa (forms) essentially differ among themselves in the complex of qualitative morphological and pomological characters. In other cases, the difference is in a deviation from the norm by one or two characters. By using the methods of field and expeditionary investigations, it is rather difficult to discover if the differences found in individuals are a reflection of genotypic changes or if they are a result of the influence of abiotic and biotic environments (climate, soil, vegetative communities). If there is a population of apple trees of different aged individuals, but with equal edaphic and biotic conditions, then a difference in the effect of a single character would be difficult to determine. For example, trees growing in the neighborhood might be damaged or undamaged by scab or powdery mildew.

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Or, trees might have early or late vegetation, or large or small fruits, and so on. These examples corroborate the existence of different qualities of trees in a single apple population, and it is probably safe to assume that these differences in quality are genotypic. We tried to elucidate the very important theoretical and practical question whether wild form variabilities are genetically or environmental controlled. We studied phenotypic differences of vegetatively propagated progeny under other various conditions. We also conducted a hybrid analysis of a number of hybrid families of some species and forms of wild apple by crossing them with cultivated varieties. For this purpose, in 1946 to 1968 (with the participation of N. K. Volkova) at the altitude of 1350 m above sea level, an experimental orchard was established, in which 39 selections of wild apple were planted on 1.15 ha. Then, in 1970 to 1973, we planted other selections at an altitude of 900 m (Dzhangaliev 1974a, 1976b). The selected forms were grafted onto the rootstocks of wild apple seedlings and were planted in their permanent locations in the orchard. During vegetative propagation, the hereditary features of mother plants readily transmit to new clonal generations. Being the result of short-term phenotypic variability, characters, which appear as a result of the influence of the environment on the organism, are not preserved. Clones planted in the orchard are equally provided with nutrient substances by the roots of their rootstocks, and this allows them to react equally in alien conditions at this new location, and thus, to more rapidly adapt and reveal their hereditary peculiarities. Thus, the use of clonal material in the experimental studies removed the influence of genetic recombination and selection so that objective data could be obtained on the behavior of separate genotypes (forms) under cultivation. Selections 198 and 270 are M. kirghisorum, the rest are M. sieversii. All selections in cultivation preserved inherent individual hereditary features, and almost all do not suffer from winter drought. Selections 28, 177, 197, 238, and 270 carry immunities to scab and apple powdery mildew. Fruits of early ripening selections 132, 134, and 197, as in nature, ripen by the end of August; and late-maturing selections 28, 35, and 36, in the middle of September. Average long-term data show that the yields of wild selections were 52.0 to 121.6 kg per tree, but yields were considerably less with 39.5 to 42.4 kg, for cultivated varieties. Selections 28, 177, and 198 carried genes for large fruits under cultivation with weights of 181 to 200 g, while ‘Aport’ (syn. ‘Alexander’) fruits weighed 215 g. Small-fruited selections remained small under cultivation. The investigations showed that color, form, flavor, storage life, transportability of fruits, and winter-hardiness of trees are also preserved in

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cultivation. The vegetation period of wild selections is five to seven days shorter than in cultivated varieties, which is explained by their ecological adaptability to mountain conditions. The mother trees of selections 28, 35, 177, 197, 198, 123, 260, 269, and 271 during the severe winters of 1950 and 1951 with temperatures falling to minus 33.5°C, and similar low temperatures in the following years, were absolutely not injured, while some damage did occur on trees of the cultivated variety ‘Aport’. These selections of wild apple continue to exhibit their peculiarities and characters in culture, and people show great interest in propagating them to establish resistant and productive orchards for different uses. Studying variability of wild apple features under cultivated conditions showed that radical changes in structure and function of selections do not take place. This high level of apple evolutionary independence from external factors testifies to its stability and perspective for the introduction of valuable selections into cultivation by means of vegetative propagation. By grafting, it is possible to establish hybrids of any genetic complexity. These selections, however heterozygous they may be, can form a new homogeneous generation, similar to the maternal generation, except for some possibly bud mutants or varieties. From one tree of a valuable clone of wild apple, it is possible to obtain hundreds of thousands of buds, which can be propagated by different means of grafting that are well-known from practices in forestry and horticulture. In all of the methods of vegetative propagation, the descendants develop from vegetative buds or from the mother tree meristems by means of mitotic divisions of cells. Here, grafting plays a crucial role for selection as a factor for the preservation and propagation of natural hybrids in their heterozygous state. Selected trees also may be propagated by stem or root cuttings from clones. The numerous examples of sprouting, stool shoot renewal and layers of horizontal branches discovered by us in these wild trees in nature testify to the possibility of using a number of vegetative means of propagation for wild apple clones under cultivation. Specific features of these clones determine the characteristics of apple selections. In introducing these clones into cultivation, there is no need to obtain pure lines, which transmit their positive features to the seed progeny. Instead the fixed hereditary characters of selected trees possess economically valuable properties and these are of interest for selection and production. Valuable peculiarities of one or another tree are firmly fixed in the vegetative progeny, and these are new clonal varieties. Depending on the economic importance of the peculiarities of the mother trees in nature,

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there could be several such variety-clones. The vast apple polymorphisms provide an opportunity for such a choice. As Tables 2.43–2.46 and other materials show, mutations may be selected in apple in relation to a number of economically important characters: the time of ripening, size, form, color, storage life, dessert quality, chemical-technological properties, and yield of fruits. Some forms may be introduced directly into cultivation, while others may be used in breeding to increase the resistance and productivity of old cultivated apple varieties. For example, in breeding for late winter varieties, the resistant selections 28, 35, 269, and 43 (Dzhangaliev 1968) with good keeping quality and high-eating quality are of special interest. Apple trees with globular crowns, thick foliage canopies, that are impervious to the sun rays, and also having beautiful flowers and pink fruit are characterized as highly ornamental by M. G. Popov. A number of wild apple forms may also be of great interest to the food industry. On the basis of chemical-technological fruit investigations, the following trees were singled out from 324 selections for seed preparation and planting material and for use in breeding (Dzhangaliev 1973d, 1976c). High-vitamin. Selections 49, 328, and 329 from Dzhungarskei Alatau. They exceed cultivated varieties in vitamin C content by six to seven times (64.5–70.8 mg/100 g fresh weight). Several selections possessing multivitamins (C, B9, P), which are very seldom seen, were included in this group. Dessert Apples. Selections 28, 43, 35, 269, and 1037 are from Dzhungarskei Alatau, selection 238 is from Zailijskei Alatau and the Karatauskayaform is from Karatau. These selections have pleasant sourish-sweet or sweet fruits with a weight of not less than 60 g and an attractive appearance. Selections 28, 238, and 269 have high ascorbic acid contents, and may also be referred to as the first group and are recommended for dietary nourishment. Relative to their chemical composition, these selections are similar to cultivated table varieties, and selections 238 and 269 contain large amounts of tannin, which is a positive factor connected to the P vitamin activity of substances of polyphenolic nature. Processing for Cider. Selections 5, 18, 25, 48, and 60 are from Dzhungarskei Alatau. Fruits are rich in sugars (10.3–11.9%), pectin (0.73– 1.34%), and tannin (0.60–0.69%), with sweet-bitter and bitter flavor. Processing for Canning. All selections, depending on tannin content and acid, may be used for different kinds of products. Since these clones are cross-pollinating trees, which are under the influence of local growing conditions, genotypic changes may appear in the descendants. These

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changes disturb the homogeneity of individual trees, which points to the necessity of repeated selection from groups of clones. Some selections of M. kirghisorum have good fruit quality but are not winter hardy under the less favorable conditions of the open plain. It is possible to obtain greater winter hardiness by selecting hardy seedlings from seed-sown progeny. P. I. Lapin (1959), using specimens of Robina pseudoacacia and Gleditschia triacanthos, demonstrated this technique. A study of hybrids obtained by artificial crossing of the different wild selections with cultivated apple varieties (Dzhangaliev and Katseiko 1969; Dzhangaliev 1976b) showed the possibility of their use in hybridization, and that the wild selections are distinguished by individual hereditary peculiarities. The first hybrid generation was very heterogeneous and the many separate economically valuable characters made it good material for selecting new types. A hybrid analysis of 16 hybrid families bred from cultivated varieties crossed with wild selections of M. sieversii from Zailijskei Alatau, selections 46, 73, 114, 128, and X2 (with late flowering, good productivity and satisfactory fruit flavor), and also M. niedzwetzkyana crossed with cultivated varieties showed that these wild forms transmit winter hardiness to their seedlings in 25 to 48 percent of the cases. But selection 46 is not winter hardy. This points to the necessity of making the right choice of parents when making crosses. Wild maternal forms produced 75 percent precocious seedlings and 25 percent medium precocious hybrids, while in reciprocal crosses, 71.2 percent of the hybrids had medium-length periods to initial bearing. Consequently, relative to precociousness, wild forms are widely varied, and it is necessary to choose precocious forms as parents. Among the hybrids, there were 75 percent high-yielding (50 kg or more per tree) or medium-yielding (20–50 kg) trees. Selections 73, 128, and M. niedzwetzkyana were the parents of many such hybrids. It is especially valuable that most of the hybrids ripen in winter or autumn. Selections 46, 73, and M. niedzwetzkyana are distinguished for their seedlings, which ripen in autumn or early winter. Good color, flavor, storage life, and fruit shapes of wild apples predominate in the progeny. Promising hybrids are characterized by their winter hardiness, precocious bearing (in the fourth year), high-yielding (87–152 kg from a 15-year-old tree), tasty, large fruits (130–200 g) that keep until June. Crossing between cultivated varieties and wild apple forms, both of which have economically valuable characters, and then selecting good “F1” segregants, is a useful strategy. Crossing of wild fruit plants with cultivated varieties provides for “hybrids” as parents for the next generations. Further segregants can be obtained by self-pollination or by

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crossing with a cultivated variety which possesses economically valuable characters, or by distant hybridization among different species and genera as well. Selection variability relative to a number of characters is genetically imprinted. When growing under other soil-climatic conditions, the individuals of a population keep their own specific distinctive features. Some of these features are rhythms of their vital processes, differences in productivity, the features of their morphological tree structures, and pomological characters of the fruits. Consequently, “form” differences expressed by a number of characters are not simple phenotype reactions to the habitat. Thus, analysis of the data of hybrid progeny of wild apple forms crossed with cultivated varieties show that characteristics act independently. The main objective in crossing old cultivated varieties with newly discovered forms of wild apple is to select varieties that are immune to the most dangerous and widespread diseases and pests. In the apple stands of Kazakhstan, there are individual trees, that are little damaged by pathogens, even when they grow beside trees, that are highly susceptible to scab and apple powdery mildew. Undoubtedly, the resistance of these forms is the result of natural selection in areas where disease is always present. Historically, during the selection process of natural forests, evolution of both the host plant and the parasites occurs (Zhukovsky 1971), and this causes intraspecific differentiation to take place, which helps highly immune forms of apple trees to survive and to be preserved. These possess considerable field resistance to pathogens and parasites. Using them in breeding may be a great success. Field resistance to diseases is widespread in some other wild apple species as well. Williams (1968) reported that clones of Malus prunifolia and Malus floribunda possess genes for resistance to Venturia inequalis. Resistance of the mountain apple populations to severe climatic conditions during flowering, fruit development and seed development is very important. The temperature fluctuates widely and is often even below zero during flowering. Here, only those individuals that possess some generative adaptability to severe conditions of the habitat are able to produce fruit. The individuals of named populations are important for selection. Thus, the urgent problem is to create graft plantations of valuable wild apple trees and permanent forest seed plots on the basis of highly productive and high quality populations of fruit forests. A literature review testifies to the fact that such projects on wild apple species and other fruit plants are not conducted on a republic or country-wide scale. Now is the time to fill the genepool with outstanding trees on the basis

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of specialized forest-fruit seed farms and to move to wild species selection on a strict scientific basis using sophisticated selection and genetic techniques. VII. UTILITY AND BIOCHEMICAL CHARACTERIZATION OF WILD APPLE FRUIT The chemical composition of apples is quite variable depending on several factors. Under the influence of the environment, these variables are mostly of a quantitative character. The qualitative features of fruit are systematic characters inherent in a species and lower taxa. Much data on fruit chemistry and its changes depending on weather conditions during the growing season have been accumulated (Sitnikov 1937; Tserevitinov 1949; Sapozhnikova 1955; Lukovnikova 1958; Lomakin 1962; Shuruba 1972 and others). The effects of geographical zones were studied by Vecher and Bukin (1940), Grebensky (1941), Arasimovich (1948), Doroganevskaya (1948), Malyshev (1948, 1957, 1962); Tserevitinov (1949); Ermakov and Lukovnikova (1959) and others. The effects of irrigation were investigated by Shifrina (1959) and Mizgireva (1962), and time of ripening by Franchuk and Kulik (1955) and Arasimovich (1962). All of these studies were devoted to cultivated apple varieties. But, there were a few investigations concerning the chemistry of wild apples that considered a limited range of indexes: sugar, acidity, and vitamin C. These indexes were not linked with the numerous other factors that have an influence on fruit quality. The processing characterization of wild apples is insufficient and this is what prevents their rational use. Processing characteristics of fruit are determined by their chemical composition, which greatly depends on species and intraspecific peculiarities, relative to their growing conditions. Thus, each type bears specific characters in each growing region. Studies were made of the chemical and processing peculiarities of wild apple fruits of Kazakhstan. These studies found valuable selections, the most promising of which were ready for direct introduction into cultivation, for selection, and for determining a rational direction for their use. Selections of M. sieversii were investigated. In some cases, when apples of other species have also been analyzed, a corresponding reference has been cited in the book. The biochemical compositions of wild apple fruit selections were studied from 1948 through 1973. The fruits for analyses were harvested at the stage of picking maturity during expeditions to apple woods in Zailijskei, Dzhungarskei, Talasskei Alatau, Karatau, and Tarbagatai. Fruits of the regionally cultivated varieties, ‘Suislepper’ (summer),

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‘Renet Burhardt’ (autumn) and ‘Aport Alexander’ (winter) were harvested as controls. The biochemical indexes of wild apple fruit selections were studied in relation to the growing site, meteorological conditions during the growing season, the time of ripening, and flavor type. The following indexes were determined in the fruits: dry substance by drying to a constant weight; reducing sugars, sucrose and total sugars by Bertran’s method; titratable acidity by alkali titration; soluble pectin, protopectin and the sum of pectin substances by the weight of calcium-pectate method; tannin and pigment substances by the method of Nejbauer-Levental (Ermakov 1952); cellulose by direct weight method (Belozerskij 1951); vitamin C by the hydrogen sulfide method (Devyatinin and Iosikova 1954); total catechin by the vanillin method (Vigorov 1964); vitamin B9 by the fluorimetric method (Andreeva 1953); and microelements according to N. G. Zyrin (1962), and A. A. Kvetkina and Z. I. Shlavitskaya (1963). From 1961 to 1967, the processing qualities of wild apple fruit types were studied. The following kinds of products were made from them: natural juice, juice with sugar, cider, calvados (brandy), strong wine, compote, jam, apple sauce, fruit filling for pies, marmalade, soufflé, and fruit filling for caramel (Dzhangaliev 1973e). Chemical analyses of the finished products were made using the same indexes as those used for fruit analyses. A quality estimation of these products was made for cider by the Republic Testing Committee, and for the other products, by the Commission of Specialists. Preserved products were evaluated by a 5-point system, and wine-making products by a 10-point system. Table 2.49 shows that the average dry substance content of M. sieversii fruits during 1948 to 1966 was higher than that of cultivated apples. Dry substance fluctuated between 14.26 and 17.61 percent during those years. The greatest quantity of dry substance in fruits was found during 1948, 1949, and 1951. M. sieversii fruits are characterized by a lower average content of dry substance than those of M. baccata and M. manshurica, in which the dry substance contents were 35.93 and 23.79 to 32.53 percent, respectively (Branke 1935). Compared to the dry substance content of M. pumila fruit, which is between 11.07 and 14.80 percent (Kostyk 1950; and Dolidze et al. 1966), dry substance content of M. sieversii fruit is higher. According to this index, M. sieversii is similar to the fruits of wild apple in southern Kirghizia, which range in dry substance content between 10 and 19 percent (Bezzubov 1949) and in Tajikistan, with 15.91 percent (Speranskij 1936). Wild apples are inferior to cultivated apples in the amount of soluble sugar. The difference in sugar content is not great (1.18%) and is affected by the reducing monosugar content. In some years (1962 and 1965), the

248

Year

1948 1949 1950 1951 1960 1961 1962 1964 1965 1966 1948– 1966

1949– 1966

M. sieversii

Cultivated

22

68 67 16 25 11 20 37 26 32 22 324 14.49± 0.20

17.61 16.78 16.50 17.14 15.60 15.18 15.22 14.26 15.10 15.09 16.17± 0.14

Dry weight

6.62± 0.15

– 4.57 4.01 4.18 4.34 5.66 6.18 5.63 6.65 5.28 5.23± 0.05

Monosugar

2.90± 0.12

– 3.16 3.13 3.68 2.84 3.17 3.20 2.21 2.66 3.68 3.11± 0.02

Sucrose

Sugar

9.52± 0.58

8.43 7.73 7.14 7.86 7.18 8.83 9.38 7.84 9.31 8.96 8.34± 0.04

Total sugar

0.86± 0.08

1.55 1.08 1.15 – 1.25 1.04 0.96 0.86 1.32 1.15 1.19± 0.013

Pectins

0.75± 0.05

– – – – – 0.92 1.10 1.02 1.02 1.10 1.04± 0.06

Cellulose

0.065± 0.004

0.475 0.551 0.476 – 0.317 0.412 0.420 0.434 0.456 0.428 0.466± 0.032

Tannins and pigments

Content (% by fresh weight)

0.58± 0.03

1.11 1.12 1.30 1.09 1.10 1.04 1.02 1.02 1.20 1.21 1.11± 0.04

Titratable acidity as % malic acid

16.41

7.59 6.90 5.49 7.21 6.53 8.49 9.20 7.69 7.76 7.40 7.51

Sugar:acid ratio

4:20 PM

Number of specimens (forms)

Chemical composition of M. sieversii and cultivated fruits.

8/8/02

Apple type

Table 2.49.

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sugar content of wild apple fruits was not inferior to those of cultivated fruit (Table 2.49). A great number of forms with high sugar contents were selected. Thus, in 1962, there were 13 of the 37 investigated forms (35%), and 1965, 14 of the 32 (44%) of the forms that had high sugar content. Pectin is intensively accumulated in the fruits of wild apple. According to average long-term data, pectins in wild apple fruits were higher than in those fruits of cultivated varieties. In most cases pectins were higher than 1 percent and in some forms, pectins were 3.28 percent. In the literature, there are indications of a somewhat lower pectin content in the wild apple fruits of Zailijskei Alatau, from 0.10 to 0.99 percent (Popov et al. 1935). In this case, the difference in the number of specimens investigated should be taken into consideration. These authors made fruit analyses of only 22 apple forms and, certainly, they could not have covered all of their variations. We investigated more than 300 specimens. Comparatively, M. manshurica fruits have little pectin, 0.38 to 0.59 percent, and M. sibirica fruits are low in pectin, 0.15 percent (Branke 1935). Fruits of M. pumila from Georgia (Dolidze et al. 1966) and M. sylvestris from the town of Kropotkin (Saburov and Grzhivo 1931) have pectin contents similar to the fruits investigated by us. The high content of pectic substances in wild apple fruits should be considered as a positive fact, because this allows their use in the confectionery and fruit-processing industries for the making of products with a jelly-like consistency without adding jelly. There is more cellulose in wild apple fruits (0.92–1.10%) than in cultivated varieties (0.75%) (Table 2.49). However, there are other data which show that 2.47 percent of cellulose was found in M. sieversii fruits (Saburov and Grzhivo 1931), 2.3 to 2.5 percent in M. pumila (Sorokin 1947), and 5.92 percent and 3.49 to 8.21 percent in M. sibirica and M. manshurica respectively (Branke 1935). The fruits of wild apples sharply differ from cultivated apples in the quantity of tannin substance. According to average long-term data (Table 2.49), the tannins in wild apples were seven times higher than in fruits of cultivated apples. Fluctuations of the average annual tannin content ranged between 0.317 and 0.551 percent. Variations between forms were rather great. A large amount of tannin substances adds bitter flavor and astringency to fruits, making them almost inedible for fresh use. However, varieties with high tannins are good for cider production because these substances prevent wines from oxidation and promote finishedproduct clarification. In wild apples of other species, approximately the same amount of tannin substances was found: in fruits of M. orientalis, 0.46 to 1.09 percent (Dolidze et al. 1966); M. sylvestris, 0.23 to 0.58

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A. DZHANGALIEV

percent (Saburov and Grzhivo 1931; Akimushkin et al. 1935; Vecher and Bukin 1940); and M. sibirica, 0.29 percent (Savitskij 1938). Wild apple acidity is two times higher than that of cultivated apples (Table 2.49). Acidity was slightly higher in 1950, but in other years acidity differed little from the average long-term indexes. The different forms had a large range of acidities. Some forms with fresh-sweet flavor have very low acidity (0.10–0.15%). The literature reports a higher acidity for wild apple fruits in comparison with cultivated apples (Branke 1935; Tserevitinov 1949; Khetagurov 1958; Dzhangaliev 1973e). The low sugar and high acidity of wild apple fruits results in a low sugar:acid ratio. The ratio for wild apples was less than half that of cultivated apples (Table 2.49). Large fluctuations in the sugar and acid contents in the forms of wild apple result in very wide differences in the sugar:acid ratio. Relative to their chemical compositions, a great variety exists in the fruits of M. sieversii forms. The extreme limits for each of the studied indexes were much wider than they were for cultivated apples. Variations in tannins, pectins, acidity, and sugar-acid ratios were especially wide. Fruits and berries are rich sources of natural vitamins. Strawberry, current, and raspberry are especially rich in ascorbic acid, P-active substances, and folic acid. However, they are perishable products and the period for their consumption is limited to one to two months. Apples can be preserved for long periods after being picked from a tree without much change in quality. Under the proper conditions, many late-ripening varieties can be preserved until a new crop is harvested. Consequently, apples are still the only fruits which may be used fresh during the whole year. The interest for their vitamin content is therefore natural. Numerous investigations conducted in different regions of the former Soviet Union showed low vitamin content in cultivated apples (Petrova et al. 1937; Tserevitinov 1949; Vigorov 1961, 1964, 1968b; Glasova and Sabov 1964; Keseli 1966; Gachechiladze 1968; Sedova 1968; Korobina 1969; Gutmanis 1970 and others). The apples of Kazakhstan are no exception. According to our investigations, the vitamin C content of the fruits of cultivated varieties mainly grown in the Alma-Ata region fluctuate from 3.17 to 18.38 mg/100 g fresh weight (Dzhangaliev and Dsju 1969). Such a scanty amount cannot meet the people’s requirements for ascorbic acid. Because the vitamin content of wild apples is considerably higher than that of cultivated apples, they may serve as a serious substitute. In the fruits of wild apple forms, ascorbic acid is 2.4 times higher than that in local cultivated varieties (Table 2.50). Fluctuations from year to year are 14.4 to 29.0 mg/100 g fresh weight, while fluctua-

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2. THE WILD APPLE TREE OF KAZAKHSTAN Table 2.50.

251

Vitamin C content of M. sieversii fruit forms and of cultivated varieties.

Apple

Year

Average content (mg/100 g fw)

Variation in forms and years

Wild

1948 1949 1950 1960 1961 1962 1964 1965 1966 1948–1966

29.02 15.21 14.40 15.08 17.25 28.29 19.75 14.83 15.83 20.46±0.360

7.30–70.80 7.00–32.50 6.50–22.90 9.66–33.82 10.19–37.57 12.35–55.92 4.50–37.43 4.22–43.74 7.17–29.86 4.22–70.80

Cultivated

1949–1966

8.45±1.02

3.17–18.38

tions between cultivated apple forms are very great, 4.2 to 10.8 mg/100 g fresh weight, which demonstrates wide individual differences and perspectives of form selection relative to vitamin C. L. I. Vigorov (1961) combined all apple varieties into groups according to the vitamin C content: 1 = very poor, with a content up to 5 mg/100 g fresh weight; 2 = poor, 5 to 10 mg/100 g fresh weight; 3 = fruits with lower than average content, 10 to 20 mg/100 g fresh weight; and 4 = fruits with average content, 20 to 30 mg/100 g fresh weight. According to these classes, in 1948 cultivated apples of the Alma-Ata region belonged to the third group; while 8 forms of 68 investigated belonged to the fourth group and 13 clones contained more than 30 mg/100 g fresh weight of ascorbic acid. In 1962, 12 of 37 clones belonged to the fourth group and 14 of them contained more than 30 mg/100 g fresh weight of ascorbic acid. High-vitamin selections may be used both for food, if the flavor qualities of their fruits allow, and for selection work. Fruits of wild apple selections are valuable sources of substances possessing vitamin P activity. Measurements of P-active catechin in fruits conducted from 1964–1966 showed that wild apples contained 7.4 times more P-active catechin than cultivated apples (Table 2.51). The variability of this index is very great both in years and in forms, depending on genetic make-up. The fruits of the 1964 crop possessed especially high P-vitamin activity. There are very few references about P-vitamin activity of wild apples in the literature (Vigorov 1968b). Also, there are no publications on the folic acid content of apples. Our investigations showed that there are no large differences in the content of folic acid

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252 Table 2.51. range.

A. DZHANGALIEV Vitamin content of fruits of M. sieversii and cultivated varieties, mean and

Apple

Year

Wild

1964 1965 1966 1964–1966

Cultivated

1964–1966

Vitamin B9 (folic acid) (mg/100 g fw)

P-active catechins (mg/100 g fw)

0.300±0.030 (0.130–0.500) 0.092±0.080 (0.045–0.166) 0.111±0.006 (0.074–0.188) 0.166±0.011 (0.045–0.500)

1002±1530 (206–2060) 783±640 (146–1561) 371±430 (59–720) 740±270 (59–2060)

0.155±0.024 (0.049–0.370)

100±200 (–240)

between fruits of wild and cultivated apples, however, wild fruits do contain a bit more than the cultivated apples. There are also differences between fruits of cultivated and wild apples in their microelement content. Mn (16.8 mg/kg), Fe (137.4 mg/kg), Zn (24.8 mg/kg), and B (13.4 mg/kg) is a bit higher in fruits of wild apple than cultivated apple Mn (11.7 mg/kg), Fe (114.7 mg/kg), Zn (17.4 mg/kg), and B (9.7 mg/kg), Mo (0.16 mg/kg) is a little bit more and Cu (12.2 mg/kg) and I (5.8 mg/kg) are less than those in local varieties (Dzhangaliev 1973e). A. Composition of Apple Fruit Forms with Different Ripening Dates According to the time of ripening, apples are divided into summer, autumn, and winter types. In spite of some exceptions in this division, each group is distinguished by its own chemical composition, which is related to individual hereditary features that regulate metabolism. An accelerated process of vitality is characteristic of summer varieties. As a consequence, soon after harvesting fruits and often when they are still on the tree, over-maturity begins in them, resulting in the decay of sugars, pectins, and organic acids. On the other hand, a delayed course of all processes is characteristic of winter varieties. They usually ripen only in storage (Metlitsky and Tsekhomskaya 1956). Delayed sugar disintegration and high content of protopectin with its slow change into soluble pectin are noted in winter varieties (Sapozhnikova 1955; Gaponenko and Protsenko 1964; Kunyaeva 1964).

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In the apple woods of Kazakhstan, we selected the forms of very early, early, middle and late seasons of ripening. The group of midseason maturing forms (46.2%) constituted the main mass. The dry matter content (Table 2.52) of wild apples ripening in midseason was higher than that of both the early ripening and the late ripening apples (average data for four years). In the fruits of cultivated varieties, dry matter increased from early ripening to late ripening fruits. The same results were obtained on cultivated varieties by E. I. Eliseev (1959) and G. K. Kovalenko (1967). Conflicting results on the sugar content in fruits of different ripening dates have been published. The greatest amount of sugar was found in summer varieties, and the least in winter varieties by M. N. Sitnikov (1937), F. V. Tserevitinov (1949) and K. K. Gutmanis (1970). Contrary data were obtained by G. K. Kovalenko (1967), A. I. Ermakov and G. A. Lukovnikova (1959), and E. I. Eliseev (1959), in which they found that the sugar content in winter and summer apples was nearly equal, or greater than in autumn varieties (E. V. Sapozhnikova 1955). Such contradictions are probably related to biological peculiarities of the variety. Wild apple investigations in Zailijskei Alatau, conducted by M. V. Andrejchuk (Popov et al. 1935), showed that more sugars are contained in late-ripening apples (11.05%) than in middle-ripening (7.9%) and early-ripening (6.94%) apples.

Table 2.52. Dry matter content of wild apple fruits of different ripening seasons. Dry weight (% fresh weight) Time of ripening Year

Early

Middle

Late

1961 1964 1965 1966 1961–1966

Zailijskei Alatau 14.49 15.86 14.18 12.84 16.48 16.19 15.07 15.26 14.97 15.06

14.63 16.47 14.54 14.21 14.68

1962 1964 1965 1966 1962–1966

Dzhungarskei Alatau 13.76 15.92 14.94 15.71 14.82 15.44 13.78 16.47 14.09 15.85

15.16 14.04 14.32 15.19 14.78

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A. DZHANGALIEV

Our four-year investigation on wild apple fruits of Zailijskei Alatau (Table 2.53) partially confirm M. V. Andrejchuk’s data. However, in several years the sugar content of midseason apples was higher than it was in late-ripening apples. The lowest sugar in most years was in earlyripening fruits at both growing sites (Table 2.53). Early ripening fruits also contain both less sucrose and less monosugars. Sugars were also

Table 2.53.

Sugar content of wild apple fruits of different ripening dates. Content (% dry weight) Time of ripening

Sugar

Monosugars

Sucrose

Total sugar

Monosugars

Sucrose

Total sugar

Year

1961 1964 1965 1966 1961–1966 1961 1964 1965 1966 1961–1964 1961 1964 1965 1966 1961–1966

Early Zailijskei Alatau 4.92 5.41 6.15 5.59 5.50 2.99 1.99 2.27 3.22 2.45 7.93 7.40 8.42 8.81 7.95

Dzhungarskei Alatau 1962 5.60 1964 5.76 1965 7.29 1966 5.59 1962–1966 5.83 1962 2.35 1964 2.05 1965 2.20 1966 2.82 1962–1966 2.42 1962 7.95 1964 7.81 1965 9.49 1966 8.41 1962–1966 8.25

Middle

Late

6.40 5.13 6.92 5.47 6.02 3.04 1.80 3.50 3.88 2.92 9.44 6.93 10.42 9.35 8.94

5.10 6.36 6.25 6.10 5.94 3.91 4.88 2.48 2.57 3.17 9.01 11.24 8.73 8.67 9.11

5.82 5.43 7.37 5.27 6.17 3.78 2.60 2.59 3.91 3.34 9.60 8.03 9.06 9.18 9.51

7.54 5.71 5.29 4.69 6.10 2.49 2.43 3.22 4.58 3.08 10.03 8.14 8.51 9.27 8.18

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lower in midseason apples than they were in late apples, agreeing with E. V. Sapozhnikova’s data (1955). There are some indications about a high sucrose content in winter fruits in the literature (Sapozhnikova 1955; and Kunyaeva 1964). Similarly, our data also show high sugar in the late apples from Zailijskei Alatau (Table 2.53). Apples of different ripening dates have different pectin substance contents. Most pectins, especially protopectin, is characteristic of winter keeping varieties (Arasimovich 1962; Gaponenko and Protsenko 1964). Early-ripening forms from both the mountain sites contained more soluble pectins than either mid-season or late apples (Table 2.54). Protopectin was higher in late-ripening forms from Zailijskei Alatau and in mid-season fruits from Dzhungarskei Alatau. The total amount of pectin was high in all apples. According to the average data for four years, earlyripening forms from Zailijskei Alatau contain the most pectin. Fruits of different ripening dates from Dzhungarskei Alatau had little difference in their pectin quantity. Cellulose is a less variable index. Its variations both according to years and to the time of ripening were not large (Table 2.54). A little more cellulose is contained in early-ripening wild apples, but less in lateripening apples. According to the long-term data, the titratable acidities of early-ripening and middle-ripening forms of M. sieversii from Zailijskei Alatau are equal, but in late-ripening forms, it is a bit lower (Dzhangaliev 1973e). Among the wild apples from Dzhungarskei Alatau, a lower acidity is noted in the early-ripening fruits. But, acidity in midseason and late-ripening fruits is almost equal to or higher than it is in early-ripening fruits. These latest data agreed with M. V. Andrejchuk’s results (Popov et al. 1935) on the wild apples of Zailijskei Alatau, in which acidity increased with later maturity. R. M. Kunyaeva (1964), G. K. Kovalenko (1967), and K. K. Gutmanis (1970) demonstrated increasing acidity in late-ripening apples. E. I. Eliseev (1959) discovered the opposite, higher acidity in early-ripening apples. Relative to the content of tannins and pigment substances in M. sieversii fruits, a common regularity was noticed. According to the average of long-term data, their highest amounts are in the early-ripening fruits, and the least in the late-ripening ones (Dzhangaliev 1973e). The same regularity that is, high early, is also noted in the accumulation of the P-active catechins in M. sieversii fruits (Table 2.55). Ascorbic acid, in 1961 and 1964, was most concentrated in late-ripening fruits. However, in 1965, ascorbic acid was greater in mid-season apples, and in 1966, in early-ripening fruits. According to average four-year

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256 Table 2.54.

A. DZHANGALIEV Polysaccharide content of wild apple fruits having different ripening dates. Content (% fresh weight) Time of ripening

Polysaccharide

Year

Soluble pectin

1961 1964 1965 1966 1961–1966 1961 1964 1965 1966 1961–1966 1961 1964 1965 1966 1961–1966 1961 1964 1965 1966 1961–1966

Protopectin

Total pectins

Cellulose

Soluble pectin

Protopectin

Total pectins

Cellulose

Early Zailijskei Alatau 0.491 0.456 0.486 0.896 0.513 0.769 0.473 0.760 0.581 0.641 1.260 0.929 1.244 1.477 1.154 0.880 1.130 1.190 0.700 1.040

Dzhungarskei Alatau 1962 0.514 1964 0.435 1965 0.378 1966 0.652 1962–1966 0.526 1962 0.511 1964 0.387 1965 0.870 1966 0.411 1962–1966 0.505 1962 1.025 1964 0.822 1965 1.248 1966 1.063 1962–1966 1.031 1962 1.430 1964 1.110 1965 1.290 1966 1.180 1962–1966 1.280

Middle

Late

0.414 0.255 0.639 0.833 0.470 0.605 0.554 1.048 0.260 0.607 1.019 0.809 1.687 1.093 1.077 0.890 0.920 1.000 1.140 0.950

0.333 0.420 0.290 0.405 0.370 0.595 0.526 0.607 0.892 0.870 0.928 0.946 0.897 1.297 1.090 0.950 1.010 0.880 0.820 0.900

0.378 0.293 0.410 0.828 0.472 0.526 0.678 0.862 0.302 0.690 0.904 0.971 1.272 1.130 1.091 0.980 1.140 1.030 1.090 1.030

0.520 0.166 0.393 0.850 0.496 0.490 0.578 0.908 0.403 0.550 1.010 0.744 1.301 1.253 1.046 0.940 1.000 1.190 1.080 1.020

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2. THE WILD APPLE TREE OF KAZAKHSTAN Table 2.55.

257

Content of biologically active substances in wild apple. Content (mg/100 g fresh weight)

Biologically active substance

Vitamin C

Vitamin B9

P-active catechins

Vitamin C

Vitamin B9

P-active catechins

Time of ripening Year

1961 1964 1965 1966 1961–1966 1961 1964 1965 1966 1961–1964 1961 1964 1965 1966 1961–1966

Early Zailijskei Alatau 13.860 24.700 14.080 15.330 17.990 – 0.260 0.090 0.070 0.161 – 1483 1127 59 1172

Dzhungarskei Alatau 1962 27.620 1964 21.090 1965 16.510 1966 22.450 1962–1966 23.380 1962 – 1964 0.280 1965 0.149 1966 0.099 1962–1966 0.166 1962 – 1964 1295 1965 1190 1966 402 1962–1966 826

Middle

Late

15.470 24.420 19.510 12.660 17.850 – 0.310 0.085 0.097 0.191 – 908 466 297 621

19.000 33.920 10.590 10.760 15.750 – – 0.240 0.085 0.141 – – 346 572 491

29.190 24.170 11.270 11.600 20.360 – 0.380 0.067 0.110 0.133 – 1228 737 466 729

24.070 16.740 34.410 13.650 21.320 – 0.380 0.142 0.145 0.207 – 999 1149 321 695

data, late-ripening forms had the least vitamin content, while early and mid-season forms had practically equal amounts of ascorbic acid. Also, according to average long-term data on apples from Dzhungarskei Alatau, vitamin C was highest in early-ripening fruits (Table 2.55). There are comparatively few data about the folic acid content of apples in the literature. Our investigations showed that in apple fruits from Zailijskei

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A. DZHANGALIEV

Alatau, vitamin B9 was highest in mid-season apples, but lower in lateripening apples (Table 2.55). Thus, in conclusion, late-ripening wild apples from Zailijskei Alatau and mid-season apples from Dzhungarskei Alatau are characterized by their high contents of total sugars, sucrose and protopectin in comparison with apples of other ripening dates. High contents of cellulose, tannin, pigment substances, P-active catechins, and vitamin C are specific to early-ripening apples. A knowledge of the chemical composition of wild apple fruits according to the time of ripening will aid in determining wild apple use as a raw material for industrial processing and to decide on the duration of fruit storage. B. Composition of Wild Apple Fruits in Relation to Environmental Growing Conditions As investigations on varieties of cultivated apple show, different factors influence the chemical composition of the fruits. We studied the influence of the growing region, habitat, and meteorological conditions during the growing season. Special attention was given to a comparative study of the chemical composition of wild apples from Zailijskei and Dzhungarskei Alatau where the main masses of apple forest are concentrated. Because the forests are situated in the mountains, we selected the various samples for chemical analysis at the same altitudes in the middle-mountain zone in order to exclude the influence of the altitude factor on the time of fruit ripening. If the dates of apple ripening are related to hereditary factors, trees growing under the same ecological conditions, often side by side, may have different dates of fruit ripening. We disregarded the data which show that when moving from the north to the south and from the east to the west, the content of dry substances and sugars increases, while acidity reduces in cultivated fruits (Sorokin 1947; Tserevitinov 1949; Ermakov and Lukovnikova 1959). Also, it has been shown that the northern and the mountain apples are richer in vitamin C than the same varieties when grown in low places (Gorskaya 1937; Kudryashov 1953; Dragavtsev 1956; Malinovskaya 1968). In spite of the considerable influence of environmental factors on the chemical composition of fruits, the main role is played by the individual variety’s peculiarities, which are firmly imbedded, irrespective of growing conditions. Thus, according to E. P. Franchuk and A. A. Kulik (1955) high-sugar cultivars preserve this quality under any conditions. T. A. Kezeli (1966), L. I. Vigorov (1968b) and others showed that the vitamin C content remained steadily high.

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Table 2.56. Chemical composition of wild apple fruits from different growing regions (1961–1966). Content Variable Dry weight (% FW) Monosugars (% FW) Sucrose (% FW) Total sugars (% FW) Soluble pectin (% FW) Protopectin (% FW) Total pectins (% FW) Cellulose (% FW) Tannins and pigments (% FW) Titratable acidity as % malic acid Sugar:acid ratio

Zailijskei Alatau

Dzhungarskei Alatau

14.89±0.34 5.88±0.20 2.93±0.21 8.81±0.23 0.439±0.047 0.659±0.044 1.098±0.071 0.94±0.07 0.380±0.069 1.12±0.18 7.87

15.04±0.30 6.02±0.32 2.98±0.27 9.00±0.30 0.501±0.045 0.569±0.046 1.070±0.079 1.10±0.05 0.465±0.053 1.08±0.18 8.33

As it is seen in Table 2.56, the lower humidity and lower air temperature in Dzhungarskei Alatau resulted in a slightly greater accumulation of dry substances, monosugars, soluble pectin, cellulose and tannin substances than in Zailijskei Alatau. The sucrose content and acidity in apples from different growing regions were practically equal, but the protopectin quantity and the total pectin substances in the fruits from Dzhungarskei Alatau were a bit less in comparison with those of the fruits from Zailijskei Alatau. Climatic conditions in Dzhungarskei Alatau proved to be more favorable for the accumulation of biologically active substances (vitamins) in the fruits (Table 2.57). Of the factors that determine the climate, humidity exerts the greatest influence on growth and formation of fruit quality (Sharapov 1954, 1962; Vasilyev 1959; Shuruba 1972). There are some data in the literature which show a considerable influence of meteorological conditions during the growing season on the chemistry of cultivated apples. A higher accumulation of dry substances and sugars and a lower acidity in fruits were noted during droughty years (Tserevitinov 1949; Sharapov 1954; Arasimovich and Vasilieva 1960; Arasimovich 1962). Vitamin C content was higher in the fruits in rainy years and is lower in droughty years (Kropacheva 1961; Vigorov 1968a). Table 2.58 shows the analysis of the chemical composition of fruits in 1962 and 1966. These years had approximately equal amounts of heat (mean yearly air temperature was within 8.1–8.8°C), but differed in moisture content. In 1962, there was minimum precipitation in com-

260

15.88±2.580 (4.40–37.43)

1964– 1966

17.28±2.330 (4.24–43.74)

16.39±1.650 (7.22–29.86)

16.28±2.550 (4.22–43.74)

0.165±0.011 (0.045–0.380)

0.092±0.007 (0.074–0.112)

0.086±0.007 (0.045–0.140)

0.270±0.020 (0.160–0.380)

Zailijskei Alatau

0.167±0.011 (0.051–0.500)

0.116±0.007 (0.074–0.188)

0.090±0.010 (0.051–0.166)

0.340±0.040 (0.130–0.500)

Dzhungarskei Alatau

Dry weight

16.47 15.21 15.11 15.07

Precipitation

1962, dry (686 mm) 1966, wet (1070 mm)

1962, dry (425 mm) 1966, wet (520 mm)

9.22 8.90

11.24 9.22

Total sugars

0.96 1.14

0.95 1.19

Pectins

1.04 1.18

Dzhungarskei Alatau 1.10 0.42 1.12 0.45

Titratable acidity as % malic acid

0.78 1.35

Tannin and pigment

0.47 0.34

Zailijskei Alatau 1.01 1.03

Cellulose

Content (% fresh weight)

27.70 16.40

38.90 13.30

Vitamin C (mg/100 g FW)

699±930 (59–1823)

237±460 (59–330)

662±880 (146–1360)

872±144 (206–1823)

8.86 7.61

14.41 6.82

Sugar: acid ratio

770±810 (154–2060)

401±400 (154–720)

889±410 (232–1561)

1155±1620 (29–2060)

Dzhungarskei Alatau

P-active catechins Zailijskei Alatau

Chemical composition of wild apple fruits in different years based of area and precipitation.

13.33±3.230 (7.17–22.49)

1966

Table 2.58.

13.18±1.810 (4.40–29.22)

1965

20.05±2.790 (4.50–33.49)

Dzhungarskei Alatau

Vitamin B9

4:20 PM

19.50±2.700 (6.81–37.43)

Zailijskei Alatau

Vitamin C

Content (mg/100 g fresh weight)

Vitamin content of wild apple fruits from different growing regions.

8/8/02

1964

Year

Table 2.57.

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parison with other years, and these were considerably less than the average long-time norms (806 mm in Zailijskei Alatau and 485 mm in Dzhungarskei Alatau). However, in 1966, considerably more rain fell (1070 mm in Zailijskei Alatau and 520 mm in Dzhungarskei Alatau). In wild apple fruits from Zailijskei Alatau, a comparison shows that there were considerably higher contents of dry matter and sugars and fewer acids in the drought year of 1962 than there were in the wet year of 1966. In fruits from Dzhungarskei Alatau, the differences in dry matter, sugars and acids were not so great, though the same tendency occurred. Probably, these trends were related to the differences between the 1962 and 1966 precipitation amount, which was much less in Dzhungarskei Alatau than in Zailijskei Alatau. Our data on vitamin C content contradict data in the literature. The vitamin content in wild apples in a drought year was considerably higher than it was in a wet year. This difference was apparently associated with differences in metabolism, which are related to the responses of wild and cultivated fruits to their environments. The cellulose contents of fruits in the dry and wet years were approximately equal. C. Composition of Wild Apple Fruits That Have Different Flavor Types In order to determine the processing value of wild fruit crops from existing plant communities and to determine the choice of methods for their use, a system of fruit classification according to flavor types was devised (Dzhangaliev 1951, 1968, 1973e). During the inventory of fruit forests of Zailijskei and Dzhungarskei Alatau, a study of the different flavor fruit types was made. The study showed that 20.8 percent of the apples were sour, 20.8 percent were sourish-sweet, 19.0 percent were sweetish-sour, 16.6 percent were sourishbitter, 12.8 percent were sweetish-bitter, 8.7 percent were sweet, and 1.3 percent were bitter. Thus, in apple woods the forms with sour fruit flavor prevail, which should be taken into consideration when determining the trend of their use. An analysis of the chemical composition of different flavor types (Table 2.59) showed that the content of dry matter, cellulose and pectic substances were not related to fruit flavor. Differences in sugar content were also insignificant. The main differences relative to fruit flavor types were tannin substances and acidity. On the basis of these results (Dzhangaliev and Bojkov 1973), the limits of the contents of acid and tannin substances were established as objective criteria for assigning fruits to definite flavor types (Table 2.60). The fruits from different growing

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Table 2.59. Chemical composition of different-flavored wild apple fruits harvested in 1948–1966. Content (% fresh weight)

Titratable acidity as % malic Sugar: acid acid ratio

Flavor fruit type

Dry weight

Total sugars

Pectins

Tannins

Cellulose

Sweet Sourishsweet Sour Sweetishsour Bitter Sweetishbitter Sourishbitter

17.2

9.36

1.07

0.32

1.04

0.30

31.20

15.9 15.7

8.88 7.28

1.26 1.10

0.29 0.34

0.94 1.01

0.84 1.70

10.57 4.28

15.8 16.4

8.73 8.52

1.11 1.32

0.32 0.80

0.94 1.28

1.22 0.91

7.15 9.36

17.3

9.61

1.25

0.61

1.13

0.61

15.75

15.8

8.18

1.15

0.63

1.11

1.45

5.64

Table 2.60. Minimum acid and tannin contents of apples relative to fruit flavor types. Content

Flavor fruit type

Titratable acidity as % malic acid

Tannins (%FW)

Sweet Sourish-sweet Sweetish-sour Sour Sweetish-bitter Bitter Sourish-bitter

0.6 0.6–1.0 1.0–1.4 1.4 0.7 0.7–1.1 1.1

0.5 0.5 0.5 0.5 0.5 0.5 0.5

places, grouped only according to acidity and tannin substance content, have very similar indexes according to the other components of the chemical composition. Fruits with a sour flavor are characterized by the lowest content of monosugars, total sugars and low sugar:acid ratio. Fruits of sweetishbitter type contain the highest amount of sucrose and total sugars. Sourishsweet fruits have the lowest content of vitamin C and tannin substances. The high indexes are peculiar to the fruits with sweetish-bitter, bitter and

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Table 2.61. Vitamin content of different flavor types of wild apple fruits (1964–1966). Vitamins (mg/100 g FW) Flavor type

C

B9

P-catechins

Sweet Sourish-sweet Sweetish-sour Sour Sweetish-bitter Bitter Sourish-bitter

Zailijskei Alatau 12.48 0.172 9.02 0.188 19.66 0.083 19.38 0.139 – – 22.83 0.214 21.99 0.233

826 459 333 713 – 1422 1162

Sweet Sourish-sweet Sweetish-sour Sour Sweetish-bitter Bitter Sourish-bitter

Dzhungarskei Alatau 16.99 0.182 9.27 0.183 13.43 0.166 18.58 0.188 13.66 0.169 27.66 0.216 25.43 0.172

611 494 569 590 994 1512 1017

sourish-bitter flavor types. Bitter and sourish-bitter types also contain the highest amount of P-active catechins and vitamin B9 (Table 2.61). Thus, the classification of fruits according to flavor types reveals the main chemical peculiarities of wild apples. D. Processing Fruit Characteristics of Promising Wild Apple Forms The great diversity in wild apple populations allow for the selection of clones which are not inferior to some of the recognized cultivars, and according to a number of indexes, they even may be better. Clones with large fruits, attractive appearance and good fruit flavor were selected. Also, a number of clones rich in vitamins were singled out for use in breeding for high-vitamin varieties and for introducing into cultivation (Dzhangaliev 1973d). Using chemical fruit composition (Table 2.62) as a guide, eating types were selected, all distinguished by their low content of tannins, which determines their suitability for fresh consumption. The sugar:acid ratios approach optimal levels in all selections except 238 and 1037. Vitamin content is close to that of cultivated varieties. Fruits of selected forms are very attractive in appearance, possess rather

264

28 35 43 269 1037 238 1043 1051

Sourish-sweet Sourish-sweet Sourish-sweet Sourish-sweet Sweetish-sour Sweet Sweet Sweet

Flavor type 14.98 15.03 13.65 14.14 12.71 16.12 15.41 15.93

Dry matter 9.75 9.10 9.13 9.31 7.62 10.80 10.40 9.96

Total sugars 0.850 1.035 1.062 0.899 0.650 0.940 – –

Pectin 0.100 0.104 0.103 0.201 0.043 0.405 – –

Tannin and pigment 18.05 10.22 13.04 15.26 8.28 49.09 26.67 31.12

Sugar:acid ratio 0.54 0.89 0.70 0.61 0.92 0.22 0.39 0.32

Titratable acidity as % malic acid

10.10 9.04 4.50 13.61 8.94 17.69 4.90 5.00

C

0.047 0.181 – – 0.034 – – –

B9

Vitamins (mg/100 g FW)

263 213 321 – 214 – 273 328

P

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Content (% fresh weight)

Chemical fruit composition of promising wild apple selections suitable for eating fresh.

8/8/02

Selection no.

Table 2.62.

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2. THE WILD APPLE TREE OF KAZAKHSTAN Table 2.63. industry.

265

Chemical composition of wild apple selections, promising for the canning

Content (% fresh weight)

Selection no. 313 212 324 325 326 215

Flavor type

Dry matter

Total sugars

Total pectin

Tannin and pigment

Sourish-sweet Sourish-sweet Sourish-sweet Sourish-sweet Sweet Sweet

23.5 20.3 17.8 19.8 22.5 21.6

12.40 11.25 10.64 10.64 10.05 10.05

1.52 2.81 2.37 2.69 2.93 2.00

– 0.352 0.454 0.488 0.534 0.535

Titratable acidity as % malic acid

Vitamin C (mg/100 g FW)

0.72 0.81 0.78 0.93 0.42 0.38

13.00 20.72 28.38 13.84 63.80 70.80

pleasant flavor and have good keeping quality. Their size is average or large. Sweetish-bitter forms, rich in sugars, pectin and tannin substances with an average acidity are of great value as raw material for cider production. High tannin substances cause rapid clarification and stability of cider during storage (Dzhangaliev and Bojkov 1973, 1975). Selections with high contents of pectin substances (Table 2.63) are promising for the canning industry. These forms have average acidity, contain much sugar and few tannin substances. High pectin substances make it unnecessary to add jelly components during processing, and secondly their biological activity as antioxidants is beneficial for human health. E. Changes in the Chemical Composition of Wild Apples During Storage One of the most valuable attributes of apple is their ability to ripen during storage. By regulating storage conditions such as temperature, relative humidity, and atmospheric composition, the storage life of many apple varieties can be prolonged for a full year, especially late ripening varieties (Saburov and Antonov 1958; Kunyaeva 1964; Gusejnov 1965; Metlitsky 1965; Bruev 1966). Through long-term investigations on different apple cultivars in various zones of the country, the main processes of metabolism during storage were discovered (Metlitsky and Tsekhomskaya 1956; Kolesnik 1959; Arasimovich 1962). The keeping quality of wild fruits has never been studied in detail. Judging from the late ripening forms and the great variety of forms, there possibly are fruits which have long storage lives. According to M. G.

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Popov et al. (1935), fruits of two wild apple forms from the Alma-Ata region were well preserved until March without quality deterioration. In order to study keeping quality, weight change and chemical composition during the storage process in 1967, apple selection 28 and ‘Aport’ were kept in a fruit storage with artificial air cooling at temperatures between +2 and –2°C and the relative air humidity of 95 percent. The fruits of the autumn-winter variety, ‘Aport Alexander’, served as controls. Apples of selection 28 excellently endured transportation in motor vehicles from Dzhungarskei Alatau (a distance of 500 km). Their good flavor persisted until June. In contrast, the fruits of ‘Aport’ that were placed into storage directly from the orchard, were taken out of storage in February, because of the great amount of fruit rot and flavor deterioration. The results of keeping quality under different temperature regimes testify to the fact that a temperature of 0°C is the most favorable for preserving the nutrient properties and market quality (Table 2.64). During the storage of ‘Aport’ fruits under the same conditions, weight losses because of natural damage were 1.5 times, and waste was 2.2 times greater than of wild fruits (Dzhangaliev 1973d). Thus, the wild fruits proved to be more resistant than ‘Aport’ to destructive influences due to a microbiological and functional character. The dynamics of the main indexes of chemical composition of selection 28 and ‘Aport’ were studied under optimal storage conditions (air temperature 0°C and packed in boxes with a layer of wood shavings). During storage, all substances constituting fruit composition suffer quantitative changes. The dry matter content in fruits of selection 28 declined more intensively than that in ‘Aport’. This shows that natural losses in fruits of ‘Aport’ are mainly conditioned by evaporation. This humidity Table 2.64.

Keeping quality of selection 28 fruits under different storage temperatures. After 131 days of storage

Variable Dry matter (% FW) Sucrose (% FW) Total sugars (% FW) Natural losses (% FW) Waste (% FW) Titratable acidity as % malic acid Vitamin C (mg/100 g FW) Vitamin B9 (mg/100 g FW)

At harvest

+2°C

0°C

–2°C

15.25 2.82 9.91 – – 0.77 11.71 0.04

13.20 0.99 8.81 2.00 16.30 0.37 6.25 0.09

13.44 1.60 8.94 1.50 13.50 0.49 7.98 0.07

14.67 1.33 9.15 1.40 15.00 0.59 6.99 0.09

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loss disguises to some extent the decrease in dry matter content. Evaporation losses are considerably less in fruits of selection 28. The amount of sucrose, malic acid and vitamin C decline evenly during storage. These substances are physiologically active and are the first to be used in respiration. The character of these chemical losses may give some indication of the keeping quality of the fruit (Dzhangaliev 1953; Luvov and Kalugina 1955; Metlitsky and Tsekhomskaya 1956; Arasimovich and Vasilieva 1960; Kunyaeva 1964; Metlitsky 1965; Shirokov 1970, and others.). The fruits of the selection 28 use these substances more economically than ‘Aport’. During storage, the content of pectic substances reduces, too. However, this process proceeds unevenly. The amount of soluble pectin at some stages of storage increases, apparently at the expense of protopectin. In March, an increase in pectin commonly occurs. V. V. Arasimovich (1962) explained that this is due to a possible release of pectins from complex compounds, which result from hydrolytic decay or from pectin formation as a consequence of the incomplete oxidation of sugars to galacturonic acid. The latter is more probable because this increase in pectins occurs simultaneously with a reduction in sugars. The content of vitamin B9 (folic acid) during storage in the fruits of selection 28 increases greatly, probably, at the expense of its release from a complex of compounds. P. G. Tavadze and G. T. Gachechiladze (1972) also observed an increase in the amount of folic acid during the storage of the apple variety, Demiralma. Thus, the fruits of the wild apple selection 28 are characterized by their good keeping quality, which is related to their metabolic features, which provide for a more economical use of their energy substances for respiration. Its tough skin, which is entirely covered with intensive anthocyanin color, penetrating even into the flesh, protects apples from infection by microorganisms (Rubin and Artsikhovskaya 1948; Kapustinsky 1950). F. Processing Characteristics of Wild Apple Fruits The necessity of using wild fruit in the food industry is not only related to their natural attributes, but also because the chemical composition of cultivated varieties does not always meet production requirements. Fruits of cultivated varieties have pleasant flavor and attractive appearance as a result of long-term selection and breeding. Because of breeding, cultivated varieties have lost a number of valuable qualities, which their wild predecessors possess, including a high content of organic acids, tannin and pectins, which are used in industry, and also tree resis-

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tances to unfavorable environments, diseases, and pests. In some countries with developed fruit processing industries (France, Spain), processing varieties have been bred using wild apple species as parents (Vecher and Bukin 1940). Thus, in France, cider varieties are cultivated in great numbers using fruits which have 1 percent tannin, 2.7 percent pectic substances, 0.57 percent acids, and 19.6 percent sugars (Alexandrov 1960). In the apple woods of Kazakhstan, numerous clones grow which are distinguished by biochemical indexes of their fruits. These may be used as raw material for different purposes by food and flavor industries, depending on their fruit quality. However, up to the present time, wild apple fruits have been little used in the processing industry. This is because of an absence of fundamental data on their chemical composition and their processing potential, which take account of the biochemical features of this wild raw material. The available processing procedures are designed for using cultivated fruit varieties as raw material. Following the Decision of the Soviet Ministers of Kazakh SSR No. 435 of June 15, 1967, “Methods of using wild fruits and the waste of cultivated apple varieties suitable for food production,” Dzhangaliev and Bojkov (1969a,b) elaborated cider production methods and methods of beverage production of the calvados type under the name “Zhiger” for which the republic’s technical conditions were established (RTU Kaz. SSR 756–67). On the basis of our materials, Kazgipropishcheprom established “The technical economic basis of complex processing of wild fruits and waste of cultivated varieties, suitable for food production in the Kazakh SSR for 1970–1980.” Our investigations demonstrated the possibility of obtaining high quality food products, which are superior to the quality of comparable products from cultivated varieties. 1. Cider. The cider industry in the former USSR, in spite of a great demand for this drink, has developed very slowly, and in Kazakhstan, it is not commercially produced at all. This lack of production is related to an absence of a special raw material base and to the short-comings of existing technologies for drink production, making production unprofitable. The clarification of cider materials and making a uniform product are the greatest difficulties in its production. The aim of our investigations was to ascertain the suitability of different types of wild apple fruits for cider production. Cultivated varieties served as controls. In the process of cider production, the chemical compositions of fresh and fermented apple juices and ready-to-drink samples were analyzed.

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The analyses showed that the chemical composition of juice was preserved after mashing and pressing the raw materials (Dzhangaliev 1973d). The type of fruit-flavor influences the composition of juices. However, relative to the content of some of the substances, the juice essentially differs from fruits. There are more sugars and acids in the juice, but only half as much tannin. This low tannin is caused by the great intensity of the oxidation processes, taking place at the moment of cell destruction. The juice turns brown with this loss of a great number of tannin substances. Sulfite was added to the pulp, immediately after mashing to prevent oxidation, which also promoted the preservation of natural fruit aromas in the drink. We tested the new methods of tannin utilization from pomace, which utilizes the tannin substances by almost 100 percent and the method of making a dry pomace, which can be preserved for long periods (Dzhangaliev and Bojkov 1975). Organoleptic evaluation of the tested cider samples from wild apple fruits showed that the drinking quality depended on the flavor type of the fruits. Ciders from sweet fruits are insufficiently flavored because of low tannins and low acidity. Ciders from sour fruits are excessively sour and have insufficient alcohol. Ciders from bitter fruits are more harmonious. The drink from mixed juices of different flavor types won the highest rating. The combination of the attributes of all flavor types of fruits enhanced the high qualities of cider: freshness, balance, and flavor completeness. A processing trial evaluation confirmed the efficacy of cider production from wild apples using our technology. 2. Calvados (Hard Cider). In the USSR, calvados production is new. It was first produced in 1964 under the name of “Lithuanian calvados.” The technology of its production is similar to the French technology. The biochemical features of wild apples are suitable for the production of calvados, which keeps the original flavor qualities of the fruit. The name “Zhiger” was given to this drink. During the fine tuning of the production technology of “Zhiger,” special attention was given to several points in calvados production: to the losses of aromatic substances; to the great losses of alcohol; and to the rise in the costs of production, owing to the long aging in oak barrels. The special aspect of our technology is a ten-day alcohol infusion of dry apple pomace at a temperature of 45°C, instead of the long-term aging of alcohol in oak barrels. Chemical analyses of the fresh apple pomace after must separation showed a considerable content of valuable nutrients in calvados: 3.08 to 6.71 percent sugars, 0.19 to 1.68 percent organic acids, and 0.155 to 0.907 percent tannins. During the process of infusion with pomace, the

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apple alcohols become saturated with extracts, and they gain an intensive attractive color, apple aroma and flavor. The best aroma, the aroma of fresh apples, is made by the infusion of alcohol with fresh pomace. However, in order to use the pomace during the entire year and to facilitate washing sugars and acids from the pomace, all of which impedes the production of the standard drink, the pomace is first dried. The liquid infused with pomace acquires the aroma of dried apples. Heat treatment of the alcohol at a temperature of 45°C increases its color intensity and softens the flavor of the drink. According to preliminary data, the production of calvados from wild fruits is 40 percent cheaper than that from cultivated apples. The Republic Testing Committee of the wine industry of Kazakh SSR issued a high approval rating for the drink “Zhiger” and the proposed manufacturing technology, recommending its production in the document, TU MSKH Kaz. SSR No. 46–131–69. For its production, it is necessary to use a mixture of wild apple forms from different flavor groups. 3. Natural Juices and Juices with Sugar. One of the most valuable kinds of products obtained from wild apples are apple juices, that contain a considerable quantity of biologically active substances. The fruits of several forms of sweet, sourish-sweet and sweetish-sour types, that contain a small amount of tannins, can be used as raw materials for the production of natural apple juice. The fruits of the other flavor types are not suitable for apple juice, because some have high acidity (sour and sourish-bitter), and others have a bitter flavor that is preserved in the juice. Any deficiency in wild apple flavor can be covered up during the production of the apple juice by adding sugar. According to the MRTU 18/30–65 law, the fruits of all flavor types and their mixtures are suitable for juice production. The production of juice from wild apples is 30 percent cheaper than from cultivated apples. 4. Canned Fruits. For compote production, sourish-sweet, sweetishsour, and sour forms of wild apples may be used (Table 2.65). Fruit jam of good quality was made from sweet apples and satisfactory jam from sourish-sweet and sweetish-sour forms. Wild apple fruits of sourishsweet, sour and sourish-bitter flavor types may be used for fruit pies. Good applesauce was produced from fruits of all flavor types, but the best sauce came from sourish-sweet and sour apples. For cooking jam, the fruits of all flavor types are suitable, except the sweet type, which does not give the necessary consistency. Mixing with the fruits of other flavor types may neutralize flavor shortcomings of some of the wild

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271

Taste ratings of canned products from wild apple fruits. Rating (2=low; 4=high)

Flavor type Sweet Sourish-sweet Sweetish-sour Sour Sweetish-bitter Bitter Sour-bitter

Compote

Fruit jam

Applesauce

Jam

Pie filling

3 4– 4– 4– 2+ 3– 3

4 3+ 3+ 2 3– 3+ 3

4 4+ 4– 4+ 4– 4– 4–

3+ 4+ 4– 4+ 4+ 4– 4

3– 4– 3 3+ 3– 2 3+

fruit types. Thus, mixing bitter forms with sweet and sour forms can dilute bitter fruit flavors. The results of these investigations showed the following main aspects about chemical composition of the wild apples in Kazakhstan. According to average long-term data, wild apple fruits are distinguished from cultivated varieties by somewhat lower (1.18%) sugars and by more of dry matter (16.17 to 14.49%), more tannins (1.19 to 0.86%) and more cellulose (1.04 to 0.75%). Acidity of wild apples is almost two times higher than those of cultivated fruit and their tannin content is seven times higher. Wild apples are characterized by high amounts of biologically active substances: vitamin C is 2.4 times and P-active substances 7.4 times higher than in the fruits of cultivated varieties. In addition, wild apple fruits contain more microelements (Mn, Fe, Zn, and B) than cultivated apples. The dependence of wild apple chemical composition on the time of ripening has been demonstrated. Individual genes carried by the tree condition this dependence. Late-ripening apples from Zailijskei Alatau and mid-season apples from Dzhungarskei Alatau are characterized by their high contents of total sugars, sucrose, and protopectins. High contents of cellulose, tannins, P-active catechins, and vitamin C are generally characteristic of early-ripening apples. A knowledge of the kinds of chemical compositions of fruits in relation to the times of ripening helps to define what uses can be made of the raw material for the fruit processing industry and to identify the time when a supply of fruit will be available. Differences in the chemical composition of wild apple fruits from different places of origin were determined. Under the harsher conditions of Dzhungarskei Alatau, fruits accumulate more dry matter, sugars and

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cellulose than those in Zailijskei Alatau. The Dzhungarskei apples also contain more C, P, and B9 vitamins. All forms of wild apple fruits were classified into one of seven flavor types: sweet, sourish-sweet, sweetishsour, sour, sweetish-bitter, bitter, and sourish-bitter. Their acidity and tannin content serve as objective criteria for categorizing fruits into a definite type. This classification reflects the main features of the chemical composition of the wild apple fruits of Kazakhstan. During these expeditionary observations, fresh fruit types of apples were selected. Clones with high-vitamin content fruits were selected. Also, it was ascertained that the wild apple fruits of Kazakhstan are excellent raw materials for wine making, canning, and the confectionery industries. VIII. PRESERVATION OF WILD APPLES The natural communities of woody shrub vegetation, wherein wild apple trees grow, form the normal conditions for the preservation and reproduction of the species. These conditions cannot be replaced by any artificial methods of propagation. Thus, it is necessary to preserve wild apples in their natural habitats. In enumerating the problems of wild apple species preservation, we assumed the following conditions based on our complex ecological and biological investigations of the apple woods in Kazakhstan. Apple woods are a component of the plant communities within the mountain regions of southeastern Kazakhstan, and in some cases, they are the main component of the landscape. Apple woods have a role in the formation of the favorable climatic, soil, and ecological environments. They are a valuable natural resource and a source of seed and planting material for forest-fruit nurseries. They include a rich variety, consisting of great genetic value for selection, for direct introduction into cultivation and for the establishment of resistant plantations of valuable forms for various uses and for selections in the breeding of new varieties. These natural apple species may also be of interest for ornamental horticulture. The problems of preserving wild apple species and numerous intraspecific taxa in Kazakhstan are of great importance, far exceeding the bounds of regional interest. A great potential wealth is being formed under different and often harsh conditions in the mountain regions. The possibilities of their yet undiscovered uses point to the feasibility of creating “standards for wild plants.” Such standards would be carefully based on biological principles for treatment of Malus species in Kazakhstan (Dzhangaliev 1975a,b, 1976a,b).

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At present, the rate of human development in the territory has sharply increased creating a real threat of disappearance or degradation, not only for Malus species, but also for the other groups that form the vegetation. As a result of this economic activity, many fruit tree habitats, especially the type III and IV forest plots, have been replaced haphazardly by farms developed without any thorough biological evaluation. This leads to a negative economic effect on the environment. Today, apple habitats are subjected to intensive economic pressures, which are caused by the exploitation of rich mineral resources in the mountains. In addition, recreational activity has increased from populations near suburban forests, that are included in the green zone. In this zone there is a considerable extent of wild apple woods, that are natural. Fruit ecosystems possess wonderful aesthetic features and some transition areas are especially valuable for establishing recreation zones. In this respect, they have great social importance. The environment within recreation zones may be cleaned up and these regions offer a scientificcognitive and cultural-aesthetic interest. But, under conditions of frequent visits to these green zones, it is necessary to have territory planning and a recreation volume based on the vegetation’s capacity for natural self-renewal. Felling of trees and trampling of vegetation negatively impact fruit-forest renewal. The lower stories of the trees are especially affected, and this leads to terracing of the plot. Recreational activities stress the environment, impoverish the living topsoil, make sparser understory and regrowth, thin wood stands and decrease stand longevity and productivity. These activities not only change the plant community but also affect the physiologically active part of apple root systems, the soil. The root system, being in the upper 20 centimeters of the soil horizon, also undergoes a negative stress from recreational activity. The preservation of Malus species may be successfully accomplished by protecting the vitality of populations and the conditions for natural development of the plant communities in which they live. Part of the whole problem of the rational use and enrichment of the vegetative cover is developing good growing conditions and preserving those landscape elements and vegetative communities with which their existence is connected. Solving these problems is possible through a process of community planning aimed at both nature preservation and economic organization. This can be accomplished by working out methods on a rational basis for useful plant resources, crop harvesting, forest reclamation and planning agricultural land use in the mountains. The special role of preservation of species and intraspecific varieties belongs to government reservations. The existing network in the mountains of the Kazakhstan is absolutely insufficient for preserving the flora

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and the various plant communities, including those of wild fruit plants. The Alma-Ata reservations in Zailijskei and Aksu-Dzhabaglinskei in Talasskei Alatau mainly represent intrazonal natural complexes. In the region of the wild fruit forests of Kazakhstan, there is no zonal-specific reservation. A full preservation of genepools and complexes of vegetation is needed that includes natural fruit plant communities. It is necessary to establish a more divisional network of small reservation territories, embracing the wide variety of types and zones of fruit plants. Apple woods of types III and IV that have the maximum productivity of fruit forests and the 15 selected seed plots of excellent and normal stands should be the targets of preservation. Malus sieversii is a species with a sufficient source of raw materials. Profitable crop harvesting is possible only in the most productive apple woods of III and IV types. It is impossible to harvest a crop from apple wood type I, which grows on rocky stone soils of the southern slopes, in easily destroyed habitats of thin stands near the upper borders of the area (higher than 1600 m). Harvest is also not possible in apple wood type V, growing under the most extreme conditions for the species, where the soil is saturated with water. In highly productive apple woods, crop harvesting should be efficiently organized and practiced in accordance with license rules. In years of light cropping, concentrated harvesting from fruiting trees in small parts of the area should not be done because these trees are the sources of seed spread by animals for natural renewal. Special attention should be paid to groves of M. sieversii in Tarbagatai where this species is seen in “pure aspect” within its extreme northeastern extent in Kazakhstan. The arid hollows of Karatau with Pistacia vera L. deserves corresponding attention. It forms thin overgrowth in the very dry low mountains, built from motley-colored gypsum (uppertertiary and chalk) rocks. The extremely drought-resistant Pyrus regelii (Rehd.) provides a great potential for mountain forest reclamation, selection and rootstock purposes. The drought-resistant Amygdalus spinosissima grows in a very wide span of the mountains, from the lowest premountains at an altitude of 500 m, to the high mountains at a level of 1500 m, and occupies habitats among stones and on rocks. The preservation of endemic plants deserves special attention. Certain endemic plants grow inside fruit plant communities. These include the relict and endemic brushwood Atraphaxis muschketovii Krassh., which is seen in the central part of Zailijskei Alatau among the overgrowths of Rosa, Spiraea, Prunus, Celtis, and Acer semenovii. Separate bushy trees of M. sieversii and others grow here. In the narrow endemic of eastern Tien Shan, the brushwood Rhamnus songorica (Gontsch), spreads on

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slopes with northern orientations on the mountain ridge, Ketmentau, in the overgrowths of Acer semenovii and mixed with Prunus. With the aim of preserving a fully natural plant genepool, it is necessary to pay serious attention to a large number of different types, and especially to the many valuable races of apple in populations described above. Although in their evolution these types and races did not reach the rank of independent species, they are still of great scientific and practical interest. Forms of M. kirghisorum and M. sieversii, as well as cultivated orchards, cover the ancient and modern valleys and slopes in the gorges of Malaya and Bolshaya Almatinka, Koturbulak, Kamenskoe plateau. The landscape of suburban forest vegetation here enters into the green rest zone in the environs of Alma-Ata in Zailijskei Alatau. Therefore, the preservation of this vegetation should be intensified. Here and in analogous landscapes around cities and industrial centers with suburban, mountain vegetation, the organized economic measures for preservation include landscape forest-parks with model flower plots. A road path network could be created in the forest plots and around the orchards and plantations of beautifully flowering woody shrubs and grassy plants where bunches of flowers could be sold to guests. A study of the processes of natural renewal under such conditions showed that M. sieversii is successful in a natural way and does not need special preservation measures. Apple plots can be used as natural floristic displays in standard plots and as decorations for forest reserve country cottages. For the preservation and renewal of wild fruit plants in the mountains of Kazakhstan today, it is impossible to be limited only to natural methods, even if they are complete, because the process of afforestation has already gone too far. In many places, even natural plant communities are not preserved, that otherwise might have served as the initial material for their continued natural renewal. Therefore, for the preservation and renewal of fruit forest variety, it is necessary to use artificial plantations. Studying the race collections of chosen M. sieversii, M. kirghisorum, and M. niedzwetzkyana forms from natural populations revealed that their selection, genetic and production values and other useful peculiarities and characters are preserved in an artificial environment, even when they are propagated under other conditions. These forms are easily propagated by seeds or from vegetative parts of a mother plant. For this reason, the possibility of plant repatriation in natural habitats exists, in the regions of former distribution or in the places where species have suffered degradation. Not only the forms that have undergone trial in artificial environments, but also maternal material selected from excellent

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and normal trees out of seed plots in nature may provide genepool preservation and stock for wild plant recovery. An essential factor during the re-establishment of natural fruit components in the mountain flora of a given region is their placement in plantations. The diversity of wild fruit is colossal. The practical value of these forms is highly variable. They have differences in ecology and hereditary characters. Therefore, in establishing plantations, it is necessary to choose the most valuable and well-adapted seeds and planting material for the local conditions. In selecting species and race-forms, it is important to take into consideration the natural regularity of the distribution of natural vegetation in the composition of the land plots, as well their arrangement in the mountains. A good indicator of the ecological environment for plantations is the structure of the zonal distribution of wild fruit plants. Under the dry conditions of the Karatau ridge, where the main limiting factor is the lack of humidity, Malus sieversii with an understory of Prunus divaricata (Led.) forms thin stand overgrowths in the moist low parts of slopes with a northerly direction. The next belt of less moisture is occupied by the drought-resistant Pyrus regelii and the low-density thin overgrowths of Crataegus pontica, and on severe boghara, which include the driest and well-heated parts of the slopes, with loess and sometimes stone soils, Pictacia vera and droughtresistant Amygdalus spinossima form thin-stand overgrowths. There are analogous regularities with respect to the zonal distribution of natural fruit plants in other mountain regions, but as far as plant distribution on the slopes of the high mountain systems, the temperature regime and humidity are most important. Once forested, lands on the mountain slopes in the apple forest belt have great potential for sowing and planting valuable apple forms for reclamation and fruit plantations on former natural territory. On the treeless meadow-steppe plots, that border wild fruit forests, favorable conditions exist for artificial plantations and subsequent natural forest renewal. However, the degree of favorable ecological environment in the apple belt is far from equal in every area. As the altitude increases, vertical layering is clearly observed, expressed in the change of climatic conditions (from hot and dry to cool and wet), soils (from chernozem to mountain-forest and mountain-meadow) and vegetation (from desert to meadow and forest). Apple heat requirements during different phenological phases and during the entire vegetative period at different altitude zones, allowed us, with the help of agro-meteorological methods of analysis, to estimate the thermal resources of the mountain regions. It was determined that as a consequence of changing altitudes, radiation, the spectral composition of sunlight and other factors, the heat requirements of fruit plants also change. Differences in the climate and soil at comparatively small dis-

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tances create a strong pressure to distribute species and form fruit forest communities up across the vertical zones during natural reforestation. We distinguished the following zones on the territory of the mountain regions. The middle-mountain zone is one of the main zones for sowing and planting valuable forms of fruit trees. This zone is situated between the altitudes of 1100 to 1600 m, coinciding with the lower part of landscape belt of deciduous forests, forest-steppe and grassy meadow steppes. Soils are dark-colored mountain-forest, mountain-steppe, gray soils and leached chernozems. The average annual air temperature is 7.6°C; the average monthly temperature in the coldest month (January) is –4.8°C; and the average monthly temperature in the hottest month (July) is 20.2°C. The absolute minimum air temperature was –32 to 33°C, and the maximum, 35 to 37°C. The annual amount of precipitation is 800 to 850 mm. More than 70 percent falls from April to October, that is, during the period of plant vegetation growth. The frost-free period is 150 to 170 days in duration. Spring frosts cease at the end of April or beginning of May, and the first autumn frosts come in the first half of October. The absence of catastrophic winter freezes, sufficient rainfall and heat accumulation, without long hot summers and without the need for irrigation make it possible for the following plants to grow: very valuable forms of apples, pears, cherries, apricots, raspberry, strawberry and currant. In this zone, it is possible to grow the highly valued apple forms which ripen during the autumn-winter period and which have a large amount of biologically active substances. These fruits are suitable for use in storage and shipping (selections 28, 35, 177), and also as raw material for cider production. Early types, which ripen two to three weeks earlier, when grown in the next lower mountain zone, lose their importance and their use is reduced to a minimum. There had been great hopes for the growing of plantations of selected plum forms. But the plum varieties Manfor and Rodina when grown here in cultivated plantations are low yielding because of light freezing of fruit buds and therefore, they have absolutely no prospective future use. The low-mountain zone is situated between the altitudes of 850 and 1100 m and in a belt of feather grass-motley, grass bushy steppes on common mountain chernozems and on dark-chestnut soil alluvial cone. This zone has the most potential for land development having artificial fruit plantings. The average annual air temperature is 8.7°C. The absolute minimum temperature was –38°C. The total annual precipitation is 512 to 615 mm, most of which (364–431 mm) falls during the warm period. The duration of the frost-free period is 149 to 174 days. In respect to climate, this zone is entirely favorable for growing many valuable species and forms of fruit plants. Severe heat stress and the

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continental climate are characteristics of this zone in contrast to the middle-mountain zone. Here, winters are more severe and may cause serious damage to less cold resistant species and forms. However, severely cold winters have seldom been observed in recent years. The winter of 1951 was exceptionally unfavorable for wintering (the temperature fell to –34°C), and many cold-tender cultivated fruit plants died. Even under conditions of this severe winter, trees of wild apple all survived. In some patches, fruit buds were damaged, but these were exceptions. The trees with damaged fruit buds were situated at a lower altitude on land with unfavorable relief, and in lower places that were badly protected by other forest communities. This low-mountain zone may be used for planting resistant apple forms that ripen during the winter and autumn-winter periods. Here, the relative importance of pear may increase because the thermal conditions for pear are more favorable than in the middle-mountain zone. The best alycha and cherry forms, and the late-ripening forms of strawberry have good potential here. The growing of low yielding strawberry varieties, Rubesal and Lui Gotje, and the low quality variety, Komsomolka, and the late low yielding variety, Pozdnaya leopoldsgalskaya, should be replaced by the best strawberry forms, which are adapted to these local conditions. The same holds for the disease resistant raspberry varieties, Usanka and Marlboro, and to a number of other low yielding varieties. They could be replaced by the currently grown, selected forms of Ribes meyeri and the common red currant. When grown in this zone, the Middle-Russian apple varieties, Borovinka and Slavyanka, have absolutely no value. These varieties are vastly inferior to the selected forms of wild apple with respect to their main biological and economic features. The premountain zone is situated between the altitudes of 700 and 800 m and coincides with the belt of the feather grass-sheep’s fescue steppes on dark-chestnut soils. According to its climatic indexes, this zone is more continental than the low mountain zone. The average annual temperature is 7.8°C. The average monthly temperature in the coldest month (January) is –8.9°C, while the average monthly temperature in the hottest month (July) is 22.8°C. The absolute minimum air temperature is –41°C, and the maximum, 42°C. This premountain zone is less wet than the low-mountain zone. The average annual total precipitation is 500 mm, mostly falling in the summer period. The duration of the frost-free period is 160 to 180 days. The soil-climatic conditions of this zone are more favorable for the growing of apple forms that ripen in the autumn and summer. Lower parts of this zone are good for planting more warm-loving species of fruit plants. The apple, ‘Grushovka Moskovskaya’, grows in the forest orchards of

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this zone. It is a good sort for the middle belt of Russia. This variety appeared to be unproductive with fruits of fair flavor. For some years, the trees of this variety continued to grow until late-autumn frosts. Considering all of its biological and cropping features and characters, Grushovka is inferior to the valuable wild apple forms. The valuable forms of pear, alycha, cherry, and strawberry show great potential for development in the premountain zone. The predominately plain zone is situated at the altitudes of 600 to 750 m in the desert-steppe landscape zone, which has wormwood, sheep’s fescue, and grass vegetation. Meadow-chestnut soils and common chernozem prevail. In comparison with other zones, the predominately plain zone is more continental and dry. The average annual air temperature is 6.6 to 7.7°C. The absolute minimum temperature was –43°C, and the maximum, 42°C. Severe heat stress, very low humidity, sharp temperature fluctuations, severe winters, and a small amount of precipitation are less favorable environmental conditions for the mass development of stands than are those found within the premountain and low-mountain zones. The plantings of summer and early ripening apple forms, winter-hardy varieties of strawberry, and summer and autumn pear forms are possible here. The planting of winter pear is not advisable. The more resistant forms of alycha, cherry and forms of early and middle ripening strawberry are of value here. The plain zone includes separate oases between a desert of sheep’s fescue, wormwood, and saline vegetation. This zone is situated at altitudes below 600 m. Soils are cheronzem, takyr, meadow-bog, solonchalks, and hilly sands. The average annual air temperature is 8.8°C. The absolute minimum temperature was –44°C, and the maximum, 44°C. The duration of the frost-free period is 150 to 160 days. The precipitation is half that in the predominately plain zone. The severe conditions in this zone make any mass artificial plantings of many fruit-berry plants very difficult. It is possible to plant selected frost-drought-resistant plants on a limited scale. The intermountain-valley zone is situated at the mid-point of the river, Ili, between the altitudes of 500 to 900 m, in desert and desertsteppe zones with sheep’s fescue-wormwood-ephimer and wormwoodsaline vegetation on chernozem, meadow-chernozem and partially light-chestnut soils. This zone is protected from the influence of northern winds by the Dzhungarskei Alatau ridge, and as a consequence, favorable climatic conditions are formed here. Among the multiple types of soil, light chernozems and meadow-chernozems are most often seen. The frost-free period is 170 to 180 days. The precipitation is 125 to 198 mm. Considering the climatic indexes of great heat accumulation and an

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abundance of sunny days, and especially in regard to the total heat unit accumulation, it is possible to plant fruit plants that require much heat. Sowing seeds which have been collected in seed wood stands from excellent and normal trees, and planting of economically valuable clones of apple stands on adjacent territories are the most rational means for the reintroduction of forests, which have been removed as a result of human economic activity. These regional mountain territories with their zonal distribution of fruit plantations can be the basis of forest reproduction projects. IX. CONCLUSION The natural fruit forests of Kazakhstan contain promising material with many useful qualities for botanical, geographic, and genetic selection. In this sense, the significance of these genetic stocks reaches far beyond the limits of any narrow regional interest. Populations of Malus species in the mountains of Kazakhstan, especially the genetic stock of the phenotypically plastic varieties, are necessary to maintain for selection and for genetic investigations. Rational utilization, that is, the protection and exchange of genetic material of this region’s vegetation, is becoming increasingly significant as a solution to major international biological problems of this time. In natural populations, practically an unlimited number of plants grow that are valuable for the introduction and selection of fruit-plant forms. Until this time, such plants have not been objects of study and they are still absent from cultivation. Apple tree forests greatly vary in floristic composition and economic value, as well as in the natural conditions and habitats that they occupy. In accordance with the natural laws of dispersion, an ecological type of apple tree has appeared, adapted to the moderately cold climate of the forest-steppe region of the Northern Hemisphere and characteristic of the Eurasian vegetation formation. The diversity of contemporary natural fruit communities is concentrated in the mountains at elevations between 800 to 2000 m above sea level; in forest-meadow, forest-steppe, and steppe zones; in deciduous forests, at the edge of forests, in combination with shrubbery, and as a composition of motley grass steppes or large herbaceous meadows. In considerable number and often of great quality, apple trees inhabit extensive natural areas. In addition, with their close conformity to the natural law of specific feature repetition, a rationale is provided for considering them a forest type in classification (Aestilignosa). The apple forests of Kazakhstan, as relict forest types, appear as part of the specific composition of the landscape, are tightly linked with the

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natural complex of biogeocenosis and maintain its balance. In the areas that they occupy, they establish an entangled, interrelated complex with various aspects of the ecological community, including, undoubtedly, not only those of the tree and associated plants, but the environment as a whole. The scattered natural habitats of apple trees, their presence in different soil and climatic conditions, and the complication of their organizational structures determine the considerable variety of habitat types, including: very dry meadowsweet, dogrose, apple; dry herbaceous, brush with hawthorn; fresh herbaceous, brush with hawthorn and aspen; and fresh-grass, mixed grasses, and moist herbs. From the types of apple known in Kazakhstan, mainly M. sieversii forms the apple forests of mesophylic autotrophic trees. This species can be considered as a xero-mesoform exomorph. Its xeroformic traits are most obvious in very dry and dry conditions of growth. In cool and especially moist conditions of growth, to a considerable degree, M. sieversii throws off its “xeromorphic armour” and its morphological features come nearer to those of the mesophytic species, M. kirghisorum. Fruit trees establish their own local microclimates, which are considerably different from the microclimates in adjoining treeless territories. The layered structures of apple trees promote the redistribution of light and atmospheric precipitation. Light goes through a series of organic green filters, which absorb most of the physiologically active radiation. Atmospheric moisture, on the other hand, due to flow and evaporation on the multiple layered canopy, gets to the soil under the apple tree in a lesser amount than on open ground. In this way, apple trees protect the soil from overheating and erosion, caused by the surface flow of atmospheric precipitation. In the heart of apple forests the humidity rises and the air temperature lowers, positively influencing all ecological functions during conditions of insufficient moisture. The resources of heat and moisture in the middle range mountains (1100 to 1500 m) are particularly favorable for the vegetative growth of apple forms there, those with summer, fall and fall-winter ripening fruit. At heights above 1600 to 1800 m, in the majority of years, the only forms that ripen have a very short period of vegetative growth and early fruit ripening. In the understory of the forest, not only do climatic conditions change, but also the soil conditions change. Investigations reveal the interrelationships of the processes that occur in the understory of apple tree forests, and confirm the enrichment in fertility of mountain chernozem and dark-chestnut soils that naturally occur. A large mass of organic matter acting on the surface of soils under fruit forests, promotes its enrichment with humus and nutrient matter and improves the water-physical

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characteristics of the soil, establishing conditions for an abundant population of soil fauna and microflora. Favorable hydrothermal conditions in ecosites promote the intensive decomposition of fallen litter and the leaching of nutrient matter in the soil. These processes, in turn, strengthen the intensity of the biological cycle of organic matter and nitrogen in the understory soil. Rooting out the forest and plowing up virgin soil abruptly change all the processes of the biological cycle within an apple forest. The entry of vegetative residues into the soil is reduced, the biological, water-physical, and chemical characteristics of the soil decline and the microclimate deteriorates. Malus sieversii is well adapted to mountain leached chernozem and under these conditions, trees of this species will create root systems which are distinguished by extreme thickness, sufficient depth and numerous horizontal branches. The structures of the root system make these apple trees well adapted to mountain slopes in the southeastern mountain ridges of Kazakhstan. The normal natural renewal of apple trees takes place by seed and by vegetative means. Rootshoot, airshoot and adventitious root formation and self-anastomosis provide examples of autovegetative renewal in a natural population of trees. An apple population, as a relatively independent association of trees occupying a definite area, consists of trees of seed and vegetative origin of various ages. Within populations, processes of natural renewal and hybridization occur. Having close mutual contact within populations, the roots of peripheral trees grow together, promoting the formation of large roots. All this anchors populations to their territory and helps them resist during the struggle with other competing species. The process of selfthinning within the population is regulated by the density of the stand, according to the age and developmental period of individual trees and their active growth and intensive differentiation. Naturally determined changes in the characteristics of growth and productivity are observed in trees that comprise populations of M. sieversii. The life cycles of populations can be divided into growth periods based on these changes; beginning with their origin from seed or vegetative offshoot through the juvenile period, to fruiting, and dying adult trees. The presence of sufficient re-growth, optimal growth and productivity for each of the distinctive age groups are testaments to the life force of populations of M. sieversii and to the absence of species degradation. In populations of wild apple numerous forms grow which are sharply distinguished from one another by a series of traits and characteristics: the duration of their vegetative period; their general habit; their winter resistance; their disease resistance; their productivity; and the pomo-

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logical characteristics of their fruit. Polymorphism of apple populations is a consequence of the breadth of the species natural habitat, the great variety of climatic, soil, and ecological growth conditions, and the numerous crossings between species and their various hybrids. Introgression of hybrids and spontaneously arising mutations, which promote further speciation of Malus in versatile mountain conditions, play a great role in the heterogeneity of dynamic populations. Evolving in complex, ecological conditions of mountains for millenniums, the wild apple species of Kazakhstan possess valuable hereditary variability, which is the foundation for further selection. A narrow connection exists between various populations of apple and the environmental conditions in various parts of their growing regions, where they have developed historically. The descendants of trees from distinctive areas and from vertical zones are qualitatively different, and this fact must form the basis for dividing forest culture districts in the mountain territory for the reintroduction of populations. Highly productive and stable individuals are produced in populations of apple forests, due to diverse processes of interbreeding and individual selection. The significance of these individuals is especially great for the improvement in fruit population productivity. They are maternal trees for selection in the establishment of forest plantations of apples. As vegetatively reproducing plants, natural apple tree hybrids can play a special role, since economically important forms can be easily cloned and successfully propagated at an industrial scale. Research reveals that first generation apple hybrids manifest heterozygotic effects and possess special value. Hybrids can be successfully utilized as seed sources as well. The utilization of the intraspecific diversity of Malus is secure and may serve as a means to raise the productivity of the forest and to contribute towards forming apple ecosystems in the natural environment. For an increase in the productivity of deforested land in the mountains, it is necessary to significantly increase the density of green fruit plantations. This is especially important for places that are poor in forest regions, as Kazakhstan, where wooded territory, including saxaul trees (Haloxylon, Chenopodiaceae), makes up only 3.3 percent in all. For introduction into culture, it is rational to sow and plant material from the biologically most valuable, stable and productive populations of apple. Apple trees are easily accessible in the low mountains, where they are subjected to negative influences caused by the economic activities of people over a prolonged period of time. Damaged apple forests demand reconstruction by means of the creation of new artificial apple ecosystems with higher biological productivity. By sowing and planting high quality forms of Malus seed in plantation, the production of bio-

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mass will increase as observed in other agricultural plantations. The greatest success may be attained by the placement of fruit plants in situations similar to the natural conditions of the vertical zones in the district mountain territory as described in this chapter. As shown by research, forms of apple under other conditions preserve their economically important genetic traits and characteristics. Their introduction into culture brings long-term, high quality productivity and stability to the species of fruit plants of highest importance for people. The colossal saturation of intraspecies apple diversity in the midrange mountains is the result of contact between populations of M. sieversii and M. kirghisorum, with the greatest number of reproductive trees in the most favorable sites for apples. Dominant mutations of highyielding forms of trees with valuable pomological traits of fruit present a special interest for selection using between populational crosses. A prospective direction in the selection of forms is also intra-populational, based on the high degree of isolation between populations that occurs on adjacent territories, wherein appear responses of various genotypes to the influence of local conditions of the environment. Here as a result of the shift in frequency of distinct genotypes towards greater adaptation of the population to the concrete conditions of the environment, one can expect the appearance of local adaptations and the formation of a specific genetic base in the population. In this sense, the high mountain populations of Zailijskei, Dzhungarskei, and Tarbagatai are of value for the selection of early-ripening and precocious winter-hardy tree forms. The extreme conditions of the high mountains reveal all the potential possibilities of the species for adaptation to severe habitats. Populations there have the ability to preserve and secure the territory by means of fruit bearing and natural seed and vegetative renewal. High mountain apple forests, formed under the influence of low temperatures, possess a shortened vegetative period. The apple populations growing in the extreme lower part of high altitude mark (below 900 meters) of the steppe zone also deserve attention in the sense of being early fruitbearing and drought resistant. Numerous and diverse biological, chemical, and industrial forms of apple grow in the wild fruit forests of Kazakhstan. Their fruits are characterized by a higher content of biologically active substances and microelements. Based on these indices, they greatly excel over the cultivated sorts. We succeeded in identifying new selections for introduction to cultivation, distinguished by regular productivity with good storage quality. Fruit of the isolated types and forms are excellent raw material for distillation, canning, and the confectionery industry. For certain kinds of

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production, their utility depends on their flavor. Recommendations are given for the industrial production of cider, juices, and calvados, after the new technology developed in Kazakhstan. An insufficient study of the ecological role and biological features of apple trees has brought about a disturbance between the established interrelations of the apple forest and human living conditions. In particular, there is ignorance concerning the diversity of forms, the vegetative reproduction of apples, and the practice of forestry for natural populations of wild apples. As a result of the ineffective management of forestry in the past (reconstruction of fruit ecosystems with the goal of creating forest gardens, repeated grafting, chopping out, with removal other species of trees and bushes, etc. from the apple forests), a decrease in area of the apple forests, disturbance and destruction of tens of thousands of valuable trees, and the degradation of plant communities have occurred. The protection of apple forests may be successfully accomplished by preserving the vitality of populations and by creating conditions for the development of their characteristic ecosystems. This question is part of the general problem concerning the rational utilization, protection, and enrichment of all vegetative cover. Answers should include the preservation of elements of the landscape and the vegetative associations on which apple forest existence depends. In a series of places in the forest free zone, adjacent to wild ecosystems, it is advisable to sow and plant valuable forms of wild apple. The diversity of forms of the populations of wild fruit plants, their durability, their high decorative quality, and their fruit with high levels of biologically active substances show great potential for ornamental plantings, forestry, and the establishment of resistant fruit plantations. LITERATURE CITED Abeev, A. C., and N. Y. Bondarev. 1967. Grafting of a wild apple by green cuttings of cultural varieties. Forestry 1. Abolin, P. I. 1930. From eremic steppes of near Balkhash region up to snow tops of Khantengri. Trans. Inst. Soil Sci. Geoactivities. SA State Univ. The geobotanic and soil description of a southern part of the Alma-Ata district of Kaz. SSR 5(1). Kazakhstan series. Leningrad. Afanaseva, E. A. 1966. A node of potent black soils under herbaceous and arboreal coenosises. Soil Sci. 6. Aiton, W. 1789. Hortus Kewensis or a catalogue of the plants 2. London. Akimushkin, V. G., P. A. Stepanov, and F. S. Salynskij. 1935. Wild fruit forests and their usage. Voronezh. Aleksandrov, A. D. 1960. Horticulture in France. Timiryazev Agr. Acad. Rep. 53. Aliev, M. 1965. Wild fruit and nuts of Azerbaijan. Forestry 10.

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Andreeva, N. A. 1953. Fluorometric method of definition of folic acid and some data about its distribution. Biochemistry 18(5). Apoyan, L. A. 1964. Root system of seedlings of different ecotypes of wild and some other species of apple in nursery in conditions of Ararat valley of Armenia, SSR. Trans. Arm. Sci. Res. Inst. Viticulture. Wine Production and Fruit-farming 6 and 7. Araratyan, A. G. 1940. To the problem on organization of fruit forests in Armenian SSR. Bul. Bot. Garden 2. Arasimovich, V. V. 1948. A problem of sugar. Biochem. Cultivated Plants 8. Arasimovich, V. V., and L. A. Vasilieva. 1960. Transformation of carbohydrates in apples in storage. Inform. Moldova Filial Acad. Sci. USSR 2. Arasimovich, V. V. 1962. Biochemistry of cultivated plants of Moldova. Biochemistry of an apple 1. Arnold, F. K. 1891. Russian forest 11. St. Petersburg. Ascherson, P., and P. Graebner. 1906–1910. Synopsis der Mitteleuropäischen Flora VI. Leipzig. Bd. Avsaragov, A. Kh. 1965. Protection of wild fruit forests. Agr. Prod. Northern Caucasus Central Black Soil District 7. Azimov, Kh. 1973. The most valuable species of wild fruit trees of Kashkadarya river basin. Scientific. Trans. Tashkent State Univ. 439. Bailey, L. H. 1916. Nomenclatorial transfers. Rhodora. Bailey, L. H. 1949. Manual of cultivated plants. New York. Baranova, E. A. 1951. Regularity of formation of adventitious roots in plants. Trans. Main Bot. Garden 2, Moscow. Batalin, A. 1893. Notae de plantis asiaticis. Batygin, N. F. 1950. About routes of development of wild fruit tree forests. Res. Notes Leningrad State Univ. Biol. Ser. 23. Belozerskij, A. I. 1951. Practical guidance on plant biochemistry. Sov. Sci. Beskaravajnyj, M. M. 1956. Formation of biogroups in pine plantations of Kamyshinsky forest irrigation-support station. Agrobiology 1. Beskaravajnyj, M. M. 1958. The new facts of accretion of different tree species in conditions of Kamyshinskij region. Agrobiology 6. Bezpoludenov, I. A. 1954. Soils of wild fruit thickets in foothills of Zailijskei Alatau. Extended Abstr. Ph.D. Thesis, Alma-Ata. Bezsonov, A. I. 1910. Soils of parts of Djarkentskij and Vernenskij regions of Semirechenskij district. Trans. Soil-botanic Expedition of Petersburg Univ. for 1908, 6, St. Petersburg. Bezzubov, A. D. 1949. Chemico-technological and economic characteristics of production of fruit forests of Southern Kirghizia and their usage. 1, Moscow, Leningrad. Blaja, D., and I. Ivan. 1960. Variabilatatea formelor de mâr, pâr si cires sâlbaltic din jurul Bistritei si valoarea lor pentru selectie. Comunicari de Botanicu (1957–1959). Bucuresti. Bogoyavlenskaya, S. V. 1962. Purchase and processing of wild fruits and berries. Moscow. Boissier, E. 1872. Flora orientalis. Borkhausen, M. B. 1803. Theoretisches praktisches Handbuch der Forstbotanik und Forsttechnologie. Borodina, N. A., and V. I. Nekrasov. 1966. Trees and bushes of the USSR. Moscow. Brandis, D. 1874. The forest flora of north-west and central India. London. Branke, Yu. V. 1935. Chemistry of wild fruits and berries of a Far East region. Vladivostok. Brimble, L. J. F. 1946. Trees in Britain. Wild ornamental and economic, and some relatives in other lands. London.

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Tsitsin, N. V. 1972. About development of search, test and introduction in cultivation of economically valuable plants of natural flora. Bul. Main Bot. Garden Acad. Sci. USSR 83. Turbin, N. N. 1971. Genetics of a heterosis and methods of plant breeding on a combining ability. In: Genetic fundamentals of plant breeding. Moscow. Turchin, F. V., M. A. Guminskaya, and E. G. Klyshevskaya. 1955. Research of nitrogenous metabolism of plants with application of an isotope N15. Soviet Plant Physiol. 2(1). Turkin, N. A. 1935. Assimilation of wild fruit species. Fruit-vegetable Farming 6. Turskij, M. K. 1912. Sylviculture. 6th ed. Moscow. Tyurin, I. V. 1930. To a problem on genesis and classification forest-steppe and forest soils. Science Notes (Kazan). 45(3, 4). Uglitskikh, A. N. 1932. Elements of a technique of morphological identification of wild apple trees. Krasnodar. Unchiev, N. D. 1947. To use wide wild fruit gardens. Makhachkala. Uranov, A. A. 1974. Plants and environment. In: Life of plants 1. Usatyj, P. P. 1959. Usage of wild apples and pears in Krasnodar region. In: Proc. Scientific Conf. Post-Graduate Students and Young Scientific Employees Dedicated to XXI CPSU Congr. Leningrad. Usenko, N. V. 1953. Fruit and berry plants of forests of Far East. Khabarovsk. Utekhin, V. D. 1964. Materials on intensity of a fructification of trees and bushes in reservation Aksu-Dzhabagly (Northwest Tien Shan). In: Geography of a fructification of forest wood species, bushes and berries. Moscow. Uvarov, F. V. 1949. Fruit-and-berry trees and bushes for field-protective afforestation of Kuibyshev district. Garden & Kitchen Garden 3. Van Eseltine, G. P. 1933. Notes on the species of apples. The American crab apples. Tech. Bul. N. Y. State Agr. Expt. Sta. 208. Vashchenko, D. L. 1958. Forest gardens. Agriculture of Caucasus 7. Vasilichenko, I. T. 1948. Wild Gissar apple as a material for breeding and hybridization. Proc. Tadjik Filial Acad. Sci. USSR 6. Vasilichenko, I. T. 1952. Wild apple species in Middle Asia. In: Proc. First All-Union Conf. of Botanists and Breeders, March 24–27, 1950. 2, Moscow, Leningrad. Vasilichenko, I. T. 1959. To knowledge of the east-Asian apple species (Malus Mill). Botanical materials of a Herbarium of V. L. Komarov Botanical institute. Acad. Sci. USSR. 19. Moscow, Leningrad. Vasilichenko, I. T. 1963. New for cultivation apple species. Moscow, Leningrad. Vasilichenko, I. T. 1971. Nature protection in republics of Middle Asia and Kazakhstan. In: Problems of preservation of botanical objects. Leningrad. Vasilyev, A. V. 1938. Wild fruit and alimentary tree species of Abkhazia. Trans. Inst. Abkhazia Cultures Georgian Filial Acad. Sci. USSR 5. Vasilyev, V. F. 1932. The review of wild and fruit trees run wild and bushes of Crimea. Trans. Appl. Bot. Gen. Breed. 27(1). Vasilyev, L. A. 1959. Accumulation of carbohydrates at apple maturation in miscellaneous zones of fruit-farming of Moldova. Biochemistry of fruits and vegetables. Collection 5. Vavilov, N. I. 1927. Geographic regularity in distribution of genes of cultivated plants. Trans. Appl. Bot. Gen. Breed. 17(3). Vavilov, N. I. 1931. The wild relatives of fruit trees of the Asian part of the USSR and Caucasus and the problem of the origin of fruit trees. Trans. Appl. Bot. Gen. Breed. 26(3). Vekhov, N. K. 1932. Vegetative propagation of woody and bushy plants. Leningrad. Vecher, A. S., and V. N. Bukin. 1940. A biochemistry of apples. In: A biochemistry of cultivated plants. Moscow, Leningrad.

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Vigorov, L. I. 1961. Vitamin content of apples of Middle Ural and perspectives of apples as a source of vitamins. In: Trans. First Seminar on Bioactive Compounds of Fruits and Berries. Sverdlovsk. Vigorov, L. I. 1964. Bioactive compounds of fruit-berry plants and primary goals of their researches. In : Trans. Second Seminar on Bioactive Compounds of Fruits and Berries. Sverdlovsk. Vigorov, L. I. 1968a. Ray protective compounds of apples. In: Trans. Third Seminar on Bioactive Compounds of Fruits and Berries. Sverdlovsk. Vigorov, L. I. 1968b. Vitamin content of apple fruits of different species in connection to peculiarities of cultivated varieties. Trans. Inst. Plant and Animal Ecol. 62. Viktorovsky, G. P. 1935. The wild apple Malus dasyphylla Borkh. In: Fruit Trees of Middle Tadjikistan. Leningrad. Vilensky, D. G. 1946. Soils of fruit forests of Fergana mountain. Newsletter Moscow State Univ. 3 and 4. Vinogradova-Nikitina, P. Z. 1929. Fruit and alimentary trees of Transcaucasus forests. Trans. Appl. Bot. Gen. Breed. 22(3). Vintergoller, B. A. 1964. About contemporary state of the art and strengthening of preservation of stands of wild apple. Inform. Acad. Sci. Kaz. SSR Ser. Biol. 2. Vintergoller, B. A. 1965. Foliaceous forests of northern Tien Shan (geobotanic essay). Extended Abstr. of Ph.D. Thesis. Alma-Ata. Voejkov, A. D.1909. An origin of fruit trees varieties. St. Petersburg. Vorobiev, D. P. 1935. A vegetative coverage of Southern Sikhote-Alin and wild fruit and berry plants in this region. Trans. Far East Filial Acad. Sci. USSR. 1 Moscow, Leningrad. Vorobieva, M. G. 1966. An apple of the Kirghiz. Trans. Sary-Chelik Reservation 2. Vysotsky, G. N. 1952. About hydrological and meteorological influence of a forest. Moscow, Leningrad. Watson, P. W. 1825. Dendrologia Britannica. 1. London. Weir, W. 1940. Soil Science. Its principles and practice (rev. ed.). Chicago. Wenzig, Th. 1874. Pomariae Lindley. Linnaea 4. Willdenov, C. G. 1794. Pyrus prunifolia. Phytographia, Erlangae 1. Williams, E. B., D. F. Dayton, and J. R. Shay. 1966. Allelic genes in Malus for resistance to Venturia inaequalis. Proc. Am. Soc. Hort. Sci. 88:52–56. Williams, W. 1968. Genetical fundamentals and plant breeding. Moscow. Wood, A. 1872. Classbook of botany. New York. Yunovidov, A. P. 1951. Accretion of root systems in a forest. Agrobiology 4. Yuzepchuk, S. V. 1939. A genera of an apple Malus. In: Flora USSR 9. Moscow. Zabel, H. 1903. Handbuch der Laubholzkunde. Berlin. Zakotin, V. S., and A. R. Atanasov. 1972. About interrelations between growth and development of above-ground and root systems of an apple in an annual cycle. Inform. TAA. 5. Zapryagaeva, V. I. 1949. Experience of non-irrigated fruit-farming and silviculture in mountainous Tadjikistan. Trans. Tadjik Filial Acad. Sci. USSR 17. Zapryagaeva, V. I. 1964. Wild fruit trees of Tadjikistan. Moscow, Leningrad. Zelensky, O. V., and N. I. Vavilov. 1966. Problem of mountainous agriculture. Moscow. Zhavoronkov, P. A. 1938. The Siberian berry-like apple and breeding of winter hardy varieties. Moscow. Zhukovsky, P. A. 1971. Cultivated plants and their ancestors. Leningrad. Zhuravskij, A. I. 1914. To a problem on best apple rootstocks (wildings). Progressive Gardening and Vegeculture 5. Zonn, S. V. 1954. Influence of a forest on soils. Moscow.

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Zonn, S. V., and A. K. Aleshina. 1953. To a problem on decomposing of abscised leaves of oak forests and interaction of its ash components with soils. Trans. Inst. Forest Acad. Sci. USSR 12. Zonn, S. V., and E. A. Kuzmina. 1960. Influence of coniferous and foliaceous species on physical characteristics and water relationships of leached black soils. Trans. Silvicult. Lab. 1. Zonn, S. V., and E. A. Kuzmina. 1964. Materials to the conjugate characteristic of a water regime of physico-chemical properties of turf-podsol soils under oak, fir-tree and best wood forests. In: Stationary biogeocenotic research in a southern subzone of taiga. Moscow. Zonn, S. V., V. N. Mina, and N. D. Varlygin. 1953. A water region of soils under forests and field protective plantations in steppe. Transa. Inst. Forest. 12. Zyrin, N. G. 1962. The definition of the gross contents of trace elements in soils by a spectral method. Moscow.

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3 The Wild Fruit and Nut Plants of Kazakhstan* A. D. Dzhangaliev, T. N. Salova, and P. M. Turekhanova Kazakhstan Academy of Science Interbranch Laboratory for the Protection of Germplasm Main Botanical Garden Almaty, Republic of Kazakhstan

I. INTRODUCTION II. POME FRUITS A. Apple 1. Sievers Apple [Malus sieversii (Ledeb.) M. Roem.] 2. Kirghiz Apple (Malus kirghisorum Al. et An. Theod.) 3. Niedzvetzky Apple (Malus niedzwetzkyana Dieck.) B. Pear C. Cotoneaster D. Hawthorn 1. Alma-Ata Hawthorn (Crataegus almaatensis Pojark.) 2. Doubtful Hawthorn (Crataegus ambigua C. A. Mey. ex A. Beck.) 3. Altai Hawthorn (Crataegus cholorocarpa Lenne et C. Koch.) 4. Ponti Hawthorn (Crataegus pontica C. Koch.) 5. Blood-red Hawthorn (Crataegus sanguinea Pall.) 6. Dzhungarskei Hawthorn (Crataegus songarica C. Koch.) 7. Turkestan Hawthorn (Crataegus turkestanica Pojark.) E. Mountain Ash 1. Persian Mountain Ash (Sorbus persica Hedl.) 2. Siberian Mountain Ash (Sorbus sibirica Hedl.) 3. Tien-Shan Mountain Ash (Sorbus tianschanica Rupr.)

*Originally translated from Russian by I. N. Rutkovskaya. Supported by grant SCA no. 580500-4-F100, U.S. Department of Agriculture, Agricultural Research Service.

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III. STONE FRUITS A. Apricot B. Cherry 1. Tart Cherry (Cerasus) 2. Bird or Sweet Cherry [Padus avium Mill.; syn. Prunus avium (L.) L.] 3. Mahaleb Cherry [Padellus mahaleb (L.) Vass.; syn. Prunus mahaleb L.] C. Plum 1. Sogdijskaya Plum, Alycha (Prunus sogdiana Vass.) 2. Spiny Plum, Blackthorn (Prunus spinosa L.) IV. SMALL AND VINE FRUITS A. Barberry 1. Bykovskiy Barberry (Berberis bykovianas Pavl.) 2. Iliyskei Barberry (Berberis iliensis M. Pop.) 3. Integrifolious Barberry (Berberis integerrima Bunge) 4. Karkara Barberry (Berberis karkaralensis Korn. et Potnap.) 5. Kashgar Barberry (Berberis kaschgarica Rupr.) 6. Monetary Barberry (Berberis nummularia Bunge) 7. Oblong Barberry (Berberis oblonga Schneid.) 8. Siberian Barberry (Berberis sibirica Pall.) 9. Spherical-fruited Barberry (Berberis sphaerocarpa Kar. et Kir.) B. Currant 1. High Currant (Ribes altissimum Turcz. ex Pojark.) 2. Dark-Purple Currant (Ribes atropurpureum C.A. Mey.) 3. Odorous Currant (Ribes graveolens Bunge) 4. Different-pubescent Currant (Ribes heterotrichum C.A.M.) 5. Hispid Currant [Ribes hispidulum (Jancz.) Pojark.] 6. Yanchevskiy Currant (Ribes janczewskii Pojark.) 7. Meyer’s Currant (Ribes meyeri Maxim.) 8. Black Currant (Ribes nigrum L.) 9. Rock Currant (Ribes saxatile Pall.) 10. Turbinate Currant (Ribes turbinatum Pojark.) 11. Red Currant (Ribes vulgare Lam.) C. Gooseberry D. Grape E. Raspberry and Blackberry 1. Common Raspberry (Rubus idaeus L.) 2. Sakhalin Raspberry (Rubus sachalinensis Leul.) 3. Blackberry (Rubus caesius L.) 4. Stone Berry (Rubus saxatilis L.) F. Strawberry 1. Alpine Strawberry (Fragaria vesca L.) 2. Green Strawberry (Fragaria viridis Duch.) G. Vacciniums 1. Bilberry (Vaccinium myrtillus L.) 2. Bog Bilberry (Vaccinium uliginosum L.) 3. Lingonberry (Foxberry or Cowberry) (Vaccinium vitis-idaea L.) 4. Microcarpous Cranberry (Vaccinium microcarpus L.) 5. Quadripetalous Cranberry (Vaccinium palustris L.)

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V. OTHER FRUITS A. Elderberry B. Honeysuckle C. Mulberry D. Oleaster 1. Narrow-leafed Oleaster, Russian Olive (Elaeagnus angustifolia L.) 2. Sharp-fruited Oleaster (Elaeagnus oxycarpa Schlecht.) 3. Turkmen Oleaster (Elaeagnus turcomanica N. Kozl.) E. Rose 1. Acicular Rose (Rosa acicularis Lindl.) 2. Albert Rose (Rosa alberti Regel.) 3. Begger Rose (Rosa beggerana Schrenk) 4. Dog Rose (Rosa canina L.) 5. Corymbose Rose (Rosa corymbifera Borkh.) 6. Dzharkent Rose (Rosa dsharkenti (Chrshan.) 7. Fedchenkovskiy Rose (Rosa fedtschenkoana Regel) 8. Glabrous-leafed Rose (Rosa glabrifolia C.A. Mey. ex Rupr.) 9. Hissar Rose (Rosa hissarica Slob.) 10. Kokand Rose [Rosa Kokanica (Regel) Regel et Juz.] 11. Lax Rose (Rosa laxa Retz.) 12. Samarkand Rose (Rosa maracandica Bunge) 13. May Rose (Rosa majalis Herrm.) 14. Dwarf Rose (Rosa nanothamnus Bouleng.) 15. Acute-prickly Rose (Rosa oxyacantha Bieb.) 16. Pavlov Rose (Rosa pavlovii Chrshan.) 17. Wide-prickly Rose (Rosa platyacantha Schrenk) 18. Pedate-color Rose (Rosa potentilliflora Chrshan. et M. Pop.) 19. Schrenkovskiy Rose (Rosa schrenkiana Crep.) 20. Silverheim Rose (Illiyskei) (Rosa Silverhjelmii Schrenk) 21. Spinose Rose (Rosa spinosissima L.) F. Sea Buckthorn G. Viburnum VI. NUTS A. Almond 1. Ledebour Almond (Amygdalus ledebouriana Schle.; syn. Prunus ledebouriana) 2. Petunnikov Almond (Amygdalus petunnikovii Litv.; syn. Prunus petunnikovii Litv.) 3. Thorny Almond [Amygdalus spinosissima Bunge; syn. Prunus spinosissima (Bge.) Franch.] 4. Dwarf Almond (Amygdalus nana L.; syn. Prunus nana Stokes) 5. Common Almond [Amygdalus communis L.; syn. Prunus communis (L.) Arcangeli] B. Hazelnut C. Pistachio D. Siberian Pine E. Walnut LITERATURE CITED

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I. INTRODUCTION The problem of preserving, multiplying, and rational use of forest resources of the Kazakh Republic (Fig. 3.1) acquires especial significance in light of the “Appeal to The People of Kazakhstan 2030” by President N. A. Nazarbaev, who proclaimed: “The symbol of Kazakhstan should not be deserts but forests and orchards.” The genetic resources of fruit forests in Kazakhstan are rich (Table 3.1). The wild fruit forests of the Republic have both biological and economic value for fruit production and play an important ecological role in the low-mountain and middlemountain zones of Tien-Shan, the Altai, Tarbagatai, Dzhungarskei Alatau, and Karatau. These forests, distributed along mountain slopes at 1000 to 1800 m above sea level, play water-regulating, soil-protecting, and climate-regulating roles, that form a self-regulating natural system. The direct use of these forests by man is basic; however, much more important is the inexhaustible wealth inherent in the genepool of the polymorphic species. The discovery of the genetic potential of wild fruits of Kazakhstan was realized through studies of intraspecific polymorphism. A species is a heterogenic, genetic system, which must be understood in order to learn how to preserve it. Kazakhstan occupies a vast territory in Central Asia where two great floristic regions, boreal and Mediterranean adjoin. Here, almost all natural floristic zones of the world are present; from dry subtropics to alpine tundras, and the flora of higher plants numbers more than 6000 species including endemic and relict species. The Kazakhstan territory is also the northern extremity of the Central Asian center of origin of cultivated plants which includes 38 important agricultural crops. In the Kazakhstan flora, the progenitors of 17 of these crop species, and also related species, include 157 species. Finally, 20 percent of the world’s cereals, more than 20 percent of the vegetable and spice plants, and 90 percent of the major temperate-zone fruit crops are found here. This confirms that Kazakhstan is the center of origin of major fruit crops. As Kazakhstan occupies the northern regions of Vavilov’s South Western Asian Center, the wild fruits here possess the genetic bases for stressresistance including resistance to diseases, pests, and wind. At present, the genetic resources of wild fruits in Kazakhstan are recognized as a world source of resistant genes for breeding of new fruit cultivars. On the slopes of the Kazakhstan Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kirghiz, Terskei, Kungei, Talasskei Alatau, Ketmentau, and Karatau more than 125 species grow, including 31 genera and 12 families, among which are relict plants, endemic species, and fragments of ancient landscapes, such as nut, apple and apricot stands. Natural fruit forests, an

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The main mountain systems of Kazakhstan and border areas with Uzbekistan and Kyrgyzstan.

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Economically valuable plant species growing in fruit forests of Kazakhstan.

Achillea filipendulina Lam. Achillea millefolium L. Achillta micrantha Willd. Aconitum leucostomum L. Allochrusa gypsophiloides (Regel) Schischk. Amygdalus ledebouriana Schlecht. Archangelica decurrens Ldb. Arctium tomentosum Mill. Armeniaca vulgaris Lam. Artemisia spp. Atraphaxis muschketovii Krasn. Berberis iliensis M. Pop. Berberis karkaralensis Kornilova Biebersteinia multifida DC. Bupleurum aureum Fisch. Camelina microcarpa Andrz. Celtis caucasica Willd. Centaurea turkestanica Franch. Chelidonium majus L. Cichorium intybus L. Codonopsis clematidea (Schrenk.) Corydalis semenovii Regel Corylus avellana L. Cotoneaster caratavicus Pojark. Cousinia grandifolia Kult. Crataegus ambigua C. F. Mey. Delphinium ssp. Dracocephalum nutans L. Dryopteris filix-mas (L.) Schott. Eremurus ssp. Euonymus semenovii Rgl. Ferula leucographa Korov. Ferula ssp. Fragaria vesca L. Fraxinus potamophila Herd. Geranium ssp. Glycyrrhizza uralensis Fisch. Gymnospermium altaicum (Pall.) Spach. Helichrysum maracandicum M. Pop. ex Kirp. Heracleum dissectum Ldb. Humulus lupulus L. Hypericum perforatum L. Inula helenium L. Inula macrophylla L. Iridodictyum kolpakowskianum (Regel) Rodion.

310

Iris alberti Regel Juniperus seravschanica Kom. Juno coerulea (B. Fedtsch.) Pojark. Jurinea almaatensis Iljin Lamium album L. Leonurus turkestanicus V. Kresz. Lepidolopha karatavica Pavl. Libanotis schrenkiana C. A. Mey. Lonicera iliensis Pojark. Malus niedzwetzkyana Dieck. Malus sieversii (Ledeb.) M. Poem. Mentha asiatica Boriss. Muretiea transitoria Korov. Nepeta ssp. Origanum tyttanthum Gontsch. Origanum vulgare L. Oxycoccus microcarpus Turcz. Paeonia hybrida Pall. Pistacia vera L. Polemonium caucasicum N. Busch. Polygonum bistorta L. Potentilla ssp. Prangos ssp. Pseudoeremostachys severtzovii (Herd.) M. Pop. Rhaphidophyton regelii (Bunge) Iljin Rheum maximowiczii Losinsk. Rheum wittrokii Lundstr. Ribes janczewskii Pojark. Rosa pavlovii Chrshan. Rumex tianschanicus Losinsk. Salvia sclarea L. Scaligeria alloides (Rgl. et Schmalh.) Boiss. Spiraeanthus schrenkianus (Fisch. et Mey.) Maxim. Tanacetum vulgare L. Taraxacum officinale Wigg. Tulipa brachystemon Regel Tulipa greigii Regel Tulipa kaufmanniana Regel Tulipa kolpakovskiana Regel Tulipa ostrovskiana Regel Tulipa regelii Krasn. Tulipa tarda Stapf. Ungernia severzovii (Regel) B. Fedtsch. Urtica dioica L. Vitis vinifera L.

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integral part of the vegetative formations of the mountain systems in the southeastern region of the Republic, vary according to floristic, coenotypic compositions, diversity of natural conditions, and habitats. Other major species of higher plants are closely connected with the fruit forests, among which are many endemic, relict, vanishing and genetically unique plants, the overwhelming majority of which are listed in The Red Books of the former USSR and the Republic of Kazakhstan. Modern fruit stands are the result of successive development of many valuable and useful tertiary vegetation that require preservation. The wild fruit plants of Kazakhstan may be classified as follows: Pome fruits: apple, pear, mountain ash, hawthorn, cotoneaster, and quince Stone fruits: apricot, cherries, and plum Small and vine fruits: barberry, bilberry, blackberry, blueberry, cranberry, currant, foxberry, gooseberry, grape, raspberry, stoneberry, strawberry Other fruits: elder, honeysuckle, mulberry, oleaster, sea buckthorn, viburnum Nuts: almond, pistachio, hazelnut, walnut, siberian pines Sixteen species of Kazakhstan fruit plants are registered in The Red Books of the former USSR and in Kazakh SSR. II. POME FRUITS Pome fruits represent an economic-botanical plant group that includes a great number of species and cultivars, particularly diverse for apple and pear. The most important species for fruit growing are apple, pear, quince, and mountain ash. Pome fruits are generally trees, seldom shrubs as found in quince. In comparison with other fruit crops, they are distinguished by high winter-cold tolerance and longevity. Propagation is done typically by grafting, and at other times by suckers and cuttings. Fruits of many pome fruits, in contrast to those of stone fruits and small fruits, can be preserved fresh for a long time. In Kazakhstan, pome fruits are represented by five genera; Malus Mill. (apple), Pyrus L. (pear), Sorbus L. (mountain ash), Crataegus L. (hawthorn), and Cotoneaster Medik. (Cotoneaster), all members of the Rosaceae Juss. A. Apple Among fruit crops of temperate-zone countries, undoubtedly apple occupies the first place in terms of both area under cultivation and total pro-

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duction. Its importance may be compared, for example, with that of wheat among the cereals, cabbage among leafy vegetables, or sunflower among field oil-bearing plants. The total area of cultivated apple plantations in the world is 3 million ha, and total annual yield averages 11 million tons. The value of apple fruits in human nutrition is considerable. Sugar content has a range of 3.5 to 18.5 percent. Organic acids and vitamins are important (vitamin C is about 4.5 mg/100 g of fruit). In addition, minerals are important (potassium is 248 mg/100 g of fruit). Apple fruits are used fresh, for production of juices, compotes, pastille, jam, cider, and dried fruits and they are also recommended for anemia, avitaminosis, heart-vascular diseases, and as a dietetic agent. The genus Malus includes deciduous trees and shrubs. All species occur naturally only in the Northern Hemisphere. Apple is winterhardy, light-demanding, but undemanding as to soils. Many apple species are grown as ornamental plants and are used in protective forestation. All species are good honey plants. In Kazakhstan there are three Malus species (see Table 3.2). 1. Sievers Apple [Malus sieversii (Ledeb.) M. Roem.]. This is the main forest-forming species of the fruit forests in Kazakhstan. It grows on northern, northeastern, and northwestern moist slopes and also in alluvial plains of the mountain rivers at 800 to 1500 m above sea level. Here on moist, rich-in-humus, chernozem soils, in open stands and under optimal light conditions, trees are 12 to 14 m high with trunks 70 to 80 cm in diameter. Crowns of trees are about 10 to 12 m in diameter and

Table 3.2.

Malus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Malus sieversii (Sievers apple)

Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Ters-kei, Kirghiz, Tallaskei Alatau, Ketmentau, Karatau

Food, decorative

M. kirghisorum (Kirghiz apple)

Dzhungarskei, Zailiyskei Alatau

Food, decorative

M. niedzwetzkyana (Niedzvetzky apple)

Karatau, (Berkara Gorge) Western Tien-Shan, Mashat mountains, Dzhungarskei Alatau (everywhere as single trees)

Decorative, valuable for selection

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spreading, a typical form for many apple cultivars. Similar trees are found in valleys (gorges) of the rivers Malay Almatinka, Kotur-Bulak, Talgar in Zailiyskei Alatau, and Pikhtovoe gorge in Dzhungarskei Alatau. Individual apple trees also grow on dry southern slopes. Here development is changed greatly; growth is 2 to 3 times weaker, trees become “dwarfish,” shrubby with 2 to 7 trunks, and they are no taller than 3 to 5 m. Older branches are first grayish-brown, then brownish-gray, and two-year shoots are brown, with dispersed hairs. Current-year shoots are green-brown and densely hairy or woolly. Buds are small, obovate, rubyred. Leaves are alternate, oblong or obovate, 6 to 11 cm long and 3 to 5.5 cm wide, slightly leather-textured, dark-green on the underside, strongly pubescent with serrate or crenate margins. Petioles are always shorter than leaf blades. Inflorescences have 3 to 7 flowers that are 5 to 11 cm in diameter and are often located on spurs, seldom on the ends of lastyear shoots. Flowers are from 4 to 6 cm in diameter and are borne on thick woolly pedicles. Sepals are lanceolate, green, sometimes with crimson stripes at the ends and with thick pubescence. Petals are 5, rounded or ovate, smooth or sometimes corrugated, pale-pink or pink, irregular colored, more intensive on petal margins or on the base. This species is very polymorphous in fruit form and size. The period of ripening is extensive; from early, middle, and late due to different natural forms. The earlier forms begin to ripen at the end of July, the majority ripens at the end of August, and the late forms ripen from midSeptember to early October. Rounded or oblate forms prevail though rounded-cylindrical, cylindrical, and conical fruit forms are found. Fruits are green, yellow, pink, and red. In spite of diverse skin color, fruit flesh is white to light yellow; it is juicy, tender, and of different flavors, from insipid and sweet, sour-sweet, sour to very sour with bitter flavor. Fruit surface is mainly smooth, but sometimes ribbed, and with a strong waxy bloom. Stem cavity differs in width and depth and is always well expressed. The calyx basin is shallow, wide, and mainly ribbed. Calyx is partly closed, and fruit stem varies in length. The average fruit weight is 20 to 60 g, with a maximum of 120 g. Seeds are small, brown, sometimes almost red, irregular obovate, with sharply acuminate, curved rostellum. One fruit contains 1 to 15 seeds. From 1 kg of fruits it is possible to obtain 4 to 16 g of seeds. Under natural conditions, Sievers apples are propagated by seeds (15%) and vegetative methods (85%). Root-suckers are formed from dormant buds at the base of the trunk from upper roots and 150 to 200 suckers of different age can be found around one tree. In spite of delayed growth Sievers apple is very long-lived, reaching 150 to 170 years. Fruits of Sievers apple are valuable medicinal products; they are components of many special food products and are recommended against diseases of gastrointestinal tract and avitaminosis.

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In these respects, they prevail over present cultivars. This apple species is of great interest for selection, especially in breeding for drought tolerance, winter-hardiness, and high vitamin content. There are also some ornamental forms. 2. Kirghiz Apple (Malus kirghisorum Al. et An. Theod.). Sievers apple and Kirghiz apple are very similar morphologically and ecologically. Kirghiz apple forms pure apple forests on the slopes of Zailiyskei and Dzhungarskei Alatau, and sometimes it is mixed with aspen, hawthorn, and Semenov maple (Acer semenovii Regel et Herd.). It grows mainly on the northern slopes at 1200 to 1800 m above sea level, prefers deep rich forest soils, and is considerably frost-resistant. It is a mesoxerophytic tree, 10 m tall, with gray bark and a broad, umbrella-shaped crown. Shoots are long, thin, and without spines. Leaves are of papery consistency, large (5–10 cm), long, and 3 to 6 cm wide, broadly oblanceolate, tapered to the base with denticulate margins and glabrous or thinly pubescent undersides. Petioles are short and pubescent. Flowers are large, 3 to 5 cm in diameter, and pale-pink. Flowering occurs in April to May and fruits mature in August to September. Fruits are rather diverse in size, form, color, and flavor. They are borne singly and are 3 to 8 cm long and 3 to 8 cm in diameter; shapes are globular or cylindrical and narrowed to the top. Skin color is yellow, greenish, or reddish. Flavor is from sour-sweet and acid to bitter and astringent. Forms with 10 to 12 percent sugar are found. Fruits are widely used fresh and as processed products by local people. Kirghiz apples are excellent raw material for processing cider, fruit paste, jam, stewed fruits, puree, natural juices, and those with high sugar content are good for drying. In addition, these trees have ornamental value for landscaping. 3. Niedzvetzky Apple (Malus niedzwetzkyana Dieck.). In the Kazakh Republic this species is found in the mountain forests of Karatau (Berkara gorge), Western Tien-Shan (Mashat mountains), and Dzhungarskei Alatau (Pikhtovoe gorge). They exist in these sites as single trees. It is a mesophytic tree, 5 to 8 m tall, with globular crown and reddishbrown bark. Annual shoots are dark-purple, and older branches are reddish-brown and without spines. Leaves are obovate or elliptical, slightly leathery, dark-green, sometimes violet-red, with serrate or crenate margins. In April to May, beautiful intense pink or purple flowers appear. They are 4 cm in diameter and are borne on thin whitetomentose, pubescent pedicels. Fruit skin is violet-purple with bluish, waxy bloom and the flesh is pink-purple and edible. Seeds are darkbrown with crimson blush.

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Niedzvetzky apple is winter-hardy, undemanding for care, droughtresistant, and interesting exclusively for ornamental horticulture and hybridization. I. V. Muchurin used this species to breed remarkable hybrids of red-leaved, red-flowered and red-fruited apples (‘Belfler Red’, ‘Belfer Record’, ‘Red Standard’, and others). Taking into consideration the importance of Niedzvetzky apple for hybridization, it is necessary to preserve it in its natural habitats. Wild apples of Kazakhstan are valuable genepools for use in selection. Therefore, the preservation of natural apple forests of the Republic is a problem of State importance. B. Pear Pear (Pyrus communis L.) is the most popular fruit tree after apple in countries at moderate latitudes such as France, Belgium, and Germany. Cultivated pears have spread farther to the north than the range of the wild species. It is cultivated in all countries of Western Europe and North America. Pear trees are deciduous, seldom-tall shrubs ranging from 5 to 30 m in height. Branches are usually spiny. Leaves are alternate, simple (sometimes pinnately divided and lobate), orbicular or ovate, margins entire, serrate, dentate, or crenate. Flower is bisexual, regular, white, seldom pink, averaging 3.0 cm in diameter. Inflorescence is a corymb. Fruit is apple-shaped, ovate, oblate, or pyriform. Seeds are brown or brownish with luster and slimy in the fruit. Trees bloom in April to May with fruit ripening from July to September. Wild trees are mainly propagated by seeds, whereas cultivars are propagated by grafting. Pear is more demanding than apple in regard to environmental conditions; it is warm-loving, light-demanding, relatively drought-enduring, rather frost-resistant, but it is not tolerant of excessive heat and low humidity. European pear fruits are valuable for their tender, melting flesh, fine aroma, pleasant combination of sugars and acids, high flavor, and dietetic qualities. Depending on the cultivar and the ecological and geographical growing conditions, the chemical composition of pear fruits may vary considerably. They contain 70 to 80 percent water and 6 to 15 percent sugar (mainly fructose 5–13%). Pear is inferior to apple in total sugar content but it seems sweeter because of low acidity (0.1–0.2%). The amount of ascorbic acid is 5 to 7 mg/100 g, pectin substances 0.1 to 0.6 percent, and tannin substances 0.32 percent. Pear fruits are used fresh, dried, canned, and for production of candied peels, pastille, marmalade, and wines. The genus Pyrus includes about 60 species, and half of them are inhabitants of the Caucasus and Central Asia. Possibly the most ancient center of origin of pear was in regions of China that escaped

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glaciation. However, the modern center is the Caucasus. The genus occupies a narrow-elliptical band of Eurasia from Gibraltar and the Atlas Mountains in Northern Africa through the whole Asian continent, including Japan. In Kazakhstan, there is one wild species, that is, Regel’s pear or diversifolious pear (P. regelii Rehd.). It grows as small formations in the mountain savannah and is a low, graceful, xerophytic tree, seldom shrub, 3 to 6 m high with wide-spreading branches. Shoots are reddishbrown and older wood gray. The leathery, shiny leaves are of different shapes; on one tree it is possible to find both entire and pinnatelydissected leaves with narrow-lanceolate or lanceolate lobes with small teeth. Flowers are in small corymbs, white, and 2.0 to 3.5 cm in length and width. The greenish-yellow fruits are bitter-sweet, strongly astringent, almost inedible fresh, but after storage they become soft and tart. There are four to six seeds per fruit. The dark-brown, almost black seeds are much larger (7 × 10 mm), than those of most other Pyrus species. Trees bloom in May, and fruits mature in August to September. Regel’s pear is an endemic plant of the Tien-Shan, Pamirs, and Altai. In Kazakhstan, it is found in Chu-Ili mountains, Kirghiz, Tallaskei Alatau, and Karatau. Trees grow on dry stony slopes of foothills among rocks, and sometimes in valleys on moister soil. Trees are found in mountains up to 1200 m. This is the most drought-resistant species of wild pears in Central Asia, and thus deserves attention as rootstock for nonirrigated orchards. Regel’s pear represents special value for breeding drought-enduring cultivars. It has potential for reforestation of stony slopes in southern regions of Kazakhstan. It is an ornamental plant suitable for landscaping in the southern part of the Republic. Regel’s pear is cultivated in a number of botanical gardens such as Valonezh, Tallin, St. Petersburg, Kiev, Moscow, Bishkek, Tashkent, Almaty, and Karaganda. It is propagated easily by seeds, although seedlings grow very slowly, especially in the first years. It is a medicinal, ornamental, food, and honey plant. C. Cotoneaster Genus Cotoneaster (Cotoneaster Medik.) refers to the pome group which includes 13 species in Kazakhstan: Goloskokov cotoneaster (C. goloskokovii Pojark.), Karatavskei cotoneaster (C. karatavicus Pojark.), Krasnov cotoneaster (C. krasnovii Pojark.), large-fruited cotoneaster (C. megalocarpus M. Pop.), black-fruited cotoneaster (C. melanocarpus Fisch. ex Blytt.), multiflorous cotoneaster (C. multiflorus Bunge), Antonina new cotoneaster (C. neo-antoninae A. Vassil.), oliganthous

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cotoneaster (C. oliganthus Pojark.), Pojarkova cotoneaster (C. pojar-kovae Zak.), false-multiflorous cotoneaster (C. pseudomultiflorus M. Pop.), Roborovsky cotoneaster (C. roborowskii Pojark.), pleasant cotoneaster (C. suavis Pojark.), and single-flowered cotoneaster (C. uniflorus Bunge). D. Hawthorn Hawthorn (Crataegus L.) is a large genus containing as many as 1250 polymorphous, wild species. These species inhabit the temperate, seldom subtropical, belts of the Northern Hemisphere. The exact number of species is not established; in America alone, more than 1000 species are described. Hawthorn is most often known as an ornamental shrub rather than a fruit plant. China is one of the few countries in the world where hawthorn is cultivated as a fruit crop. Many species are cultivated as plants for hedge landscapes because of the ease of pruning. They are also used as a rootstock for apple, pear, and quince. Fruits of some species have gained wide recognition for their special distinctive flavor and dietetic qualities. Fruits contain from 4 to 11 percent sugars, mainly fructose; among organic acids, malic acid prevails. Content of pectin substances reaches 0.6 to 1.6 percent, tannin and pigment substances 0.8 to 1.7 percent, ascorbic acid is 31 to 108 mg/100 g of fruit, vitamin A content is 380 to 680 mg/g of fruit. They are deciduous, seldom semi-evergreen, small trees, often shrubs, 3 to 6 m tall. Stems are covered by thorny spines (modified shortened shoots), 3 to 6 cm long. Bark is gray with light or dark tints, smooth at first, but with age becomes cracked. Leaves are from entire and pinnately-dissected to pinnately-lobed, alternate, pubescent or glabrous. Flowers are white (garden forms sometimes are pink and red) in cymose or corymbose inflorescences with unpleasant aroma. Fruit is a small pome with 1 to 5 stony seeds, and is orange-yellow, red-purple, or black. Hawthorn is winter-hardy, light-demanding, drought-resistant and undemanding as to soil. It is propagated in nature by seed or sprouts and in culture, by layers or grafting. Trees live 200 to 300 years. In Kazakhstan, seven wild species are found: three of them rare (Table 3.3). 1. Alma-Ata Hawthorn (Crataegus almaatensis Pojark.). This species grows along mountain slopes, in valleys and bottom lands, among mixed forests and shrubs. Trees are mesophytic, to 10 m tall with mostly a single trunk, though multistemmed trees of 3 to 5 m with low, umbrellashaped crowns are found. Bark on old branches and stems is gray and cracked; on younger shoots it is light-brown, glossy, and glabrous; growing shoots are green and hairy. Spines are few or absent; if present they

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318 Table 3.3.

A. DZHANGALIEV, T. SALOVA, AND P. TUREKHANOVA Crataegus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Crataegus almaatensis (Alma-Ata hawthorn)

Dzhungarskei, Zailiyskei Alatau

Ornamental, food, medicinal, honey; endemic in Kazakhstan

C. ambigua (Doubtful hawthorn)

Ustyurskei, Karatau, Mangyshlak Peninsula

Ornamental, honey

C. cholorocarpa (Altai hawthorn)

Ulutau, Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz Alatau, Ketmentau, Karatau, Western Tien-Shan

Ornamental, food, honey

C. pontica (Ponti hawthorn)

Karatau, Western Tien-Shan

Ornamental, food, medicinal, honey; rare in Kazakhstan

C. sanguinea (Blood-red hawthorn)

Altai, Tarbagatai, Dzhungarskei Alatau. Common Syrt, Semipalatinsk pinewood, Kokshetau, Torgai, Zaisan

Ornamental, medicinal, honey, food

C. songarica (Dzhungarskei hawthorn)

Dzhungarskei, Zailiyskei, Kungei, Kirghiz Alatau. Western Tien-Shan, Chu-Ili mountains

Ornamental, medicinal, food, honey

C. turkestanica (Turkestan hawthorn)

Karatau, Western Tien-Shan

Ornamental, food, honey

are strong, brown, and shiny, 1 cm long. Leaves on vigorous shoots are ovate or elliptical-ovate, wide, rhombic-shaped with an acute apex and a cuneate entire base; young leaves are slightly pubescent with tufts of hair in vein angles. Petioles are 3 cm long. Inflorescences are loose, many-flowered cymes. Flowers are 1 to 3 cm wide with white petals. Fruits are oval or almost globular; mature fruits are dark-purple, almost black, glossy and glabrous, 10 to 15 mm in diameter with juicy, purplishred flesh. Each fruit has 3 to 5 stones that are light brown with a grooved inner surface. Fruits are edible but they are not used industrially, because they are small and sugar content is lower than in other species. Alma-Ata hawthorn is an excellent ornamental plant, especially during bloom and in the fruit-bearing season. Therefore, it has been introduced

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into many botanical gardens in the Republic and abroad. Its winterhardiness allows adaptation to many environments and it bears fruit annually. Alma-Ata hawthorn is an endemic plant of Kazakhstan. 2. Doubtful Hawthorn (Crataegus ambigua C. A. Mey. ex A. Beck.). This is a very rare species in Kazakhstan. It grows in gully bottoms in chalky and gypsum gorges, on moist soils of Mangyshlak Peninsula and in Ustyurskei Karatau mountains. Plants are deciduous shrubs or small trees 3 to 4 m tall; bark of older wood is gray-brown, with gray, hairlike, scaly spots; young shoots are red-brown, glabrous, and thornless. Leaves are bright-green, wide-ovate, with 5 to 7 sharply lobate lobes. The leaves are 5 cm long and 3 to 4 cm wide, with a wide-cuneate or orbicular base. Inflorescences are dense, containing about 20 flowers distributed on 2 to 6 short spurs; sepals are ovate-triangular with short beaks at the ends. Corolla is about 15 mm in diameter. Fruits are wide, elliptical, 11 to 14 mm in diameter, and purplish-black with light dots; flesh is juicy but fruits are not used for food. Trees bloom in May and fruits mature in July to August. Plants are propagated by seeds. In culture, doubtful hawthorn is best known in Mangyshlak Experimental Botanical Garden. It is recommended for town and settlement landscaping on the Mangyshlak peninsula. It is necessary to establish a botanical preserve forest in the mountains of Mangistauskie Karatau for preserving the natural stands of doubtful hawthorn. 3. Altai Hawthorn (Crataegus cholorocarpa Lenne et C. Koch.). This species grows along gorge slopes, mainly in understory of deciduous and mixed forests, among shrub-lands and along river valleys in stony, finegrained soils. It is a water-loving plant. Plants are small trees or large shrubs about 5 to 6 m tall with 3 to 4 and sometimes up to 15 stems of different height and diameter, with rare short, straight spines, 0.6 to 2.0 cm long. Bark of old trunks is gray, on older branches, it is yellow-gray or reddish-gray with large light-gray lenticels, and annual shoots are glossy, smooth, brown-red or green-brown with numerous light lenticels. Leaves are light-green, wide, triangular-ovate to orbicular, 3.5 to 10 cm long, 2.5 to 9 cm wide, with acute apex and rounded-cuneate base, 7 to 9 lobate, with acute-toothed margins. Inflorescences are corymbose, 8 to 9 cm in diameter, and with 27 to 35 flowers. Flowers are white, 13 to 16 mm in diameter; sepals are triangular-ovate, recurved. Fruits are small, globular or flattened-globular, yellow or orange-yellow, 8 to 12 cm in diameter with mealy, sweet, tasty flesh, and contain 4 to 5 stones. Fruits are edible but they are not harvested for industrial use, although fruitbearing of Altai hawthorn is abundant and annual. Fruits accumulate

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about 18 percent sugar, contain free organic acids, mainly malic acid, pectin substances, vitamins C, A, and carotene. They are used for different kinds of processing, such as jam, jelly, pastille, and pie filling. 4. Ponti Hawthorn (Crataegus pontica C. Koch). This species grows mainly as pure stands on southern or southeastern rocky, and sometimes gravelly, slopes. It prefers loamy soils with some stones. In its droughtresistance and soil requirements, Ponti hawthorn is similar to common pistachio, except it can grow at considerably higher elevation of 1400 to 1500 m above sea level. Trees are xerophytes, 6 to 10 m tall, with a hemispheric, wide crown. Branches are dark-gray; annual shoots and sprouts are tomentose without spines. Leaves have short petioles, are firm, bluish-green, rhomboid-shaped or wide-obovate, with wide-cuneate base; 5-lobate, seldom 7-lobate, 5 to 7 cm long, 4 to 6 cm wide, entire or with few large teeth at the apex. Young leaves are pubescent, and later, glabrous. Inflorescences are compact corymbs with 6 to 14 flowers; corolla is 15 to 20 mm in diameter, pedicels 3 to 7 mm long, sepals are triangular, densely-hairy, and recurved. Fruits are large (3 cm in diameter, weight is 1.6–4.8 g); shape is pyriform or globular, flattened at the poles, yellowish-green at first, turning at maturity, to yellow, orangeyellow or dirty-yellow with numerous red or pink-bluish dots. Fruit flesh is juicy, tender, aromatic, and acid, acid-sweet, or sweet flavor. Seeds, usually 2 to 4 per fruit, are hemispheric with domed back and 2 to 3 grooves; average seed weight is 0.2 g. Trees bloom in May, and fruits mature in September. Yields are heavy, but fluctuate sharply from year to year in wild stands. The periodicity of fruit-bearing is not a biological peculiarity of Ponti hawthorn, but is probably explained by the severe droughts of southern slopes on which it grows because under irrigation it bears fruits annually. Selections of individual seedlings have been found which are consistently heavy yielding annually. The average yield per tree is 12 to 20 kg with a maximum of 80 kg. Trees are propagated by seeds. A distinctive attribute of this species is the surprisingly high survival rate of its seedlings. In maple, apple, and other mesophytic species growing with Ponti hawthorn, seedling mortality at the beginning of the dry period reaches 80 to 95 percent. However, in Ponti hawthorn loss is not more than 10 to 15 percent; in fact, most of these seedlings die because of mechanical damages, mainly by cattle trampling rather than by natural causes. Such survival is the result of a welldeveloped root system in seedlings. In the first year with the aboveground part 10 to 12 cm tall, roots are 90 cm long. Undoubtedly, under conditions of Central Asia, Ponti hawthorn may, and in fact, must become a new fruit crop for southern nonirrigated fruit-growing.

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5. Blood-red Hawthorn (Crataegus sanguinea Pall.). This species grows in forest, forest-steppe and steppe zones, in open forests, at forest borders, and on banks and alluvial plains of rivers. It is a thorny shrub, sometimes small tree, 1 to 4 m tall; trunk is 10 cm in diameter; branches are purple-brown, glossy with long (2.0–3.5 cm), strong, thick, shiny, red-brown straight thorns. Leaves are slightly dull, sparsely hairy, 7 to 9 lobate, 3 to 7 cm long, 2 to 5 cm wide, rounded-ovate, with acutely serrate margins, and base is wide-cuneate. Flowers are in corymbs on ends of short spurs; calyx has wide-triangular acute teeth; corolla is white, 12 to 13 mm in diameter. Fruits are almost globular or short-elliptical, 8 to 10 mm in diameter, orange-yellow or bright red with mealy flesh, and have 3 to 4 seeds with a hard seed coat. Fruits are edible, sourish-sweet; they contain sugars, organic acids, tannin substances, saponines, glycosides, carotene, and vitamin content is 127 mg/100 g of fruit. The trees are ornamental, widely used for hedges, or as a group or as individual trees in the landscape. It is also a good honey plant. 6. Dzhungarskei Hawthorn (Crataegus songarica C. Koch.). This species grows on rocky slopes of gorges and river valleys of the middle mountain belt. Small tree or large shrub with many trunks ranging from 5 to 9 cm in diameter. Bark of the trunks is reddish-gray to blackish, finely-split and exfoliates in thin strips. The brownish, smooth bark on older branches and reddish-brown bark on one-year shoots distinguish this species from many others. Current-year shoots are greenish, glabrous, or sometimes covered by thin, white hairs, and with straight thorns 5 to 15 mm long. The slightly pubescent leaves are wide-ovate to almost rhomboid-shaped with cuneate base and 5 to 9 divided, unequal lobes; the apical lobes being large and serrate while the basal lobes are larger often with large incisions. Inflorescences are corymbs with 28 to 35 flowers. These flowers are 1.7 to 1.8 cm in diameter. Peduncles are densely-hairy; sepals are entire, acute-triangular, recurvate and hairy; petals are orbicular and white. Fruits are orbicular or wide-oval, 1.2 to 1.4 cm long, 1.0 to 1.3 cm wide, dark-red, with orange, sweet, edible flesh. Each fruit has 2 to 3 seeds which are smooth on the ventral side with oblique longitudinal grooves. Plants are propagated by seeds. Dzhungarskei hawthorn is of limited importance as a fruit tree; it may be used in mountain forest reclamation for soil stabilization along river banks. 7. Turkestan Hawthorn (Crataegus turkestanica Pojark.). This species grows in walnut and apple stands on northeastern and northern slopes, in pistachio stands on loess deposits of southern slopes, and in almond

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forests on stone mounds of rock terraces. Trees are strongly branched with thick crowns and are 7 to 8 m tall, with gray-brown trunk, grayishbrown divaricate branches and red-brown young shoots; spines are thin and 1 to 1.5 cm long. Leaves are ovate, 3.5 cm long with 5 to 7 acute, unequally-serrate lobes; early developing leaves are reddish-green, but in summer they are bright-green; young leaves are densely pubescent on both sides, but as they mature they become almost glabrous. Inflorescences are corymbs with 12 to 15 white flowers with each flower 16 to 18 mm in diameter. Fruits are wide-elliptical or rounded, bright-red, turning to dark-cherry as they mature, with an average weight of 0.8 g; skin is coarse and thick; flesh is yellow, mealy and edible; seeds are rather large, elliptical and rounded-elliptical, scabrous, and slightly ribbed. Under natural conditions, Turkestan hawthorn is easily propagated by seeds. This inconspicuous species persists after all other woody plants are cut because it is not used for fire wood due to abundant sharp thorns. It is exclusively valuable in the mountain forest reclamation for anchoring soils along rivers and mountain slopes. E. Mountain Ash Mountain ash (Sorbus) is a popular and famous plant and, as a rule, is considered as a forest fruit that is cultivated as an ornamental and not widely distributed as a fruit crop. Nevertheless, its fruits contain from 4 to 14 percent of sugar and the vitamin C content is similar to that of black currant and lemon. In mountain ash fruits, there is twice the amount of carotene as in carrot, the main raw material for extracting carotene preparations. They contain from 2000 to 3000 mg of vitamin A/100 g of fruit. And fresh fruits are rich in valuable phytoacids. Trees or large shrubs are deciduous up to 25 m tall; bark is smooth or split, gray or reddish. Leaves are alternate, simple, entire, pinnately-dissected, lobate or compound odd-pinnate with serrate or toothed margins. Flowers are bisexual, white, light-yellow or pink, in corymbose inflorescences. They are pollinated by insects. Fruits are 2 to 5 locular small pomes. Wild trees are propagated by seeds or layers and in cultivation, by cuttings and grafting. They live 200 to 300 years, are winter-hardy, prefer moist, fertile soils and avoid injury from a variety of air pollutants. They are exceptionally decorative from spring to late autumn. There are about 100 species of Sorbus in the temperate belt of the Northern Hemisphere. Three species grow in Kazakhstan (Table 3.4). The southern habitats of mountain ash are situated in the upper borders of forest zones in the mountains.

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Sorbus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Sorbus persica (Persian mountain ash)

Karatau, Western Tien-Shan

Decorative, food

S. sibirica (Siberian mountain ash)

Tobolo-Ishim, Semipalatinsk pinewood, Zaisan, Altai

Decorative, food

S. tianschanica (Tien-Shan mountain ash)

Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz Alatau, Western Tien-Shan

Decorative, medicinal, food

1. Persian Mountain Ash (Sorbus persica Hedl.). This species grows in the understory of deciduous and juniper mountain forests and in shrublands on the slopes of northern exposure, and in hollows, where soil moisture is adequate. Persian mountain ash does not form extensive stands, but occurs as small groves or individual trees. They are small trees 2 to 5 m tall with the broad, open crown arising almost from the base of the tree. Young shoots are thickly pubescent or tomentose. Later they lose this pubescence and become glossy, yellow-red-brown. Leaves are simple, lobulate, elliptical to rounded-elliptical, 5 to 10 cm long and 5 to 6 cm wide; upper surfaces are glabrous, bright green and leathery; lower surfaces are silvery-gray with pubescence. Inflorescences have 20 to 22 flowers and are very compact, about 9.0 cm in diameter. Peduncles are densely hairy. Fruits are small, broad-elliptical, in racemes; average fruit weight is 1.3 g. The edible fruits accumulate 10 percent sugar; skin is rather thin, tender; and flesh is orange, juicy, sour-sweet, and slightly astringent. The plant is very ornamental and is also valuable in forest reclamation for conservation of soil on river banks and mountain slopes. 2. Siberian Mountain Ash (Sorbus sibirica Hedl.). This species grows on borders of pine forests and in mountains in the deciduous forest belt. Trees are 3 to 10 m tall; leaves are 10 to 20 cm long, 8 to 12 cm wide; leaflets are oblong-lanceolate, 3.5 to 5.0 cm long, 1.5 to 2.0 cm wide; upper surfaces are green and smooth; lower surfaces are gray-green, pubescent along mid-vein, and margins are subulate-toothed. Inflorescences are dense, 6 to 8 cm long, 8 to 12 cm wide. The large fruits are

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12 mm in diameter, rounded, juicy, orange-red, astringent, and bittersweet. They contain 4 to 6 percent sugar, 40–60 mg vitamin C/100 g of fruit, and 5 to 6 mg of carotene/100 g of fruit. This species is distinguished by its exceptional winter-hardiness (withstands frost of –50°C). Siberian mountain ash is sensitive to light conditions: with low light intensity, trees develop poorly and bear very few fruits. This species is propagated by seeds and is recommended as one of the most winterhardy species for use in hybridization aimed at breeding for this trait. 3. Tien-Shan Mountain Ash (Sorbus tianschanica Rupr.). This species grows at the upper borders of fir forests in the mountains and in shrublands at 2000 to 3000 m above sea level. It is a small tree or large shrub 3 to 5 m tall with a wide crown that begins near the base of the tree. Older shoots are reddish with thin, exfoliated, gray, scarious fibers. Young shoots are olive or red-brown, slightly pubescent. Leaves are compound odd-pinnate, glabrous, 10 to 16 cm long, with 6 to 8 pairs of lanceolate leaflets, each 5.0 cm long and 1.3 to 1.6 cm wide; upper surfaces are slightly leathery and lower surfaces white-green with typical metallic luster. Inflorescences are very large, 10 to 15 cm wide and loose, and have 25 to 30 flowers. Fruits are 12 mm in diameter, rounded or slightly flattened, at first yellow-red, then dark-red with bluish bloom. The bitter, astringent fruit of Tien-Shan mountain ash are edible, especially after the first autumn frosts. They contain ascorbic acid, vitamin A, tannin substances, and carotene. Trees are propagated by seeds, layers, or root sprouts. Tien-Shan mountain ash is an endemic species for Central Asia. The great importance of this species in nature is its ability to anchor banks of the mountain rivers. It may also be used for breeding winter-hardy mountain ash cultivars or as a rootstock for valuable cultivars of mountain ash. III. STONE FRUITS Wild stone fruit crops of Kazakhstan include five genera in the Rosaceae: Armenica Scop., Cerasus Mill., Prunus L., Padellus Vass., and Padus Mill. Although these five genera are usually all included within Prunus, they will be treated separately here. Stone fruits are a very interesting plant group and the source of many cultivars, which are of great importance to human nutrition. They are distinguished by early maturity, weakly expressed alternate fruitbearing, and high productivity. In Kazakhstan, fruit maturity of most cultivars is at the end of spring and the beginning of summer, when there is a great shortage of fresh fruits. Nutritional and medicinal value of

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stone fruit plants is very high: they contain sugars, organic acids, tannin substances, nitrogenous compounds, and vitamins. These fruits are excellent for fresh consumption and are valuable raw materials for the canning industry and for the production of different juices, jams, fruit paste, compotes, marinades, and dried fruits such as prunes, and dried stoneless apricots. A. Apricot The genus Prunus armeniaca L. includes only eight subspecies, among which is the common, or wild, apricot (Prunus armeniaca, vulgaris Lam.), the major cultivated species. This is the only apricot species in Kazakhstan. It is an endemic species of northern and western TienShan, a relatively rare species with strongly decreasing natural habitat. In the northern macroslope of Zailiyskei Alatau, it usually grows in gorges of the rivers Bolshaya and Malaya Almaatinka, Talgar, Aksai, Issyk and it is plentiful in Kotur-Bulak and Karakunuz. In Dzhungarskei Alatau, the northern border of its natural habitat, it is found along the banks of the Khorgos, Usek, and Koktal rivers. The total area in the Republic occupied by this species is 1675 ha. Beyond Kazakhstan, it grows in China, Tajikistan, and Afghanistan. Apricot is undemanding of care, light-demanding, heat-resistant, winter-hardy, and droughtresistant. It bears heavy crops annually. Some plants are self-pollinated while others require cross-pollination to set fruits. Vigorous growth of trees points to the fact that they possess well-developed, deep root systems. They begin to bear fruit early, at three to four years. Wild trees are propagated by seeds. The common habitat for apricots is rocky, shallow carbonate chernozems on well-drained slopes of southern exposure in mountain steppes at 800 to 1800 m above sea level. Trees occur singly, as groups, or as small groves. It is a mesoxerophytic species. Tree is usually 7 to 9 m tall, but more vigorous specimens 15 m tall occur. Trees have an orbicular crown and a tapering stem with grayish-brown split bark. Leaves are oblong-elliptical almost orbicular, 6.5 cm long, 4.8 cm wide with an apical cusp; color is bright green, shiny (emerald) and margins have acute serrations. Leaf petioles (1.4–3.4 cm) are thin, strong, grooved, and usually dark-red. Flowers are borne on short, pubescent pedicels that average 40 mm in diameter. They are actinomorphous and bisexual, with five pink, white, sometimes crimson petals, five brownred sepals, many stamens, and one pistil. Fruit is a juicy drupe with variable shapes from ovate and oval to oblate. Skin surface is from velvet with fine pubescence to glabrous; fruit size and skin color is also variable; they weigh from 3 to 27 g and color ranges from white-cream to

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bright-orange with up to 50 percent carmine blush. Texture varies from juicy, weakly fibrous, to fleshy, sometimes with coarse fibers. Seeds are bitter or sweet. Flowering is in April and fruits mature from June to August. In nutritive/dietetic importance and distinctive flavor, apricots are superior to fruits of many other woody trees and shrubs. The high content of sugars, vitamins A and C, and some minerals make apricots among the best fruit crops. Even when dried, the amount of vitamin A in apricots is reduced not less than 50 percent. Fruits of common apricot of Kazakhstan usually contain 17.5 to 22.1 percent dry matter, 6.28 to 8.08 percent sugars, 1.95 to 3.90 percent organic acids, 0.87 to 1.32 percent cellulose, 1.10 to 3.35 percent pectin substances, and 7.41 to 12.83 mg of folic acid/100 g of fruit. However, there are some registered/selected forms of wild apricot with as high as 17.6 percent sugars, 23.8 mg vitamin C, and 9.87 mg carotene per 100 g of fruit. The wild apricot of Kazakhstan is also notable for its excellent winter-hardiness, withstanding air temperatures of –37 to 40°C, and its drought-resistance, withstanding summer temperatures over 40°C and its resistance to diseases such as, Coryneum beijerinckii Ond. Sclerotinia laxa Aderh., et Ruhe., and others. There is also variation in the duration of the dormant period, flowering time, and fruit quality. It is an exceptionally polymorphous species. Undoubtedly, wild apricot of Kazakhstan deserves great attention of fruit breeders. B. Cherry Cherries here are grouped under three genera Cerasus, Padus, and Padellus although they are often included in the genus Prunus elsewhere. 1. Tart Cherry (Cerasus). In Kazakhstan, tart cherry is the most widespread of stone fruit crops. There are four species of Cerasus native to Kazakhstan and commercial cultivars of this genus are commonly grown here. It is undemanding to growing conditions, frost-resistant, droughttolerant, and relatively shade-resistant. Its valuable attribute is precocity. Trees in this genus are deciduous trees or shrubs with simple, glandular-toothed leaves. Although somewhat variable depending on species, flowers are in umbellate inflorescences. Each flower has 15 to 50 stamens, a single ovary, and a bell-shaped or tubular hypanthium. Fruit is a globular drupe with juicy mesocarp, red or black, and generally has smooth skin. Stone is mostly rounded, sometimes acuminate at one or both ends. Fruits of wild cherry represented in Kazakhstan contain 9 to 14 percent sugar, 1.06 to 1.84 percent organic acids (predominately citric and malic

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acids), 11 to 364 mg/100 g of fruit of tannin and pigment substances, 1.9 to 19.5 mg/100 g of fruit of vitamin C, along with good quantities of carotene, protein, and cellulose. Cherry fruits are twice as rich in iron as apples. The presence in fruits of three hematogenic substances: iron, folic acid, and riboflavin determines the value of cherry for prevention of anemia. Seeds contain amygdalin and 30 percent oil. Cherry fruits are used fresh and in the canning industry for production of excellent juices, jams, fruit paste, marmalade, and for dried fruits. Wild trees are propagated by seeds and shoots. There are about 150 Cerasus species in the world and four species are described in Kazakhstan (see Table 3.5). The practical value of the Kazakhstan species is in their high droughtresistance, early and heavy fruit bearing, and fruit flavor. Red-fruited Cherry (Cerasus erythrocarpa Nevski.; syn. Prunus bifrons Fritsch.). This species grows on dry, stony slopes, among various shrub species in middle and subalpine mountain belts. The tree is a low spreading shrub 0.7 to 1.5 m tall with gray bark; young shoots are thin, shortdowny, brownish-gray. Leaves are narrow-ovate or elliptical, acute-toothed with white-tomentose undersides and green, bristly-hairy or almost glabrous upper sides. Flowers with pink petals are borne singly, in pairs or sometimes as clusters of 3 to 6 on two-year-old shoots; sepals are densely hairy on the inside. Fruit is a drupe, globular or ovate-globular, dark-red, that is

Table 3.5.

Cerasus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Cerasus erythrocarpa; syn. Prunus bifrons (Red-fruited cherry)

Western Tien-Shan, Karatau

Ornamental, food, honey

C. fruticosa; syn. Prunus fruticosa (Bushy, steppe cherry)

Tobolo-Ishim, Pre-Caspian basin, The Irtish, Kokshetau, Western low mounds

Food, honey, ornamental, medicinal

C. tianschanica; syn. Prunus cerasus (Tien-Shan cherry)

Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Ketmen, Kirghiz Alatau, Western Tien-Shan, Karatau

Ornamental, honey, food

C. verrucosa; syn. Prunus verrucosa (Warty cherry)

Western Tien-Shan

Food, ornamental, honey

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pubescent when immature and nearly glabrous when mature. Flesh is edible, though with bitter flavor and dry. It is a very polymorphous species and has potential for soil conservation on stony, dry knolls due to its well-developed root system. It is recommended as a dwarfing rootstock for cherry trees under nonirrigated conditions and for breeding drought-tolerant cultivars. Bush or Steppe Cherry (Cerasus fruticosa Pall.; syn. Prunus fruticosa Pall.). This species grows in steppe or forest-steppe zones, at forest borders and in understories, on dry southern slopes, in valleys and on riverbanks. The tree is a xerophytic shrub 0.2 to 2.0 m tall. Bark has yellow lenticels, annual shoots are gray, and older branches are dark-brown. Leaves are obovate, elliptical, or wide-lanceolate, about 12 mm long and 6 mm wide; upper sides are dark or bright-green, shiny, and glabrous. Undersides are lighter green. Leaves are acute or obtuse at the apex and cuneate at the base, with serrate margins. There are 2 to 3 white flowers in an umbellate inflorescence. Fruits are variable in size (8–25 mm diameter) and shape (rounded to pyriform); color is from pink to darkred, almost black, and taste is sour-sweet, sometimes astringent. This species is one of the progenitors of the cultivated sour cherry. It is important for breeding new winter-hardy and drought-resistant cultivars and cultivated bush cherries. Because it produces abundant roots, it is used for soil conservation on dry slopes. Tien-Shan Cherry (Cerasus tianschanica Pojark.; syn. Prunus cerasus var. humilis Bean). This species grows on stony slopes and in gorges of the Northern Tien-Shan mountains at 800 to 1000 m among diverse shrub species, sometimes forming extensive colonies. This species is a branching, globular shrub 1 to 1.6 m tall, with light yellowish-gray bark on annual shoots, brownish-gray on older shoots. Leaves are narrowlanceolate or obovate-lanceolate, 3 cm long and 0.9 cm wide; both sides are glabrous, margins are denticulate; leaf base is narrow-cuneate and apex acute. Pink flowers are on spurs usually four to six to a cluster. Fruit is a drupe, globular or ovate, dark-red, and glabrous, 7 to 9 mm in diameter. Stone is ovate with acute apex, smooth on each side with a thin net of furrows at the apex and near the suture. Fruits are edible, often with high sugar content, and have a pleasant flavor. Tien-Shan cherry may be used in hybridization for breeding new drought-resistant cultivars. In nature, hybrids with alycha (Prunus sogdiana Vass.) give fruitful progeny. It is used for soil conservation on dry stony slopes. Warty Cherry [Cerasus verrucosa (Franch.) Nevski.; syn. Prunus verrucosa (Franch.) Nevski.]. This species grows as sparse colonies on

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mounds of rocks and on stony slopes, seldom on lowlands in the middle mountain belt. It is a many-stemmed, low-growing xerophytic shrub 1.5 to 2.0 m tall with knotty, brownish-gray older branches and grayishyellow annual shoots. Leaves are obovate or obovate-lanceolate, 15 to 20 mm long and 6 to 8 mm wide, sharp-toothed with acuminate apex, usually glabrous on both sides, but occasionally with appressed-downy hairs on the lower side. Inflorescences with pink flowers are dense, with 3 to 8 flowers on spurs. Fruit is a drupe, globular or wide ovate, red to dark-red, 7 to 9 mm long. Stone is warty and acuminate at the apex. It is propagated by seeds. The main value of this cherry is its use as dwarf rootstocks for cultivated cherry and plum cultivars. Fruits are used only fresh. It has potential for stabilization of slope plantings. 2. Bird or Sweet Cherry [Padus avium Mill.; syn. Prunus avium (L.) L.]. In Kazakhstan, bird or sweet cherry (Padus avium Mill) usually grows in forests or shrub-lands on moist, open sites along river banks, and on alluvial-plains. It is found in flora regions of Syrt, Tobol-Ishim, Kokshetau, Mudozhar as well as in Semipalatinsk pine forests, eastern and western low hillocks in the Altai, Dzhungarskei, Zailiyskei, Kungei, and Terskei Alataus. It is a deciduous tree or large shrub 2 to 10 m tall; bark on stems and framework branches is brown-black or dark gray, young branches are olive-colored, or cherry-red, and inner bark is yellow. Leaves are simple, alternate, ovate-lanceolate or oblong-elliptical, 3 to 15 cm long, 2 to 7 cm wide, with cuneate base, acuminate apex, and serrate margins. Flowers are small, white, in dense, drooping racemes with a pleasant aroma. Fruit is a globular drupe, juicy, black, and glossy, with tart flesh; stone is rounded-ovate, sinuate-grooved. It blooms in May to June, and matures fruit in August to September. Edible fruits contain sugars, malic and citric acids, flavonoids, tannin substances, vitamins A and C, and there is amygdalin in fruit, bark, leaves, and seeds. Bird cherry is propagated by seeds and root sprouts, and is renewed by stool shoots. Common bird cherry is a food, vitamin, and honey plant. It is a valuable forest species, improving soil structure and stabilizing river banks and mountain slopes. 3. Mahaleb Cherry [Padellus mahaleb (L.) Vass.; syn. Prunus mahaleb L.]. Mahaleb cherry grows in southeastern Kazakhstan in the Karatau and Western Tien-Shan mountains. It is a tall 10 to 12 m, deciduous shrub with brownish-gray, glossy branches. Its glabrous, light-green, slightlyshiny, wide-ovate, almost rounded leaves are 2 to 8 cm long, 1.5 to 6 cm wide; margins are glandular-crenate, the apex suddenly narrows to a short, sharp point and base is round, slightly cuneate. Inflorescences are

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7 cm long racemes with 5 to 14 white flowers, 10 to 12 mm in diameter; sepals are ovate-triangular, obtuse, and recurved. Mature fruits are darkred, almost black, ovate, bitter, inedible, 8 to 10 mm long, 7 to 8 mm wide. Stone is white or slightly pink, smooth, ovate, and contains much amygdalin. Plants bloom in May to June and bear fruit in July to August. The main value of Mahaleb cherry is its use as a rootstock for sweet cherry cultivars. It is propagated by seeds. It is also very ornamental; there are garden forms with unique globular crowns, variegated leaves, and yellow fruits. It is recommended for hybridization with cherry cultivars aimed at breeding for drought-resistance. Mahaleb cherry is also valuable for the mountain forest reclamation, due to its well-developed root system. It is suitable for soils of dry stony slopes and riverbanks. C. Plum Plum (Prunus L.) is one of the most important cultivated stone-fruit crops of the temperate zone. Plum is inferior to cherry in winterhardiness and more demanding of soil, climate, and growing conditions. It is light-demanding, grows well on loam, moist, well-drained soils, rich in nutrient substances. Plum is precocious and it bears fruits annually with heavy yields. It consists of deciduous trees or shrubs; leaves are simple, alternate, round to lanceolate, with glands on basal margins of leaf blades or on sepals. It blooms before leaves emerge, or simultaneously with them. The five-petal, five-sepal flowers with many stamens are white, fragrant, single or in inflorescences with one to three flowers. The fruit is juicy, fleshy, with a suture, weight varies and color ranges from yellow and green to red-purple and bluish-black with a more or less waxy bloom. Fruits of plums possess high flavor and dietetic qualities. They contain 13 to 25.7 percent dry matter, 6.5 to 14 percent sugars, 0.33 to 0.98 percent pectin substances, 4.6 to 14.3 mg/100 g of fruit of vitamin C, 0.34 to 1.58 percent organic acids. They are consumed fresh, but can be processed by freezing, canning, or drying. They can be stewed or made into jams and marinades. Plum can be propagated by seeds and sprouts. There are 30 species of plums distributed in temperate latitude of two centers in the Northern Hemisphere (Eurasian and North American). Two species grow in Kazakhstan (see Table 3.6). 1. Sogdijskaya Plum, Alycha (Prunus sogdiana Vass.). This species grows in gorges and on northern slopes of the mountains, in river valleys, in understory of apple/maple forests and among shrub-lands at 800 to 2200 m. Among wild fruits in Kazakhstan, plum is more heat-loving; therefore, its natural stands are found only in the southern part of the

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Prunus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Prunus sogdiana (Sogdijskaya plum, alycha)

Kirghiz, Zailiyskei Alatau, Western Tien-Shan, Karatau

Food, medicinal, vitamin, ornamental plant. Endemic plant of Western Tien-Shan and the Pamirs-Altai

Prunus spinosa (Spiny plum, blackthorn)

Mugodzhary, Pre-Caspian basin

Food, vitamin plant

republic. The most northern habitat of alycha is in the Karakunuz river basin in the western part of a ridge in Zailiyskei Alatau. Here it grows in understory of apple forests and partially in thickets of blackberry in sparse forests of Celtis caucasica Willd., and in white mulberry (Morus alba L.) forests. This species consists of deciduous shrub or short trees 2 to 8 m tall, often shrubby with low branches almost touching the soil. They usually have none or few root sprouts. Bark on old branches is dark-gray, almost black and, on young shoots, brown-red. Numerous shortened spurs are thorny spines. Leaves are narrow-elliptical, 4 to 5 cm long, 2 to 4 cm wide, with serrate margins; upper sides are dark-green and glabrous, lower sides are lighter green with pubescent glands. In autumn, they turn to a lemon-yellow color. Prior to leaf emergence, the white flowers bloom on last-year’s growth; they are about 2 cm in diameter. Fruits are globular or elliptical, yellow, pinkish, light-red, cherryred, or blue, 1 to 5 cm in diameter, acid, sour-sweet, sometimes with bitter skin. Stone is yellowish-brown or grayish, ovate or compressedovate, and fruit flesh clings to it. Sugars (4.2–9.9%) in fruits are glucose 1.9 to 4.5 percent, fructose 1.3 to 1.4 percent, and sucrose 0.9 to 6.1 percent; of organic acids, malic and citric (1.2–3.9%) dominate. The astringent alycha flavor is due to tannin substances. The seed is rich in oil (42%). The considerable amount of pectin (1.6–2.9%), enhances jellymaking properties of the fruits. Fruits are edible and are used for production of jams, marmalades, pastille, juices, nalivkas and wines. Polymorphism of alycha is exceptionally great. It is easily crossed with apricot, plum, and peach. Therefore, it is of great interest to fruit breeders. It is recommended for rootstocks for plum, peach, and apricot because it is winter-hardy and relatively drought-enduring. 2. Spiny Plum, Blackthorn (Prunus spinosa L.). This species grows in glades, forest borders, gullies, river valleys, and on mound slopes. It is

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characterized by broad adaptability and good viability. It survives on dry soils and often grows on eroded soils along banks of gorges and on stony slopes. It is a heavily spined, strongly branching shrub, or seldom a tree 4 to 6 m tall, with dark gray, slightly splitting bark. It is a very thorny plant that with its many root sprouts forms dense thickets. Young shoots are at first velvet-like-pubescent and later become glabrous. Leaves are oblong-obovate, elliptical or lanceolate with serrate or crenate margins, pubescent at first and with age, almost glabrous, dark-green, dull, and leathery. Flowers are single, white, small, pentapetalous; in spring they bloom before leaves emerge. Fruits are round or oblong-globular, black, with tart flesh. Stone is middle-sized, orbicular or ovate, wrinkled, either cling stone or nonclinging. Fruits contain 6.7 to 7.1 percent sugars (glucose and fructose), 0.8 to 1.7 percent malic acid, 8.3 to 26 mg/100 g of vitamin C, 0.68 to 1.5 percent pectin, and 0.9 to 1.7 percent tannin and pigment substances. Stone kernels contain 37 percent oil and 3 percent amygdalin. Fruits are edible and, after frost, they become sweeter; they are used for production of jams, compotes, wines, vinegar, and kvass. Blackthorn is drought-resistant and is a very polymorphous species. It is a good rootstock for dwarf plum orchards. It is suitable for stabilizing stony slopes of gorges and preventing landslides in the mountains. IV. SMALL AND VINE FRUITS Wild, small fruits in Kazakhstan are represented by 5 families, 9 genera, and 75 species; the most numerous are currant, barberry, rose, and honeysuckle. A. Barberry Barberry (Berberis L., Berberidaceae Juss.) is mainly used as an ornamental bush. It is rather winter-hardy, heat-resistant, light-demanding and resistant to air pollution. These beautiful ornamental plants are used in ornamental borders or as hedges. They bloom abundantly and bear heavy crops of fruit. Insufficient attention has been paid to them as fruit crops, since barberry fruits contain 32 to 35 percent dry substance, 3.9 to 7 percent sugar, 0.6 to 0.8 percent tannin and pigment substances, 0.4 to 0.6 percent pectin and 6 to 7 percent malic acid. Immature fruits contain berberine, a compound used as a styptic in medicine. Berries are used for production of jam, jelly, marmalade and as a filling for pastries. Barberry is also used for production of different kinds of wines, liqueurs, nalivkas, syrups, juices, and extracts. Fruits, leaves, bark, and roots are used to treat urinary diseases, colitis, chronic hepatitis, and gall stones, and as diaphoretic and astringent agents.

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Berberis has 175 species, 9 of which grow in Kazakhstan (Table 3.7) and 3 are endemic. All barberries are branching, thorny shrubs, 2 to 5 m high. Leaves are ovate or elliptical with short petioles. Inflorescences are racemes with few-flowered clusters borne axillary or terminally on abbreviated shoots. Flowers consist of six stamens, one pistil with unilocular ovary and short style, six yellow petals, each with basal nectar glands. Fruits are red-to-black, about 1 cm long, elliptical ovate, and sometimes almost orbicular, with a dry, persistent stigma. They contain one to five oblong seeds with firm brown seed coats. They are propagated by seeds, bush division, layers, and summer cuttings. 1. Bykovskiy Barberry (Berberis bykovianas Pavl.). This endemic species grows in steppe-zones near the mountains, among motley grasses and shrubs. It is a xero-mesophytic shrub, about 2 m high with darkbrown bark and furrowed angular branches with spines. Leaves are oval, entire or with thin, small, cartilaginous denticles, glabrous, and leathery. Inflorescence is a few-flowered, axillary raceme 2.5 cm long, with Table 3.7.

Berberis species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Berberis bykoviana (Bykovskiy barberry)

Steppe zones

Food, ornamental

B. iliensis (Iliyskei barberry)

Dzhungarskei, Terskei Alatau, Ketmentau, valley of the Ili river

Food, ornamental, medicinal plant; not easily cultured

B. integerrima (Integrifolious barberry)

Talasskei Alatau

Ornamental, medicinal, food plant

B. karkaralensis (Karkara barberry)

Central Kazakhstan (Kentau)

Ornamental plant

B. kaschgaria (Kashgar barberry)

Terskei Alatau, Ketmentau

Ornamental, pigment plant

B. nummularia (Monetary barberry)

Talasskei Alatau

Ornamental, medicinal plant; easily cultured

B. oblonga (Oblong barberry)

Kirghiz, Talasskei Alatau, Karatau

Food, industrial plant

B. sibirica (Siberian barberry)

The Altai, Tarbagatai, Dzhungarskei Alatau

Food, honey, ornamental, medicinal plant

B. sphaerocarpa (Spherical-fruited barberry)

The Altai, Tarbagatai, Zailiyskei, Dzhungarskei, Kungei, Terskei, Kirghiz Alatau, Ketmentau

Food, ornamental, medicinal plant; not easily cultured

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two to seven yellow flowers. Berries are dark-blue, with gray bloom, 12 to 15 mm in diameter, spherical, obtuse or pentagonal, strongly flattened at the apex. The species is related to B. sphaerocarpa Kar. et Kir., but is distinguished by its larger petioles. 2. Iliyskei Barberry (Berberis iliensis M. Pop.). This is a rare, endemic species with decreasing natural area. It grows in shrub-lands, on pebbles of river terraces, on stony slopes, in sand and salty soils. It is a stronglybranching, xerophytic, thorny shrub 2 to 3 m high; older branches have gray bark, and the thorny annual shoots are reddish-brown. Leaves are leathery, oblong or lanceolate-lobed, entire on fruiting shoots and subulate-dentate on vegetative shoots, with a well-expressed net of veins on lower sides. Multiflorous inflorescences are axillary racemes, 3 to 5 cm long, with yellow flowers, 3 to 4 mm in diameter, and ovate sepals. Light red berries are oblong-ovate 6 to 7 mm long, 3 to 4 mm wide. This species is preserved in the territories of forest husbandry and in Charyn protected forest in the Sartogai gorge. 3. Integrifolious Barberry (Berberis integerrima Bunge). This species grows on mounds and stony slopes of the mountains. It is a strongly branched, xero-mesophytic shrub, 4 m high with brownish-purple, angular shoots covered by thick, straight thorns, three to a group. Leathery, glabrous leaves are obovate, oblong, entire with short, thin burrs at the apex. Inflorescences are racemes, 6 to 10 cm long with 12 to 25 flowers, each 8 to 9 mm in diameter. Obovate sepals are the same length as the petals. Berries are purplish-red with a bluish bloom; they are oblongobovate, 7 to 8 mm in diameter and have 2 to 3 dark brown seeds. 4. Karkara Barberry (Berberis karkaralensis Korn. et Potap.). This is a very rare species with a diminishing habitat; it grows on stony mountain slopes, among separately growing pines and shrubs, and in meadowsteppe grasslands. Plants are also found as single specimens. It is a mesophytic shrub, 2 m high, with few branches, the older ones are covered by gray bark, and thorny, annual shoots are reddish-brown. Leaves are obovate, slightly leathery, with either dentate or entire margins and well-expressed vein net on the lower sides. Inflorescences are racemes, 3 to 5 cm long, with five to nine yellow flowers and ovate, 5 mm-long sepals. Bright red fruit is obovate without wax bloom. 5. Kashgar Barberry (Berberis kaschgarica Rupr.). This species grows on stony slopes and high-mountain plateaus. It is a low-growing, thorny, spreading, xerophytic shrub about 1 m high, with red-brown shoots and

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splitting bark. Leaves are oblong-ovate, 15 mm long; margins are entire or with fine, thin prickles. Inflorescences are umbellate with two to three flowers in leaf axils, each 6 to 7 mm in diameter. The black berry is wide oval, about 8 mm long, with two to three dark brown, elongated seeds. It may be used in breeding for new frost- and drought-tolerant barberry forms. 6. Monetary Barberry (Berberis nummularia Bunge). This species grows on rocky slopes of the mountains. It is a strongly branched, thorny, xero-mesophytic shrub, 4 m high, with reddish-brown older branches covered with three-part thorns, 2.5 cm long. Younger shoots are bluish. The grayish-green, leathery, glabrous leaves are obovate, entire with small denticles at the apex. Multiflowered inflorescences are axillary racemes, 6 to 7 cm long, 3 to 4 cm wide; flowers have obovate sepals and petals. The light red, inedible berries are obovate-orbicular, 5 to 6 mm long with light-brown, grayish, obovate seeds. 7. Oblong Barberry (Berberis oblonga Schneid.). This species grows on stony slopes of mountains. It is a xero-mesophytic shrub about 4 m high with dark, violet-blackish, cracked bark on older branches, and reddishbrown young shoots. Leathery leaves are elliptical, oblong-obovate, 10 cm long, 4 to 5 cm wide, entire or serrate, often with prickles at the apex; they are grayish-green on the upper sides and bluish on the lower sides. Inflorescences are racemes or panicles, about 15 cm long with 10 to 50 yellow flowers, each 11 mm in diameter. Sepals are orbicular or widelyovate. The violet-black berries, with a bluish bloom, are oblong or elliptical, about 10 to 15 mm long and 5 to 7 mm wide. 8. Siberian Barberry (Berberis sibirica Pall.). This species grows on stony mountain slopes and mounds. It is a xero-mesophytic, thorny shrub about 1 m high, with brownish-red spurs. Leaves with three to six spines on both sides, are oblong-ovate with a cuneate base, and lower sides have well-pronounced, raised venation. The red berries are broadly-orbicular, about 9 mm long and 7 mm wide. 9. Spherical-fruited Barberry (Berberis sphaerocarpa Kar. et Kir.). This species grows on mountain slopes, among shrubs, in gorges and valleys of mountain rivers and streams. It is a xero-mesophytic shrub about 2 m high, with gray, older branches, and brownish-red younger shoots. Glabrous, gray-green or bluish leaves are 7.5 cm long, 4 cm wide, obovate, entire, with pronounced, raised venation on the lower sides. Inflorescences are racemes with five to nine yellow flowers. The violet-black

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berries with bluish bloom, are spherical-oval, about 12 mm in diameter. Seeds have seed coats with reticulate surfaces. B. Currant Currant (Ribes L., Grossulariaceae DC.) is one of the main small fruit crops of the Northern Hemisphere. Its berries enjoy wide popularity both as a dessert and as an antiscorbutic product. Ribes includes more than 150 species distributed in cold and temperate climate zones of Europe, Asia, and North America. Tasty fruits distinguish most of the wild species that are ancestors of existing cultivars. The widespread cultivated currants include two sub-genera, black currant in the subgenus Coreosma and red currant in the subgenus Ribesia. All species are shrubs with exfoliating bark on old branches and smooth bark on young shoots. Leaves are alternate, serrate-lobate, toothed, and densely pubescent, sometimes with glands. The dioecious flowers are in racemes. Fruit is juicy and multiseeded. The hard seeds are covered with a gelatinous coating. The dried hypanthium and perianth are persistent at the apical end of the fruit. The plants are propagated by seeds and vegetatively, they are frost-resistant and moisture-loving. In open places fruits are borne more abundantly than in shaded areas, although it is rather shade resistant. Black currant berries have a natural, complex vitamin content. They contain 7 to 10 percent sugars, 4 percent organic acids, 0.68 to 1.02 percent pectin which promotes processing of jelly, 400 mg/100 g vitamin C, 500 mg/100 g vitamin P (a vitamin C derivative), 0.7 mg/100 g pro-vitamin A, and 0.06 mg/100 g vitamin B. Berries, leaves, buds, and young shoots are used for medicinal purposes. Leaves, flowers and buds contain a valuable ether-soluble oil. Currant is a valuable small-fruit shrub; it is also used for ornamental purposes in urban landscapes. Ten currant species grow in Kazakhstan (see Table 3.8). 1. High Currant (Ribes altissimum Turcz. ex Pojark.). This species grows on coarse-stone hillocks within the forest belt of the Altai mountain ranges. Shrub is 2 to 3 m high, with brown or reddish-brown bark that splits and separates into long shedding strips. Shoots range from glabrous to glandular-bristly. Leaves are trilobate with a shallow cordate base, 3 to 6 cm in length and width, with dark-green upper sides and light green lower sides. Both sides are glabrous or on the under side, glabrous only along the veins. Inflorescences are 2.5 to 6 cm long and have 7 to 26 bell-shaped flowers with dull purple spots and recurved sepals. Tasty berries are black-purple with purple juice, and 5 to 7 mm in diameter. Plants are exceptionally winter-hardy, and resistant to pests

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Ribes species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Ribes altissimum (High currant)

The Altai

Exceptionally winterhardy, resistant to diseases and pests. Food and vitamin plant.

R. atropurpureum (Dark-purple currant)

The Altai, Tarbagatai

Potential for breeding cultivars with large berries and vigorous shrubs but demanding in culture.

R. graveolens (Odorous currant)

The Altai

Freely crosses with black currant; potential for breeding of winter-hardy cultivars; ornamental plant.

R. heterotrichum (Different-pubescent currant)

The Altai, Tarbagatai, Zailiyskai, Dzhungarskei, Terskei, Kirghiz Alatau, Ketmentau

Potential for breeding drought-resistant, frostresistant cultivars; demanding in culture.

R. hispidulum (Hispid currant)

The Altai, Tarbagatai and Dzhungarskei Alatau

Potential for use in breeding; demanding in culture.

R janczwskii (Yanchevskiy currant)

Zailiyskei, Kungei, Terskei, Kirghiz, Talasskei Alatau, Ketmentau, Karatau

Demanding in culture.

R. meyeri (Meyer’s currant)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei Alatau, Ketmentau

Drought-resistant, heavy bearing, high vitamin C content; demanding in culture.

R. nigrum (Black currant)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei Alatau, Ketmentau

The ancestor of the most black currant cultivars; demanding in culture.

R. saxatile (Rock currant)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei Alatau

Berries are not edible. Demanding in culture.

The Altai, Dzhungarskei Alatau

Endemic species. Food, honey, vitamin plant.

European species

The ancestor of the most cultivars of red and white current; cultivated in Kazakhstan.

R. turbinatum (Turbinate currant) R. vulgare (Red currant)

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and diseases. They are heavy bearing and have a high content of vitamins P (2700 mg/100 g) and C (38–47 mg/100 g). This species deserves the attention of plant breeders. 2. Dark-Purple Currant (Ribes atropurpureum C.A. Mey.). This species grows along shady slopes of forest belts in the mountains, on riverbanks and in forest zones on the plains. Shrub is 1 m high with greyish-yellow bark and brown glabrous shoots. Leaves are orbicular-cordate, thin, glabrous on upper sides and densely-pubescent on undersides. There are three-to-five lobate with acute, well-developed upper lobes; leaf-margins are coarsely, double-toothed. Inflorescences are 3 to 4 cm long with four to six bell-shaped purple flowers, 4 to 5 mm long. Red berries are quite large at 11 to 13 mm in diameter. It has the potential for breeding vigorous, resistant, large-fruited cultivars. It is recommended for use in culture. 3. Odorous Currant (Ribes graveolens Bunge). This species grows in subalpine mountain belts on stony slopes and deposits, on rocks and glacial moraines in the Altai. It is a multibranched shrub 30 to 70 cm high with light-gray bark. All plant parts, including lower sides of leaves, are dotted with glossy, yellow glands. Leaves are small, 1 to 3 cm wide, abruptly trifoliate, thick, glabrous on upper sides with glossy, yellow glands and sparse, white pubescence on the lower sides. Inflorescences have four to ten white flowers, each 2 to 4 cm long with fan-shaped petals. The calyx tube is covered with dotted glandules. Fragrant berries are spherical, 8 to 10 mm in diameter, red-brown, smooth, and with glands on the surface. 4. Different-pubescent Currant (Ribes heterotrichum C.A.M.). This species grows on rock mounds, stony slopes in forest-steppe and subalpine mountain belts and foothills of the Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz Alatau, and Ketmentau. It is a multi-branched, low-growing shrub 50 to 90 cm high with grayishbrown separating bark on older branches and young shoots are either glabrous or densely-white pubescent. Leaves are small, 1 to 3 cm wide, orbicular, or orbicular-reniform, thick, with wide-cuneiform base, both sides glabrous or slightly pubescent, with scattered, sticky glands. Leaves have three wide-orbicular lobes with coarsely dentate margins. Densely pubescent leaf-petioles are shorter than leaf-blades. Upright inflorescences are racemose, multiflowered, 2.5 to 5 cm long, with dull purple flowers that are patelliform pubescent, about 6 mm in diameter with wide-ovate calyx lobes. Berries are orbicular, orange-yellow, pubes-

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cent or glabrous, without glands, 5 to 7 mm in diameter, sweetish-acidic or with strong bitter flavor. This species may be useful for breeding of drought-resistant and frost-resistant forms. Fruit flavor is not pleasant but is suitable for processing of fruit wines and drinks and is useful as a honey plant. 5. Hispid Currant [Ribes hispidulum (Jancz.) Pojark.]. This species grows in moist forests, on forest edges, along riverbanks, among shrubs, near swamps, on the northern plains, the Altai, Tarbagatai, and Dzhungarskei Alatau. It is a shrub 1.5 to 2 m high. Bark on older branches is dark-brown or grayish and pale-gray, while young shoots are glabrous or pubescent and covered with stalked glands. Leaves have three to five wide lobes with glabrous or sparsely pubescent upper sides and mostlypubescent lower sides. Inflorescences are 3 to 7 cm long with 3 to 12 pale yellow flowers, 5 to 6 mm in diameter. Inflorescence rachis and pedicels are sparsely pubescent and covered by stalked glands. Berries are red, orbicular 4 to 12 mm in diameter with acidic flavor. Vitamin C is 60 to 100 mg/100 g, seeds contain 13.4 to 15 percent oil, and leaves, 0.11 percent carotene. It is used for production of juices, jams, and wines. It is distinguished by berries ripening earlier than those of black currant and is used as a food, oil, vitamin, and honey plant. It is not a demanding plant in culture and is easily crossed with other species of red currant. 6. Yanchevskiy Currant (Ribes janczewskii Pojark.). This species is comparatively rare, growing in forest belts of the mountains on stony slopes, on hillocks, in gorges and river valleys. It is a shrub 1 to 1.5 m in height, with straight branches; older branches are grayish-brown with red blush; younger shoots are light-gray, glabrous, and current-year shoots are red-yellow and glabrous. Leaves are three-to-five-lobate with glabrous upper sides and the lower sides dotted with yellow aromatic glands. Leaf lobes are ovate, margins dentate, and apices acute. Inflorescence is a 6 to 10 cm long raceme with five to ten flesh-colored flowers that are 8 to 9 mm in diameter; hypanthium is campanulate; sepals are obtuse, and petals are widely-ovate. Berries are large, 10 to 13 cm in diameter, black, sourish-sweet, aromatic, with flavor superior to that of fruits of other wild currant species. Vitamin C content is 140 to 150 mg/100 g. It is of great interest for breeding and should be preserved. 7. Meyer’s Currant (Ribes meyeri Maxim.). This species grows in forest-steppe and subalpine mountain belts among shrubs on stony slopes, in gorges, and along riverbanks. It is a shrub 1.5 m high with glossy gray-brown bark. Older shoots are dark-gray while current-year

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shoots are brownish-green, glabrous or slightly pubescent, and glandular. Leaves are three- to five-lobate; on fruit-bearing shoots they are 2.5 to 5 cm in diameter, and on vegetative shoots, up to 6 cm, orbicular and slightly heart-shaped. The inflorescences are drooping racemes, 5 to 9 cm long with 4 to 12 flowers. Small, brownish-green flowers with purple spots and with glands are 3 mm in diameter and have 5 stamens. Petals are brownish-green, lobate with ciliate margins, and style is cylindrical, not widened at the base. It is distinguished by heavy bearing with violetblack berries, 7 to 8 mm in diameter. Vitamin C content is 140 mg/100 g. 8. Black Currant (Ribes nigrum L.). This species grows along moist valleys and meadows, riverbanks, near swamps, on stony slopes and hillocks within the forest belt in the mountains. It is a shrub, 1 to 1.5 m high. Bark on older branches is dark brown and on younger shoots is grayish-yellow. Leaves are five-lobate, 10 to 12 cm wide with dark green, slightly pubescent upper sides while lower sides are lighter green and dotted with golden-yellow, odorous glands. Pinkish-gray flowers are 7 to 9 mm long with dotted glands on the outside. Sepals are recurved, acute, rather wide, and petals are one-third shorter than sepals. Black, fragrant berries are orbicular, about 10 mm in diameter. They contain 100 to 300 mg/100 g of vitamin C, 8 to 10 percent sugars, 0.5 to 1 percent acids, and 0.68 to 1.02 percent of pectin substances which contributes to high jellying ability. In ascorbic acid content, black currant is inferior only to wild rose and Actinidia. This species is the ancestor of almost all black currant cultivars. Berries are used fresh and for processed products (jam, jelly, juices, wines, etc.). 9. Rock Currant (Ribes saxatile Pall.). This species grows on rocks, hillocks, stony slopes, and on rocky riverbanks of the low-hill plains, in foothills and low mountains of Kazakhstan. It is a low spreading shrub 50 to 90 cm high with grayish-brown bark on older branches which is usually pubescent, although sometimes glabrous. Thorns are usually present at the nodes and often also on internodes. Leaves are small, 1 to 1.5 cm long, bluish-green, firm, orbicular-obovate with a wide, cuneate base, shallowly trilobate or with three large-lobed teeth. Inflorescences are racemes, top-pointed, 2 to 3 cm long with small, greenish, discshaped flowers about 4 mm in diameter and with ovate sepals. Dark-red, smooth berries are orbicular, 6 to 7 mm in diameter. Blooming and fruit bearing are usually heavy. 10. Turbinate Currant (Ribes turbinatum Pojark.). This is an endemic species that grows among shrub lands, along riverbanks. It is an upright

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shrub with brown-yellow, pubescent young shoots that turn gray as they age. Glabrous, glossy, acute-lobate and acute-dentate leaves are 10 cm wide with yellow aromatic glands near the base. Inflorescences are racemes about 3 cm long, with four to seven flesh-colored flowers that are 9 mm long. Sepals are wide, obtuse, sometimes overlapping each other. Black, orbicular berries have an aromatic flavor similar to that of black currant, but without the distinctive aroma of the latter. 11. Red Currant (Ribes vulgare Lam.). This is a shrub 1 to 1.5 m high with bark on old branches peeling off in scales or oblong fibers while young shoots are straight and light-grayish. Leaves are three- to fivelobate, with upper sides glabrous and lower sides, pubescent, without glands, and margins dentate. Leaf bases are shallowly cordate. Inflorescences are racemes about 8 cm long with glabrous rachis and pedicels. Flowers are greenish with glabrous, recurved sepals. Red berries contain high levels of sugar and organic acids. This species is cultivated and is the ancestor of most varieties of red and white currant. C. Gooseberry Cultivated gooseberries (Grossularia Mill, Grossulariaceae) are widely distributed in northern Europe. Fruits are used fresh or as processed products. Grossularia includes more than 50 species, distributed throughout the Northern Hemisphere. Gooseberry is rather winter-hardy, though in severe winters without snow cover, there is some freeze damage. The vegetative period begins earlier than in other small-fruits, and berries ripen in the middle of June. Gooseberry does not form root shoots, but sprouts arise from underground stems. Acicular gooseberry (Grossularia acicularis (Smith.) Spach.) is the only species grown in Kazakhstan. It is distributed in The Altai, Tarbagarai, Dzhungarskei Alatau, Kokshetau, and Eastern low mounds. It grows on stony slopes of middle- and low-mountain belts. It is a shrub about 1 m tall with arching shoots densely covered with needle-shaped thin spines and light gray bark. Leaves are three- or five-lobate, 0.7 to 3 cm wide, glabrous, glossy. Inflorescences are racemes with greenishwhite or pink, sessile flowers. Glabrous berries are 1 to 1.5 cm in diameter, oval-orbicular or orbicular, seldom glandular and have good flavor. Berries contain sugars, organic acids, pectin, nitrogenous substances, vitamins C, B, B2, and A. They are used fresh and for preparation of jams, juices and compotes. High acid and considerable content of amide and ammoniac compounds in berries allow them to be used widely in wine making. Berries are recommended for use as medicines against kidney,

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urinary bladder, and gastrointestinal diseases, against some skin diseases, avitaminosis and blood vessel obstruction. It is a food and honey plant and is easily cultivated. D. Grape Grape (Vitis L., Vitaceae Juss.), one of the most ancient, popular, and useful fruit plants in the world, is a mostly deciduous, perennial woody vine (liana). The genus includes about 70 species, growing mainly in warm and temperate belts of the Northern Hemisphere. It is cultivated on all continents except the polar regions. Stems reach 20 to 40 m with long thin annual shoots and thick root systems that penetrate the soil to a depth of 7 m. Flowers are bisexual or dioecious, in panicle. Both crossand-self pollination occur. Fruits are juicy berries of different shapes and color, borne in clusters. Berries contain 10 to 33 percent sugar, 0.5 to 1.4 percent organic acids, 0.3 to 0.5 percent mineral substances, 0.3 to 1.5 percent pectin substances, vitamin C, group of B vitamins, and provitamin A. Berries are used fresh, dried (seedless grape, raisin), preserved, or processed in different kinds of wines (Champagne, table, dessert, cognac), juices, jams, stewed fruits, and marmalade. It has also been shown that grapes have beneficial medicinal properties. It is propagated by seeds, and in culture, mainly by cuttings and layers. A unique community of wild grape (V. vinifera L.) grows in southern Kazakhstan, the most northern area of wild grape in Central Asia. It grows on terraces of the mountain rivers with shallow, subsoil waters, and on stony mounds. It usually forms small groups among wood and shrub vegetation in the areas of Karatau 9 ridge, Boroldai mountains, gorges of Bosturgai, Bugun, Kok-Bulak, Kaurchkty, spurs of Talasskei Alatau (Mashat mountains) and the western extremity of Zailiyskei Alatau (Karakunuz gorge). The perennial liana grows prostrate on the ground or climbs with the help of tendrils along tree stems to 25 m, from where clusters of grape berries hang down. The diameter of the stem reaches 15 to 20 cm. Bark of old stems is brown, multilayered; young, hardened shoots are light brown with thin furrows. Wood is dark-brown, light, porous, with poorly differentiated heart and sapwood. Leaves are lobate with rounded outlines 5 to 20 cm in diameter, entire, three- to five-lobate or dissected. Leaf surface is either smooth or wrinkled, always with welldistinguished veins colored more intensely. Leaf size and shape of wild grape vary even within one bush. Autumn leaf color is diverse, from bright green to bright-yellow and purple-red. Abundant, late-blooming flowers (after full leaf development) are small, in loose or dense panicles. Fruit is a juicy berry, strongly varying in size, color and flavor.

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Most of berries are small (1.2 cm long, 0.7 cm wide) though larger berries can be found, to 1.5 cm long and 1.3 cm wide. Berry weight is from 1.0 to 3.2 g. Berry shapes are rounded, slightly flattened, oval, oblong, ovate, obovate and almost always symmetrical. Color is blackviolet, purple, dark-red to pink, yellowish-green or green. Berry flesh is very juicy, sweet or sour-sweet. Skin is rather coarse, thick, and with a well-expressed waxy bloom. Each berry has one to four seeds, although seedless berries are sometimes noted. Seed shape is pyriform or ovate with long beak. Clusters, which can be loose or very compact, have very diverse shapes from branching, winged, to cylindrical and conical. Cluster length is 5 to 25 cm, with from 10 to 90 berries per cluster. Fruit stem at the place of attachment is widened to a peculiar cushion; the attachment of berry to stem is always firm. In southern Kazakhstan, wild grape blooming begins in May to June, with fruit ripening from mid-August to the end of September. Berries of wild grape in the Karatau mountains accumulate about 11.4 percent of sugars and 1.95 percent organic acids. In natural conditions, wild grape is propagated mainly vegetatively, by rooting of stems; both young and old canes root well. Seedlings are seldom found, only on well-watered plots along banks of mountain rivers. Infection of wild grape by pests and diseases is not recorded. Wild “Kazakhstan” grape is highly frost-resistant and drought enduring. Therefore, it is recommended to be used as initial material for breeding. The preservation of all relict habitats of wild grape in Kazakhstan is necessary. E. Raspberry and Blackberry Brambles (Rubus L., Rosaceae) are popular small-fruit crops, famous for food and medicinal properties. Cultivars are grown everywhere there is suitable climate and soil. The cultivated brambles are considered inferior to the wild species in flavor, aroma, and medicinal properties. There are more than 600 wild species in this genus. They are distributed in all continents of the world from subpolar to temperate, subtropical and tropical regions. Wild raspberry is a typical forest or mountain-forest plant. In the Kazakh Republic, it grows on mountain slopes, riverbanks, in forest mounds of Northern Kazakhstan and in mountains from the Altai to Kungei Alatau. Raspberry populations are characteristic of cutover and burnt-over areas in the Altai taiga. Three to five years after these events, raspberry entirely covers the cleared lands that subsequently revert to forests. Raspberry populations are usually found on stony deposits in the forest belt of Northern Tien-Shan and Dzhungarskei Alatau. Four bramble species grow in Kazakhstan (see Table 3.9).

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A. DZHANGALIEV, T. SALOVA, AND P. TUREKHANOVA Rubus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Rubus idaeus (Common raspberry)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei Alatau, Ketmentau

Food, vitamin, medicinal, honey plant; easily cultured

R. sachalinensis (Sakhalin raspberry)

The Tarbagatai, Altai, Dzhungarskei Alatau

Food, vitamin, medicinal, honey plant

R. caesius (Blackberry)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz Alatau, Ketmentau, Karatau

Food vitamin, honey, pigment plant; easily cultured

R. saxatilis (Stone berry)

Tarbagatai, Dzhungarskei, Zailiyskei Alatau, Ketmentau

Food plant; not easily cultured

1. Common Raspberry (Rubus idaeus L.). This species grows on mountain slopes in the understory of open forests, along riverbanks, on forest borders and cut-over areas. It is a semi-shrub about 1.5 m high. Annual shoots are straight with drooping apex and reddish-brown thin prickles. Leaves are ternate, dark-green on upper sides with white, dense pubescence on lower sides. Inflorescences are in cymes with white flowers, and grayish-green sepals recurved from berry base. Red berries are aggregates of drupelets, often pubescent, orbicular or round-conical, juicy, and sweet with small, round stones. Plants are light-demanding and require good fertile, well-drained soils. They usually over-winter under snow cover. Due to high content of sugars (6–11%), organic acids, mineral salts, and aromatic substances, raspberry fruits are distinguished by high flavor and nutrient qualities. They are high in vitamins and ascorbic acid. Berries are used fresh and for preparation of jams, nalivka, nastoyka, juices, and syrups. Dried raspberry is used in medical treatment, especially for catarrah diseases. 2. Sakhalin Raspberry (Rubus sachalinensis Leul.). This species grows in stony places in mountains, forests, in openings among forest trees, and in shrub-lands. It is a semi-shrub about 1 m high. Shoots have a bluish bloom and are heavily covered with yellowish-brown or reddish thorns. Leaves are usually ternate, ovate-cordate or oblong-lanceolate, with upper sides glabrous and lower sides white-or-gray tomentose. White flowers are in few-flowered, terminal or axillary inflorescences. Pedicels

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on floriferous shoots are pubescent, thickly or thinly covered with thorns and glands. The edible, sweet, red berries are covered with fine, white tomentum. Stone surfaces are reticulate. 3. Blackberry (Rubus caesius L.). This species grows in forests, gullies, in shrub-lands, along roads, riverbanks, and in wet meadows. Shrubs are 0.5 to 1.5 m tall and shoots are covered with whitish bloom and numerous thorns. Cylindrical, bluish primocanes with straight or recurved thorns are arching, branched and root at their apices in autumn. Floricanes are long, and more upright. Leaves are ternate with widelanceolate stipules. Inflorescences are corymbs with five to ten white, 3 cm flowers with elliptical petals. The green, pubescent, often glandular sepals adhere to the fruits. The blackberries are 1.5 to 2 cm long and 1.5 cm wide, in clusters of 5 to 22 aggregates of drupelets covered with bluish bloom. Fruits of blackberry are used fresh and for production of jam, compote, and kissel. They contain organic acids, vitamin C, glucose, sucrose, and fructose. The citric acid content is somewhat higher than sugar, therefore, the berries are always sour. In folk medicine, berries, leaves and roots are used as an astringent substance. Pigments extracted from the berries have industrial uses, such as in food coloring. Shoots and leaves are used in tanning hides. 4. Stone Berry (Rubus saxatilis L.). This species grows in glades among forests and on forest borders, on stony slopes of mountains, and in swamps. It is a perennial, herbaceous plant 10 to 15 cm high with upright floricanes and 40 to 100 cm long procumbent primocanes. Hairs and thorns cover stems and pedicels. Leaves are ternate with long petioles and thin-needle-shaped thorns on the under-sides. Inflorescences are corymbs with three to ten white flowers. Sepals are lanceolate, recurved, and slightly pubescent. Berries are bright- or dark-red, weakly accreted among themselves, about 5 mm in diameter, of acid-sweet flavor. Berries contain sugar, organic acids, vitamin C, tannin, and pectin substances. They are used fresh and for production of jam and berry juice. F. Strawberry Cultivated strawberries (Fragaria L., Rosaceae) are widely distributed. Fruits are used fresh, and they are exceptionally valuable for processed products like jam, jelly, and beverages. Berries have an excellent flavor and aroma, and ripen earlier than other berries and fruits. They are an important source of natural vitamins. This genus includes about 50

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species, all but one of which is in the Northern Hemisphere. The sole exception is Fragaria chiloensis (L.) Dieck., the Chilean strawberry of South America. Strawberry is a perennial, herbaceous, nonwinter-hardy plant, although it winters well under snow cover. It is light-demanding, though endures a little shading, moisture-loving, adapted to weak acidity and soil alkalinity. Two strawberry species grow in Kazakhstan (see Table 3.10). 1. Alpine Strawberry (Fragaria vesca L.). This species grows on forest borders and in glades among forests. It is a perennial, herbaceous plant 10 to 20 cm high. It has horizontal rhizomes covered with brown, dead stipules. Roots and long-creeping shoots that root at nodes grow from the rhizome. Ternate radical leaves consist of ovate leaflets, with 8 to 13 teeth, the upper tooth is slightly larger than the adjacent. Leaves are darkgreen on the upper sides and grayish-green, pubescent on the lower sides. Bisexual, white flowers are about 2 cm in diameter. Sepals are acuminate. Bright-red berries are ovate, oblong-conic, about 1 cm long with delectable flavor and aroma. Strawberry is propagated by stolons, seeds, and plant division. Fruits contain sugars, organic acids, tannin, mineral substances, ether oils, and vitamin C. The content of ascorbic acid is 16 to 20 mg/100 g. There is 19 percent oil in seeds and about 9.4 percent tannic substances in roots. It has potential for breeding winterhardy, aromatic berry cultivars, resistant to diseases and pests. 2. Green Strawberry (Fragaria viridis Duch.). This species grows on open, grassy, mountain slopes, on borders and in glades of mountain forests, among aspen-birch trees, in meadows and meadow steppes. It is a perennial, herbaceous plant about 25 cm high with an upright pubesTable 3.10.

Fragaria species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Fragaria vesca (Alpine strawberry)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz, Talasskei Alatau, Ketmentau, Karatau

Food, vitamin, medicinal plant; easily cultured

F. viridis Duch. (Green strawberry)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Kirghiz, Talasskei Alatau, Karatau

Food, medicinal, honey plant; easily cultured

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cent stem. Wooly-hairy leaves are ternate with obliquely-ovate leaflets with triangular teeth. The terminal leaflet is very small and suppressed between lateral leaflets. Inflorescence is a corymb with white flowers about 2.5 cm in diameter. Leaves on flowering shoots have appressed pubescence. G. Vacciniums In Kazakhstan, various species of Vaccinium L. (Ericaceae Juss.), are indigenous including bilberry, bog bilberry, lingonberry, and cranberry (Table 3.11). The blue fruited Vaccinium native to Kazakhstan include bilberry, bog bilberry, and lingonberry. The dark-red fruited cranberry is a small evergreen shrub with thin, decumbent stems 80 to 100 cm long with small, leathery leaves and reddish flowers. Berries are juicy, sour, with a little bitter taste. Cranberry fruits are valuable food products containing: 2.3 to 5.0 percent sugars, 2.5 to 3.6 percent organic acids, 1.3 percent pectin substances, and 50 mg/100 g vitamin C. The presence in berry juice of free benzoic acid provides long storage of berries and juices. It is mainly propagated by rooting of stems. In cold-temperate, northern latitudes of Eurasia and North America, there are four cranberry species; two of these are endemic to Kazakhstan. 1. Bilberry (Vaccinium myrtillus L.). This species grows in coniferous forests on moist soils with well-developed humus horizon and often dominates in the grass-shrub belt. It is a deciduous shrub 50 to 100 cm

Table 3.11.

Vaccinium species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Vaccinium myrtillus (Bilberry)

The Altai, Tarbagatai

Food, medicinal, plant

V. uliginosum (Bog Bilberry)

The Altai

Food, medicinal plant

V. vitis-idaea (Foxberry, cowberry, lingonberry)

The Altai, Tarbagatai

Food, medicinal, technical, honey plant; easily cultured

V. microcarpus (Microcarpous cranberry)

The Altai

Medicinal plant

V. palustris (Quadripetalous cranberry)

The Altai

Food, medicinal plant

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high with greenish bark. It has rhizomes with long, creeping, numerous, strongly branching roots. The fruit-bearing shoots are erect, green, and acute-edged. Light-green, glabrous leaves are 5 to 38 mm long and 2 to 24 mm wide, ovate or elliptical, serrulate, obtuse or pointed on very short petioles. Green-pinkish flowers are borne singly on short pedicels, with ascidiform-globular ovary that is pentilocular. The black, globular berry with bluish bloom and reddish flesh, is 6 to 8 mm in diameter, and flavor is sourish-sweet. Vegetative propagation predominates over seed propagation. Bilberry fruits contain 5 to 20 percent sugar, 7 percent organic acids, 12 percent tannin substances, 10 to 75 mg/100 g carotene, 6 mg/100 g vitamin C, and seeds accumulate about 30 percent of oil. Berries are used fresh or dry, for preparation of kissel, compotes, and jams. In medicine, they are used against stomach, bronchial, angina, and anemic diseases. 2. Bog Bilberry (Vaccinium uliginosum L.). This species grows on diverse soils, in wet-coniferous forests, in swamps, and sometimes in shrub-lands. It is a light-demanding, moisture-loving, deciduous shrub 0.25 to 1.4 m high with brown or dark-gray bark. Glabrous leaves are 5 to 38 mm long and 4 to 24 mm wide, thin, firm, obovate or oblong, lightgreen on upper sides and bluish on lower sides. White-pinkish flowers are small, drooping, and ascidiform with a slight pleasant aroma. Blue fruit with prominent bloom is an orbicular berry, four- to five-locular, 9 to 12 mm in diameter, with green flesh. Propagated mainly by seeds, it has endotrophic mycorrhiza. Fruits contain 5.6 percent sugar, 1.65 percent organic acid, cellulose, pectin, tannin and pigment substances, and 30 mg/100 g of vitamins A, P, C; mature fruits are a well-known antioxidant agent. Fruits are used fresh and dry, for preparation of kissels, compotes, jams, and wines. 3. Lingonberry (Foxberry or Cowberry) (Vaccinium vitis-idaea L.). This species grows in coniferous, mixed, and deciduous forests, on poor dry soils and on peat. It is a long-lived semi-shrub 30 cm high with underground rhizomes. Evergreen leaves are 10 to 30 mm long, 7 to 12 mm wide, elliptical or obovate, alternate, thick, leathery, with upper sides dark-green and lower sides pale-green with dark-brown scattered glands. The bell-shaped, pale pink flowers are borne in 2 to 8 filamented racemes. These develop into bright-red, globular berries. The plant is very winter-hardy and begins bearing in the third year. On moist soils and slopes of northern exposure, yields are much higher (about 300kg/ha) than on southern slopes. It is propagated mainly vegetatively, by rhizomes and also by seeds. Berries contain: 8.8 percent sugars, 2.5

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percent organic acids, 0.25 percent tannin substances, 1.8 percent cellulose, 18 mg/100 g vitamin C, and seeds accumulate about 23 percent oil. Berries are used fresh, for preparation of jams, kissels, syrups, and compotes. Due to the content of benzoic acid-antiseptic, both fresh and processed products have a longer shelf life. Leaves contain high amounts of tannin substances. 4. Microcarpous Cranberry (Vaccinium microcarpus L.). This species is an evergreen semi-shrub with thin creeping stems that grows in sphagnum bogs. Small leaves, 3 to 7.5 mm long, 1 to 2.5 mm wide, are leathery, acuminate at the apex, oblong-ovate with curled margins. Upper sides are dark-green and lower sides are bluish due to waxy bloom. Small, pink flowers develop into red, globular berries, 5 to 10 mm in diameter. 5. Quadripetalous Cranberry (Vaccinium palustris L.). This species grows only in swamps, mainly in sphagnum bogs, often forming vast colonies, especially in open, well-lighted areas. It is a creeping, low bush with thin shoots. Small leaves, 8 to 16 mm long, 3 to 6 mm wide, are leathery, acuminate, ovate or oblong-ovate, grayish on lower sides and dark-green on upper sides. Red flowers are drooping in umbellate inflorescences. Ruby-red fruits, 10 to 18 mm in diameter, are globular or oblong-ovate berries, juicy, sour, and edible. V. OTHER FRUITS A number of other species with edible fruit are indigenous to Kazakhstan. These include elderberry, honeysuckle, mulberry, oleaster, rose, sea buckthorn, and viburnum. These are often better known for other uses. For example, viburnum, oleaster, and rose are best known as ornamental or landscape plants, while mulberry is best known as a food source for silkworm. A. Elderberry The genus Sambucus L. (Caprifoliaceae) has 40 species, distributed in countries with temperate and subtropical climate in both hemispheres. One species is known in Kazakhstan. Siberian elder (Sambucus sibirica Nakai) grows in coniferous and mixed forests, on slash-fire lands and cut-over areas, along gully slopes and riverbanks, in gorges and northern slopes of eastern foothills of Arkat mountain, and in the Altai and Tarbagatai.

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It is a densely-branched, deciduous shrub 2 to 4 m high with bark of older branches reddish-brown, thin, and wrinkled young branches are light-brown or violet and covered with lenticels; young shoots are pubescent. Leaves are pinnately compound with 5 to 7 lanceolate, ovatelanceolate, acuminate leaflets, each 5 to 14 cm long, 1.5 to 5.5 cm wide; margins are serrate-toothed or serrate, green on upper surfaces and light green on lower surfaces. Flowers are whitish-green or yellowish, in corymbose inflorescences. Oval drupaceous fruits are bright-red, 3 to 4 mm long. Stones are narrow-elliptical, light-brown with a high quantity of oil. It is propagated by seeds. It has been used as a medicinal, honey, and ornamental plant. B. Honeysuckle Honeysuckle (Lonicera L., Caprifoliaceae Juss.) is usually considered an ornamental, but some have edible fruit. There are 21 species of Lonicera in Kazakhstan: Albert honeysuckle (Lonicera alberti Regel.); Altai honeysuckle (L. altaica Pall. et DC.); Altman honeysuckle (L. altmanii Regel et Schmalh.); grey honeysuckle (L. cinerea Pojark.); hispid honeysuckle (L. hispida Pall. Schult.); low honeysuckle (L. humulis Kar. et Kir.); Ili honeysuckle (L. iliensis Pojark.); Karatav honeysuckle ( L. karatauiensis Pavl.); Kareh honeysuckle (L. karelinii Bunge ex P. Kir.); Korotkov honeysuckle (L. korotkowii Stapf.); parviflorous honeysuckle (L. micrantha (Trautv.) Regel.); microphyllous honeysuckle (L. microphylla Willd. ex Schulf.); monetary-phyllous honeysuckle (L. nummularifolia Jaub. et Spach); olga honeysuckle (L. olgae Regel. et Schmalh.); pallas honeysuckle (L. pallasii Ledeb.); Popov honeysuckle (L. popovii Golosk.); Semenov honeysuckle (L. semtnovii Regel.); imitating honeysuckle (L. simultatrix Pojark.); narrow-flowered honeysuckle (L. stenantha Pojark.); Tartar honeysuckle (L. tatarica L.); and Tien-Shan honeysuckle (L. tienschanica Pojark.). While Kazakhstan honeysuckles are ornamental plants, three species have edible fruits: Altai honeysuckle (L. altaica Pall. et DC.); Ili honeysuckle (L. iliensis Pojark.); and pallas honeysuckle (L. pallasii Ledeb.). C. Mulberry Mulberry (Morus L., Moraceae Link.) includes 24 species, growing mainly in warm temperate regions of Asia, but also some in the United States and Africa. Its origin is considered to be China. Mulberry has an aggregate fruit produced on a tree whose leaves are used to feed silkworms. Mulberry fruits are also well-known to have some value as a food product. Fruits can be used fresh and dried. They contain 11 to 12 per-

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cent sugars (mainly fructose and glucose); organic acids, 0.6 to 1.5 percent; cellulose, 1.7 percent; tannin substances, 0.08 percent; and nitrogenous substances, 1.5 percent. Mulberry fruits are also used for preparation of bekmes (artificial honey), and for waste-vinegar. Only one very polymorphic species, the white mulberry (M. alba L.), grows naturally in Kazakhstan. Two relict populations are known in the republic; in Syrdaria Karatau (Boroldaitau) and in the western part of Zailiyskei Alatau (Karakunuz gorge). Here, white mulberry forms small groves that are situated exclusively on river alluvial plains. White mulberry is a deciduous tree 15 to 20 m tall, with a very dense, spreading, globular, sometimes weeping crown. Shoots are clearly of two types: vegetative are oblong with large leaves and fruiting spurs are much shortened with smaller leaves. Young shoots are light-green and downy. Leaf shapes are variable; within the same tree, even on the same branch, they may be entire, symmetrical or asymmetrical, or lobate with 3 to 11 lobes incised to different depths. They are 6 to 15 cm long, ovate or wide ovate heart-shaped at the base and acute at the apex. In summer, leaves are dark-green, and in autumn, straw-yellow. The plant is dioecious with flowers in cylindrical, catkinlike inflorescences appearing simultaneously with leaves. Bloom occurs in April to May, and fruits mature in June to July. Easily detachable fruit is an aggregate of drupelets, 0.7 wide by 4.0 cm long with cylindrical, conic or orbicular shape, similar to blackberry. Its color is white, yellow, pink to violet, with a sweet flavor. The species is very polymorphous. White mulberry is winter-hardy, drought-resistant, and salt-enduring. It is relatively undemanding to soil, though it grows best on porous, fertile, sandy loam, and loam soils. When young trees grow rapidly they can live up to 200 years. Trees are propagated by seeds, sprouts, and in culture, also by cuttings and layers. It can be considered as an ornamental, food, and medicinal plant. D. Oleaster Oleaster (Elaeagnus L., Elaeagnaceae) consists of 40 species worldwide, three of which occur in Kazakhstan (Table 3.12). Most are found in temperate latitudes, except for a few that grow in the subtropics of Asia and the Mediterranean region. 1. Narrow-leafed Oleaster, Russian Olive (Elaeagnus angustifolia L.). This species grows mainly in Turgai forest along the banks and in valleys of desert rivers, among alluvial shrubs, forest-borders, sands, borders and lower sides of sandy hills and steppes. It is a shrub or small tree,

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352 Table 3.12.

A. DZHANGALIEV, T. SALOVA, AND P. TUREKHANOVA Elaeagnus species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Elaeagnus angustifolia (Narrow-leafed oleaster, Russian olive)

Valleys of desert rivers of Kazakhstan, valleys of the rivers Ili, Chu, the Syr-Darya

Honey, food, medicinal, ornamental, ameliorative

E. oxycarpa (Sharp-fruited oleaster)

Torgai, Prekaspian, western and eastern low mounds, northern Ustyurt, Pre-Altai, Turkestan, Dzhungarskei, Zailiyskei, Kungei Alatau, Karatau, western Tien-Shan

Honey, food, medicinal, ornamental, ameliorative

E. turcomanica (Turkmen oleaster)

Alluvial plain of the Syr-Darya

Ornamental, honey, food

3 to 10 m tall, with an open silvery crown; trunk and older branches have reddish-brown, glossy bark; shoots and buds are covered by silvery scales. Leaves are alternate, lanceolate, oblong-lanceolate, acuminateelliptical 2.5 to 7 cm long, 0.4 to 1.5 cm wide, grayish-green on the upper sides and silvery-white on the lower sides. Fragrant flowers are axillary on short (2 mm) pedicels, with a silvery-yellow color. Edible fruit is spherical to oblong, drupaceous, 7 to 14 mm long, 5 to 10 mm wide, yellow or with reddish-bluish, sweet-mealy flesh. It blooms in May and matures fruits in July to August. Fruits have at least 50 percent carbohydrates, much glucose and fructose, about 10.6 percent protein, and they are rich in potassium and phosphorus salts. Fruits are widely used fresh and for preparation of different seasonings. Oleaster is droughtresistant, light-demanding, air pollutant-resistant, and tolerates soil salinity. When the stem is covered by sand it forms adventitious roots; therefore, it may be widely used for stabilization of riverbanks and shifting sands. It is valuable in landscaping especially as sheared hedges to provide contrasting silver colors against backgrounds of trees with darkgreen foliage, especially conifers. 2. Sharp-fruited Oleaster (Elaeagnus oxycarpa Schlecht.). This species is one of the components of alluvial forests, and grows along banks and valleys of desert rivers and lakes, on stony areas and in lowlands of sandy hills. The tree is 3 to 16 m tall with reddish-brown, glossy bark; branches have strong spines 1 to 3 cm long and young shoots and leaves

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are thickly covered by silvery-white stellate scales. Leaves are alternate, wide lanceolate or lanceolate but those near flowers and fruits are linear-oblong, acuminate. Fragrant flowers in long clusters (up to three) are found in the leaf axils. Drupaceous fruits are ovate or round, 8 to 10 mm long and 6 to 7 mm wide, yellow or orange with sweet, mealy flesh. Stone is oblong, acuminate at ends, with eight dark stripes. Sharp-fruited oleaster is a xerophytic, light-demanding and salt-tolerant plant. Under natural conditions it is easily propagated both by seeds and root sprouts. Fruits are edible and used fresh and dried. Fruit flesh contains 40 percent carbohydrates, half of which is fructose, and kernels contain 3.7 percent oil. 3. Turkmen Oleaster (Elaeagnus turcomanica N. Kozl.). This species grows in alluvial plains of the Syr-Darya. It is a deciduous tree 3 to 8 m tall with reddish-brown bark. Young shoots and leaves are covered with thick white scales and branches are spiny. Silvery leaves are widelanceolate, ovate or rhomboid, 3.0 to 3.3 cm long, 1 cm wide. Yellow flowers are 1 to 3 in leaf axils; perianth is funnel-shaped or bell-shaped, 7 to 8 mm long, 3 to 4 mm wide. Edible drupaceous fruits are numerous, globular, 10 mm in diameter, dark-red, shiny, containing sugars, tannins, and vitamin C. E. Rose Rose (Rosa L., Rosaceae) is light-demanding, drought-resistant, and winter-hardy. It grows well in full sun where it is protected from strong winds in forests and among shrubs. It is not demanding as to soil though it prefers medium-loamy, neutral, or weakly leached soils. Rose fruits (or hips) are a natural storehouse of vitamins. They accumulate large amounts of sugars, citric acid, pectin and pigment substances, flavonoids, pro-vitamins A1, B2, P, and K. Vitamin concentration in rose fruits is 6 to 8 times higher than in black currant, and 100 times higher than in apples; High concentration of vitamin C can also be found in leaves. Fresh and dried fruits are used for different kinds of processing in both industrial and home situations. Due to high content of ascorbic acid, oils, and other biologically active substances, rose is used in medicine as drinks, decoctions, extracts, and syrups for anemic diseases, fatigue syndrome, chronic diseases of liver and gall ducts. Rose is an excellent honey plant, beautiful ornamental, with its bright-green leaves, large flowers and attractive bright-colored fruits. It is used as a hedge, plantings of water basins, for decorative lawns and slopes, and in group plantings. Rose seedlings are the best rootstocks for cultivated roses.

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There are 400 species of Rosa, of which 21 are indigenous to Kazakhstan (Table 3.13) and four are endemic. All roses are deciduous upright, or sometimes creeping, shrubs, with thorns on shoots. Leaves are alternate, compound, pinnate, pubescent or glabrous, with stipules adnate to petioles. Large bisexual flowers are borne singly or in umbellatepaniculate inflorescences with petals that are white, pink, red, or yellow. Fruit is a hip or false berry, developing from the fleshy receptacle that surrounds the carpels. It is propagated by seeds or vegetatively. 1. Acicular Rose (Rosa acicularis Lindl.). This species grows in mountain gorges, wet shrub lands, stony slopes and forest borders. It is a xeromesophytic shrub 2 m high with arching branches, and thickly covered with thorns. Compound, bluish leaves with 5 to 7 ovate or elliptical leaflets are weakly pubescent on the upper sides and densely-pubescent on the lower sides. Reddish-pink, slightly aromatic flowers, 3 to 6 cm in diameter, are usually borne singly but sometimes in groups of two to three. Red fruits with thick flesh are edible, shapes are ovate, elliptical or obovate-pyriform, mostly drooping, 2 to 3 cm long and 1 to 1.5 cm wide. Vitamin C content is 4500 mg/100 g, while seeds contain 8 to 10 percent oil, and leaves and roots are abundant in tannin substances. 2. Albert Rose (Rosa alberti Regel). This species grows in forest borders and among shrubs in forest belt of the mountains. It is a xeromesophytic, strongly branched shrub 1.5 m high with long, arching Table 3.13.

Rosa species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Rosa acicularis (Acicular rose)

The Altai, Tarbagatai, Dzhungarskei Alatau, Central Kazakhstan

Food, medicinal, ornamental; easily cultured

R. albertii (Albert rose)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kirghiz, Kungei, Talasskei Alatau

Food, medicinal, ornamental

R. beggerana (Begger rose)

Tarbagatai, Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz, Talasskei Alatau, Ketmentau, Karatau

Medicinal, ornamental, food plant; easily cultured

R. canina (Dog rose)

Zailiyskei Alatau

Ornamental, medicinal, food, technical plant; easily cultured

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R. corymbifera (Corymbose rose)

Talasskei Alatau, Karatau

Ornamental plant; easily cultured

R. dsharkenti (Dzharkent rose)

Balkhash-Alakol

Endemic species

R. fedtschenkoana (Fedchenkovskiy rose)

Zailiyskei, Kirghiz, Talasskei Alatau, Karatau

Ornamental, medicinal, food plant; easily cultured

R. glabrifolia (Glabrous-leafed rose)

Tobol-Ishim, Turgai, West low mound

Ornamental tree

R. hissarica (Hissar rose)

Talasskei Alatau, Karatau

Ornamental tree

R. kokanica (Kokand rose)

Talasskei Alatau, Karatau

Ornamental plant; easily cultured

R. laxa (Lax rose)

The Altai, Tarbagatai, Dzhungarskei, Zailiyskei, Kundei, Terskei, Talasskei Alatau, Ketmentau

Food, medicinal, ornamental plant; easily cultured

R. maracandica (Samarkand rose)

Talasskei Alatau

Ornamental, honey plant

R. majalis (R. cinnamomea sensu L.) (May rose)

The Altai, Tarbagatai

Medicinal, ornamental plant

R. nanothamnus (Dwarf rose)

Dzhungarskei, Talasskei Alatau, Karatau

Ornamental, honey plant

R. oxyacantha (Acute-prickly rose)

The Altai

Food, medicinal, honey plant

R. pavlovii Pavlov rose)

North-East Kazakhstan: Irtish river

Rare endemic species

R. plathyacantha (Wide-prickly rose)

Dzhungarskei, Zailiyskei, Kungei, Terskei, Kirghiz, Talasskei Alatau, Ketmentau, Karatau

Ornamental, honey plant

R. potentilliflora (Pedate-color rose)

Zailiyskei Alatau

Endemic species

R. schrenkiana (Schrenkovskiy rose)

Dzhungarskei Alatau

Food, medicinal, honey plant

R. silverhjelmii (Silverheim [Ilijskei] rose)

Alluvial plains of the rivers Ili, Chu

Endemic species, ornamental plant; easily cultured

R. spinosissima (Spinose rose)

The Altai, Tarbagatai, Dzhungarskei Alatau

Food, medicinal, ornamental plant; easily cultured

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branches with thin, short, straight thorns. Leaves are obovate or elliptical. White flowers are borne singly. Orange-red fruits with dehiscent sepals are smooth or with long setae, ovate or elliptical, 1.5 cm long. It is one of the species with the highest vitamin content in Kazakhstan, containing 4,000 to 20,000 mg/100 g of ascorbic acid. It is used as a rootstock for horticultural remontant roses. 3. Begger Rose (Rosa beggerana Schrenk). This species grows on mountain slopes and along banks of rivers and streams. It is a xero-mesophytic shrub, 3 m high with upright, bluish branches, which have large, sickleshaped thorns that are wide and yellowish at the base. Leaves consist of three to five bijugate leaflets that are each 3 cm long, ovate, ovate-oblong, or orbicular, glabrous or slightly pubescent on the under-sides. White flowers are 2 to 3 cm in diameter, borne in complex corymbs and panicles. Spherical, red or brownish-red fruits are 4 to 5 mm in diameter with deciduous sepals upon ripening. Fruits contain vitamins C, E, P, B2, carotene, flavonoids, and are very rich in ascorbic acid from 7,000 to 20,000 mg/100 g of fruit. Seeds are rich in oils. Roots contain flavonoids and catechin, and leaves have tannin substances as well as flavonoids. 4. Dog Rose (Rosa canina L.). This species grows on open slopes, along mountain streams, and on forest borders. It is a xero-mesophytic shrub 1.5 to 2 m high with arched branches with green or red-brown bark. Thorns are sickle-shaped and compressed at the base. Compound leaves consist of five to seven elliptical, pointed leaflets, 5 cm long. Pink-towhite flowers are usually borne singly, and are 2 to 8 cm in diameter. Red fruits are orbicular or oblong-ovate, and glabrous. Roots contain tannin substances; flowers have flavonoids and ether oil, and fruits have carbohydrates, alcohol, vitamins C, P, E, carotenoids, tannin substances, and flavonoids. 5. Corymbose Rose (Rosa corymbifera Borkh.). This species grows on open mountain slopes and on banks of mountain rivers and streams. It is a xero-mesophytic shrub with velvetlike pubescence and recurved, uniform-sized thorns. Compound leaves have five to seven ovate leaflets, pubescent on both sides or with more dense pubescence on the lower side. White-to-pale pink flowers have pinnate sepals that are pubescent on the margins. Dark-red fruits are large, ovate or orbicular, with deciduous sepals. 6. Dzharkent Rose (Rosa dsharkenti Chrshan.). This species is an endemic species that grows on dry, stony slopes, among shrub lands. It

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is a strongly branching shrub about 1.1 to 1.2 m high with acicular-setose thorns that densely cover the branches. Compound leaves have 3 to 4 bijugate, oblong-ovate leaflets, about 10 to 12 cm long, that are glabrous on the upper sides and with short pubescence along the main vein on the lower side. Yellow flowers are borne on corymbose inflorescences. Dark brown fruits with a thin fleshy layer become lighter colored during ripening. 7. Fedchenkovskiy Rose (Rosa fedtschenkoana Regel). This species grows on mountain-steppe and often on stony slopes of the mountains. It is a xero-mesophytic, vigorous shrub, 4 m high, with large, straight thorns that are broad and flattened at the base. Compound leaves have seven leaflets, about 4 cm long, almost orbicular or ovate, round-bluntpointed, glabrous and bluish. Large 9 cm in diameter white flowers are usually borne singly. At the stem end, fruit narrows to a neck and the apex is wide or ovate. The orange-red fruits are 5 cm long, covered by glandular setae, usually glabrous, and contain extremely high levels of ascorbic acid. 8. Glabrous-leafed Rose (Rosa glabrifolia C.A. Mey. ex Rupr.). This species grows on steppe and alluvial meadows, and along forest borders. It is a xero-mesophytic, erect shrub, 1.2 to 2.0 m high, with dark-red or greenish bark, straight and slightly bent thorns and setae arranged in pairs. Compound leaves have five to seven leaflets, 7 cm long and elliptical or ovate-oblong. Reddish-pink flowers are usually borne singly but, at times, as two to four on short pedicels. Red fruits are large, 1.3 to 2.5 cm long, elliptical, pyriform or ovate, seldom round, with deciduous, convergent sepals. 9. Hissar Rose (Rosa hissarica Slob.). This species grows on stony slopes, in mountain gorges, among shrub-lands. It is a xerophytic, short, branching shrub about 10 to 25 (seldom 60) cm high, with numerous, thin, slightly uncinate-curved thorns that are extended-cushion-shaped at their base. Compound leaves have five to seven almost orbicular or obovate leaflets. Pinkish-white flowers are borne singly and are small, about 2.5 to 3.0 cm in diameter. Spherical, red fruits are fleshy, glabrous or glandular-bristly, mainly on the top, and constricted at the apex. 10. Kokand Rose [Rosa kokanica (Regel) Regel et Juz.]. This species grows on stony slopes, along the banks of mountain rivers, and in shrublands. It is a xerophytic shrub with erect branches 1.5 to 2 m high, with red-brown bark and straight thorns that are small and slightly flattened

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at the base. Compound leaves with seven to nine elliptical or obovate leaflets are pubescent on both sides and thickly covered by short, stalked glandules. Yellow flowers are 1 to 2 cm long, with 4 cm pedicels. Spherical, dark-brown, almost-black fruits have divaricate sepals. 11. Lax Rose (Rosa laxa Retz.). This species grows on mountain slopes, forest borders, and banks of rivers and lakes. It is a xero-mesophytic shrub, 2 m high, with thorns that are strong, uncinate-recurved or slightly pointed, and widened at the base. Grayish-green, compound leaves have five to nine leaflets that are ovate, elliptical or oblong, glabrous or slightly pubescent on the under-sides. Pale, pinkish-white flowers are borne three to six in corymbs. Fruits are smooth, globular or elliptical, with persistent sepals. Fruits contain high levels of carotene and vitamin C. 12. Samarkand Rose (Rosa maracandica Bunge). This species grows on stony slopes of the middle mountain belt. It is a xero-mesophytic, vigorously branching, low-growing shrub with scaly bark on old branches and brown-red young branches. Thorns are approximate and firm, strongly trigonal-extended at the base. Compound leaves with seven to nine leaflets are 7 to 9 cm long, and oblong-ovate or orbicular. Goldenyellow flowers are borne singly. They are 1.7 to 2.5 cm in diameter and have lanceloate sepals that are mostly pubescent on both sides. Blackviolet, glossy, glabrous fruits are globular, 5 to 7 mm in diameter. 13. May Rose (Rosa majalis Herrm.). This species grows in meadows and river valleys. It is a xero-mesophytic shrub from 20 cm to 2 m high, with bright brown-red bark on the thin branches. Thorns are curved and usually arranged in pairs. Compound leaves are thin with three to seven elliptical or ovate leaflets. Pink flowers are usually borne singly and are 3 to 6 cm in diameter. Orange-red fruits are smooth, fleshy, orbicular, seldom ovate or elliptical, orange-red, with deciduous sepals. Edible fruits have thick flesh containing large amounts of sugars, vitamin C and carotene so that they are widely used for the production of vitamin preparations. 14. Dwarf Rose (Rosa nanothamnus Bouleng.). This species grows on stony slopes of the mountains. It is a xero-mesophytic, divaricatebranching shrub, 1.5 to 2.5 m high with thin, straight thorns widened at the base. Compound leaves have five to nine orbicular or obovate leaflets, 1.5 cm long, with both sides mostly pubescent. Pinkish-white

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flowers are borne in inflorescences of two to three. Red fruits are globular or ovate, with thin glandular-bristles. 15. Acute-prickly Rose (Rosa oxyacantha Bieb.). This species grows on mountain slopes near stone mounds. It is a low, xero-mesophytic divaricate-branching shrub with numerous, thin, needle-shaped thorns. Compound leaves with nine leaflets that are oblong or elliptical, glabrous and pale-green on the lower sides with stalked glands along the middle vein. Pale pink flowers, 3 cm in diameter, are borne singly. Bright red fruits are fleshy, orbicular, or oblong. 16. Pavlov Rose (Rosa pavlovii Chrshan.). This species is a rare endemic species that grows on wet meadows among herbaceous species. It is a mesophytic, erect shrub 45 to 60 cm high with short, 2 to 3 mm, needleshaped thorns, 6 to 10 cm long and 5 to 6 mm wide, with obtuse auricles. Compound leaves have five to seven leaflets that are obovate or widely elliptical with sparsely pubescent upper sides and densely pubescent undersides. Flowers single, seldom two, pale-pink, 3.5 to 4 cm in diameter. Large globular or conical fruits, 1.5 to 1.7 cm in diameter, are borne on 2.5 cm long, thick, glandular-bristly pedicels. Because of large fruit size and ornamental value, this species has potential for introduction into culture. 17. Wide-prickly Rose (Rosa platyacantha Schrenk). This species grows in foothills, mountain slopes, in gorges. It is a xero-mesophytic shrub, 1 to 2 m high with reddish bark. Prominent thorns are straight, pointed to the apex, abruptly widened at the base and very stiff. Compound leaves have five to nine orbicular leaflets with four to eleven large, pointed or obtuse teeth and sparse pubescence on the lower side. Yellow flowers are borne singly on 1.5 to 4 cm long, glabrous pedicels; sepals are lanceolate and tomentose on the upper sides. Black-violet globular fruits are 1 to 2 cm in diameter and crowned by nonconvergent sepals. Vitamin C and tannin levels are very high in the fruits. 18. Pedate-color Rose (Rosa potentilliflora Chrshan. et M. Pop.). This is an endemic species that grows on stony mountain slopes and in gorges. It is an erect, deciduous, xero-mesophytic shrub, 60 to 80 cm high with short, straight, needle-shaped thorns. Compound leaves have seven to nine narrow-elliptical, glabrous leaflets 9 mm long. Bright yellow flowers are borne singly, 3 cm in diameter. Glossy, glabrous fruits are 8 to 9 mm in diameter.

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19. Schrenkovskiy Rose (Rosa schrenkiana Crep.). This species grows on mountain steppes and stony slopes, among shrub-lands. It is a xeromesophytic shrub, 1 to 1.5 m high with slightly sinuate branches and paired thorns, straight or slightly curved at the apex, thin, and slightly flattened. Compound leaves have three to five small leaflets with darkgreen upper sides and whitish-bluish and glabrous under sides. Flowers are usually borne singly, but sometimes in inflorescences of two to four. Red, ovate fruits are 1.5–2 cm long and contain high levels of carotene and vitamin C. 20. Silverheim Rose (Illiyskei) (Rosa silverhjelmii Schrenk). This is an endemic species that grows in sands on the banks of the alluvial plain of the Chu and Ili river valleys. It is a xerophytic shrub, 1.5 m high, with semi-twisting branches, covered by paired, sickle-shaped thorns widened at the base. Leaves have 2 to 3 narrow-elliptical, glabrous, paired leaflets, 25 mm long, 10 mm wide. White flowers are borne singly or in corymbs. Globular fruits are smooth with very thin walls, 5 to 7 mm in diameter, and black when ripe. 21. Spinose Rose (Rosa spinosissima L.). This species grows on slopes of the mountains and in steppe meadows. It is a xero-mesophytic shrub 0.7 to 2.0 m high with branches that are thickly covered by straight thin thorns. Compound leaves have five to eleven orbicular or elliptical leaflets. White to yellowish-white flowers are borne singly. Brownishred or blackish fruits are globular or oblate. Flesh is inedible due to the high tannin content. F. Sea Buckthorn Sea buckthorn (Hippophae L., Elaeagnaceae) consists of three species in the world, one of which is known in Kazakhstan. Common sea buckthorn (Hippophae rhamnoides L.) grows on alluvial deposits and along riverbanks, on stony soils of eastern low hillocks in the Zaisan and Balhash-Alakol regions, and in Altai, Kungei, Tarbagatai, Dzhungarskei, Zailiyskei, Terskei, Kirghiz Alatau, Ketmentau, Karatau, and Western Tien-Shan. Sea buckthorn is a thorny shrub, sometimes a spreading small tree, 1.5 to 6 m high with brownish-green, yellow-brown, or black-brown bark and arching branches usually form a distinctive silvery-gray canopy. Leaves are bicolored with green or gray-green upper sides and whitishsilvery lower sides, in autumn, they turn golden. Flowers are dioecious. Staminate flowers are 5 to 8 mm long and 4 to 6 mm wide, silvery-brown,

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with a bipartite perianth covered by brown, sometimes white, scales. Pistillate flowers with tubular perianths are borne axillary on very short pedicels in groups of two to five. The fragrant, orange-yellow, drupaceous fruit is juicy and of various shapes but most often oval, 5 to 11 mm long, 3 to 10 mm wide. Seeds are ovate, grey-brown or dark, glossy. Fruits possess high food and flavor values. They contain 2.4 to 6.8 percent sugars, 1.0 to 2.4 percent organic acids, 0.058 percent tannin substances, 300 mg/100 g of vitamin C, 16 mg/100 g of carotene, and 6 to 18 mg/100 g of vitamin E. There is about 6 percent oil in fruit flesh and skin and about 12 percent in seeds. This oil possesses special medicinal properties. Sea buckthorn fruits are used for production of excellent jams, jellies, and stewed fruits. Sea buckthorn is light-demanding, frostresistant, tolerant of excessive salinity and wet soils. Under natural conditions, it is propagated by seeds and root sprouts arising from dormant buds of surface horizontal roots. At six to eight years, sprouts appear around the periphery of the plant. This vigorous, young growth suppresses the older, central growth, causing it to die. The cycle of growth and dying out on a single plant may last for 60 to 80 years. Sea buckthorn is exceptionally ornamental, and may be used in orchards or parks, singly or in group plantings. Common sea buckthorn is introduced in Altai Botanical Garden (ABG), Main Botanical Garden (MBG), Karaganda Botanical Garden (KBG). It is exceptionally easy to grow in culture and is used mainly for food, medicinal, and ornamental purposes. G. Viburnum Viburnum (Viburnum L., Caprifoliaceae) includes 200 species in temperate and subtropical belts, mainly in Eurasia and North America. One species is found in Kazakhstan. Common viburnum (Virburnum opulus L.) grows on fertile moist soils, often at forest borders, glades, shrublands, on cut-over areas and along banks of rivers, lakes, and swamps in the regions of Tobolo-Ishim, Irtish, Semipalatinsk pine forests, Kokshetau, Aktubinsk, Ulytau, Karkara, Zaisan, Altai, Tarbagatai, Dzhungarskei, Zailiyskei, and Kungei Alatau. It is a strongly branched shrub about 4 m high with brownish-gray bark and green shoots. Leaves are opposite with well-developed petioles; they are wide-ovate or rounded, three- to five-lobate, large-toothed, dark-green on upper sides and pubescent on lower sides. Flowers are white, fragrant, in terminal, flat umbellate inflorescences. Drupaceous fruits are juicy, red, oval or globular, bitterish, and each fruit has a single stone seed. It blooms from the end of May to July with fruits ripening in August to September. Fruits are rich in sugars, organic acids, vitamins, pectin, and pigment substances.

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Viburnum is easily grown, propagated by seeds, and used for food, medicinal, and ornamental purposes. VI. NUTS Nut crop species of Kazakhstan include five genera found in five families. These include almond, pistachio, hazelnut, walnut, and pinenut. Within these genera there are nine species endemic to Kazakhstan. A. Almond Almond (Amygdalus L., Rosaceae) consists of about 40 species distributed in subtropical and temperate zones of the Northern Hemisphere. This genus is often included in Prunus. This genus is widely distributed from the Mediterranean to Central Asia with a concentration of species in the eastern Mediterranean region. There are five native species in Kazakhstan (Table 3.14). 1. Ledebour Almond (Amygdalus ledebouriana Schle.; syn. Prunus ledebouriana). This is a xerophytic species that grows on slopes of steppe mountains and plateaus, in river valleys and meadow hollows of the southern Altai and Tarbagatai regions. Almond trees in Tarbagatai Table 3.14.

Amygdalus (syn. Prunus) species indigenous to Kazakhstan.

Species (common name)

Distribution

Use and comments

Amygdalus ledebouriana (Ledebour almond)

Southern Altai, Tarbagatai

Ornamental, medicinal, honey plant; easily cultured

A. petunnikovii (Petunnikov almond)

Karatau, Western Tien-Shan

Ornamental, food, honey, endemic plant of Western Tien-Shan

A. spinosissima (Thorny almond)

Karatau, Western Tien-Shan

Ornamental, food, honey plant

A. nana (Dwarf almond)

Spurs of the common mountain belt, TobolIshim, Mugordzhary, Torgai, Western low mounds, Ural foothills

Ornamental, medicinal, food plant; easily cultured

A. communis (Common almond)

Western Tien-Shan

Food, medicinal, honey plant; not easily cultured

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are distributed in the central part of southern slopes, where they compose an uninterrupted belt, 100 km long and 15 km wide. They can be found at 500 to 1500 m above sea level, among desert-steppe and shrub belt. It is a deciduous shrub 2.5 m high with divaricate branches and numerous spurs. The bark of older branches is gray or reddish-gray, and reddish-brown on younger shoots. Glabrous, lanceolate, or oblongovate leaves are large and alternate on tightly clustered spurs. Pink flowers bloom simultaneously with leaf emergence. Fruits, 1.5 to 2.5 cm long and 1.2 to 2.0 cm wide, are irregular, orbicular-ovate, somewhat compressed, with dense, stiff hairs on the pericarp. Stones are wide orbicular-ovate or oblong-ovate, shallow-netted and grooved. The kernel is inedible, bitter, containing oils, amygdalin, and emulsin. It is propagated either vegetatively or by seeds. Shrubs bear fruits annually, maturing in early August. 2. Petunnikov Almond (Amygdalus petunnikovii Litv.; syn. Prunus petunnikovii Litv.). This species grows on shallow stony slopes of the mountains in Karatau and Western Tien-Shan at 1400 to 2000 m. Most often it grows in dense colonies of several shrubs; separate shrubs are seldom found. It is a strongly branching bush, 1 m high, without thorns, and with brownish-gray bark. Glabrous leaves, 2 to 3 cm long and 3 to 5 mm wide, are alternate, linear or linear-lanceolate with acuminate apex and acute-cuneate base, and serrate margins. Pink flowers open simultaneously with leaves at the end of April. Average nut weight is 0.7 g; size is 1.2 to 2.5 cm long, 1.2 to 1.6 cm wide and they are oblique with oblique-tapered base and acuminate apex. Pericarp is densely pubescent with glossy, foxy-red tomentum. Stones are brown, irregularly rounded-ovate, oblique on dorsal suture, and shallow netted. Kernel is very bitter, containing 58 percent edible oil and 3.4 percent amygdalin. Propagation is mostly by seeds. It is a beautiful ornamental shrub that is quite winter-hardy and drought-resistant. 3. Thorny Almond [Amygdalus spinosissima Bunge; syn. Prunus spinosissima (Bge.) Franch.]. This species inhabits the low belt of Karatau and Western Tien-Shan at 400 to 1500 m above sea level. Plants grow as separate shrubs on open, warm slopes, and on bare rocks. It is most prolific on the slopes of Talasskei Alatau in the Mashat river valley. It is a rough shrub, 2 m high with strongly divaricate branches and long, straight thorns. Bark on older branches is scabrous, whitish-gray, and younger shoots are glossy, red-brown. Small, leathery leaves are lanceolate, entire, and borne on short spurs and alternately on shoots. Sessile pale-pink or bright-pink flowers bloom before leaves emerge in the

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second half of April. The drupe-type fruit ripen at the beginning of July. At first the pericarp is green and at ripening becomes shiny yellow with a pink bluish cast due to a mild pubescence. Glabrous, brown stone is ovate to lanceolate, 1.5 to 2.0 cm long and 1.0 to 1.5 cm wide with an average weight of 0.34 g. The mean kernel content is 42.2 percent of stone weight. It has a slightly bitter flavor and contains amygdalin and 16 percent oil. Thorny almond is the most xerophytic almond species and is distinguished by rapid development in spring until the middle of July when growth ceases and it enters a dormant period during which it endures heat, drought, and frost. Propagated by seeds, it is an earlyflowering ornamental shrub excellent for hedges. It serves as droughtresistant rootstock for peach and is recommended for initial material for breeding drought-resistant, winter-hardy almond cultivars. 4. Dwarf Almond (Amygdalus nana L.; syn. Prunus nana Stokes). This species is a very polymorphic species that grows in the steppe grasslands, on slopes of mounds and hills, on riverbanks and precipices, on spurs of Common Syrt, Tobol-Ishim, Kokshetau, Pre Kaspian Aktyubinsk floristic regions, Mugodzhari, Torgai, Western low mounds, and Altai foothills in the north. It is the most widespread wild almond species. It is a low shrub 0.5 to 1.5 m high with glabrous, divaricate straight branches and numerous spurs. Bark of older branches is gray or reddishgray, and young shoots are reddish-brown or whitish. Glabrous leaves with stipules are linear-lanceolate or lanceolate, 4 to 7 cm long, acuminate with serrate margins. On spurs, leaves are in congested clusters. Bright-pink single flowers with slight aroma bloom simultaneously with leaf appearance. Straw-yellow fruits are dry drupes, thickly tomentousvillous, orbicular, 1 to 2 cm long, 1.2 to 1.8 cm wide. Stone is irregularly netted, furrowed, wide-orbicular or oblong-ovate, compressed with thick ventral suture flat or slightly tapered at the base and blunt or acute apex. Kernels are bitter, inedible, contain 4.5 percent of amygdalin and 50 percent fragrant oil, and are used for food and medicinal purposes. This species is widely used as an ornamental and is quite winter-hardy. It is propagated by seed. 5. Common Almond [Amygdalus communis L.; syn. Prunus communis (L.) Arcangeli]. This species grows on southern, stony, gravelly slopes of Western Tien-Shan and is cultivated in Southern Kazakhstan. Among wild almonds, common almond is the most heat-loving species and has an extremely limited distribution. Besides western Tien-Shan, natural populations of common almond are found in Western Kopetdag. It is a small tree or shrub, 4 to 8 m high, with spreading or divaricate crown,

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seldom pyramidal, with a dense root system and a swordlike, curved stem and branches without thorns. Bark on old trunks is gray-black and on older branches is grayish-brown or brown, whereas young shoots are reddish-brown or brown. Glabrous leaves with serrate margins, glands, and stipules are narrow-elliptical or lanceolate, 4 to 6 cm long, 1.5 to 2.0 cm wide, gray-green on upper sides and lighter green on the bottom. Leaves of younger shoots are alternate and on spurs they form congested bundles. Opening earlier than leaves, the attractive pink flowers, with 3 to 5 cm pedicels, are sessile or on spurs. Fruits are 1 to 6 cm long, pericarp is pubescent, dry, green, slightly red, and dehisces along a ventral suture at maturity. Stone is free with variable densities, from very firm, with stony cells, to papery and easily broken by fingers. Stone surface is grooved-furrowed or smooth. Kernel is sweet (var. dulcis DC.) or bitter (var. amara DC.). Sweet almond kernels contain 27.7 percent water, 16.5 percent nitrogen substances, 50 to 53 percent oil (almost pure oleic acid), 3 percent free fatty acids, 10 to 11 percent nitrogen-free extractive substances, 2 to 3 percent cellulose and 1.7 percent ash. Bitter almonds are characterized by higher (4%) content of amygdalin in kernels; in other respects, its contents are similar to those in sweet almond kernels. Common almond is light-demanding, heat-enduring, drought-resistant and rather winter-hardy, withstanding temperatures of 20 to 25°C. It is propagated by seeds. B. Hazelnut Hazelnut or filbert (Corylus L., Betulaceae S.F. Gray) is one of the most popular nut crops of moderate latitudes in the Northern Hemisphere. Its nuts are used directly and it is in popular demand by the confectionery industry and for cookery. Corylus consists of 22 species, distributed in temperate forest zones of Eurasia and North America. Plants are monoecious, wind-pollinated (seldom self-pollinated). Fruits are one seeded nuts, covered by a leafy involucre. Of all the species within Corylus, common hazelnut (C. avellana L.) has the most widespread, economic importance. However, it is a very rare species in Kazakhstan, with declining numbers; it grows on the west bank of the Ural river, 50 km east of Uralsk city. It is a shrub or small tree about 5 to 7 m high with a wide, almost globular crown; young glabrous branches are gray and older branches are yellow-gray, bristly and pubescent. Large, wide-oval or rounded leaves are dark-green on the upper side and dull-scabrous, lighter green on the lower side, with pubescent veins. Leaves are 6 to 12 cm long, 5 to 8 cm wide on short petioles, and have an oblique-depressed base. It blooms in April to May before leaf emergence. Staminate flowers

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form long, drooping catkins and pistillate flowers are wrapped in bracts. Fruits are usually borne in clusters of two to five but sometimes singly. Fruit involucre is bell-shaped, dissected-lobate, and equal to the nut length. Nuts are oval and brown at maturity. Hazelnuts are very tasty and nutritious. Kernels have high content of fats, protein, carbohydrates, vitamins, and other useful substances. The amount of excellent-flavored oil reaches 70 percent, proteins, 18 percent, and sugars, 2 to 5 percent. Often reaching the age of 80 years, the tree is propagated by seeds or vegetatively by sprouts at the base of the trunk. C. Pistachio Pistachio (Pistacia L., Anacardiaceae Lindl.) is one of the most remarkable nut trees in the world. It is widely known for its fruit content, formation of leaf galls, and for its valuable wood. Pistachio oil possesses useful specific traits, competing freely with olive oil on the world market. Pistachio kernels are eaten out of hand and are used in the production of cakes, biscuits, and sherbet. It also has medicinal uses. There are about 20 species of Pistacia distributed in the Northern Hemisphere including: the Mediterranean region, Central Asia, China, and southern North America. The genus was originally a component of subtropical forests of ancient Mediterranean; subsequently, it underwent strong xerophytization. Today pistachio forests in Central Asia are residues of these subtropical deciduous forests. Common pistachio (Pistacia vera L.) is the only species of the genus that is cultivated. Pistachio forests of Southern Kazakhstan represent the most northern distribution of the center of Pistacia cultivation. It grows in Boroldajtau Karatau ridge, Mashattau, Daubaba (Talasskei Alatau), and Makbalskaya forest dacha (Kirghiz Alatau) at 700 to 1100 m above sea level, on the slopes of south-southwestern exposure of 60° slopes. Usually, pistachio trees are widely scattered, but on deep soils more dense stands of 90 to 150 tree/ha are found. Modern stands are heavily damaged by fires, fellings, and grazing. Plants are small, xerophytic, many-stemmed trees, 3 to 4 m tall (sometimes up to 7 m). Bark is brownish-gray with longitudinal cracks. Young shoots are reddish-brown, thinly pubescent or glabrous. Leaves with short petioles are odd-pinnate, with 5 wide, oval or ovate, leathery, shiny leaflets, dark-green on upper sides and pale-green on lower sides, slightly pubescent. Bud-break occurs in May and leaves remain green the whole summer. Because of that, pistachio is distinguished by its dark-green crowns against a back-

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ground of faded grass cover. Buds are small, acuminate, dark brown and, by July, flower buds can be distinguished from shoot buds. Common pistachio is a dioecious plant. Staminate flowers are in dense, wide panicles 4 to 6 cm long. Pistillate flowers are in more open, narrower panicles 4 to 6 cm long. It blooms in May simultaneously with leaf emergence, and staminate flowers bloom two to three days earlier than pistillate flowers. There is about a 25-day range in flowering time within a population, and blooming period for each tree is about three to four days. There are usually approximately half male plants and half female plants in wild stands, but under more severe environmental conditions, there are more male plants (about 70%). Pistachio trees begin to bear fruits at 7 to 8 years and produce abundantly at 15 years. When trees are 30 to 40 years of age they may yield 15 to 20 kg/tree. Active fruit bearing extends to 100 years. Seedling plants attain life spans of 300 to 500 years while plants of sprout origin have considerably shorter lives. Fruit bearing of common pistachio in populations of Southern Kazakhstan is not annual, the cause of which may be explained by severe environmental conditions. Often it will have alternate-year bearing when crops are heavy. The species is very polymorphous; each pistachio plant is a specific form. Fruit is a dry drupe with a shape nearly orbicular to strongly elongated and curved. The average weight is 0.6 g. The cream to dark-violet endocarp (stone) is large; kernels are oily, bright-green, of pleasant flavor and aroma. Pericarp is easily separated at ripening. Dehiscence of the nut shell varies, with some nuts slightly split only on one side, while others dehisce the entire length on both sides. Among wild pistachio of Kazakhstan some forms are found that are not inferior in size and weight to nuts of the best cultivars. Pistachio fruits are very nutritious; they contain 51 percent oil, 3.2 percent sugar, 3.6 percent cellulose, 3.6 percent ash, and up to 24 percent protein. Pistachio fruits possess specific tonic characteristics; therefore, local people call pistachio “tree of life.” Common pistachio is light-demanding and more drought-resistant than most other trees, a real inhabitant of the semiarid zone. Its drought-resistance is explained by deep root system, to a depth of 7 meters. Pistachio also possesses winter-hardiness, withstanding temperature of –41°C. Trees are able to grow without irrigation on very stony soils, steep and washed-out slopes and thus, represents a valuable plant for anchorage of mountain slopes. It is propagated by seeds and possesses remarkable ability to form sprouts from roots, even into old age. Common pistachio is a rare species in Kazakhstan.

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D. Siberian Pine The genus Pinus L., Pinaceae Lindl., is comprised of evergreen trees, seldom shrubs, with about 100 species, mainly growing in the temperate belt of the Northern Hemisphere. The pine genus is one of the main forest-forming trees. Seeds of Siberian pine (Pinus sibirica Du Tour), and Italian pine are edible. Siberian Pine is also know as Cedar Pine or Siberian Cedar and is referred to as “Taiga pearl,” or “Tsar of Taiga.” In Kazakhstan, it grows naturally only in the Altai, composing together with larch, fir, spruce and birch the scenic, dark, coniferous taiga. Siberian pine is a vigorous, beautiful tree, 35 m tall; crown of young trees is acute-pyramidal, of old trees wide-spreading, often polyapical; branching is whorled. Bark on young branches is ash-silvery with brown, transverse lenticels; later it is cracked, gray-brown. Young shoots are covered by thick, foxy-red pubescence. Needles are 5 to 12 cm long, triangular in cross section, dark-green with bluish bloom, usually living three to seven years, but sometimes to eleven years. Staminate (pollen) catkins form in the middle part of crown. Pistillate cones are borne in clusters of two to three on shoots in the upper part of the crown, near terminal buds. As cones mature, the color changes from violet to yellowishbrown; they are ovate, 6 to 13 cm long, 5 to 8 cm wide, and covered by densely appressed scales with a thickened, terminal protuberance. Each cone contains 80 to 140 brown seeds each, 10 to 14 mm long, 6 to 10 mm wide. Seeds become ripe two years after blooming and fall together with cones. In productive years, one large tree gives 1000 to 1500 cones. Kernels of pinenuts are a valuable food product that contains natural concentrates of proteins and carbohydrates along with cedar oil (50–56%), and has excellent flavor. Siberian pine is adapted to a continental climate; it is water-loving, and at maturity age is light-demanding. It grows on different soils though it prefers well-drained, deep-loamy, weakly-podzolic soils. It endures polluted, smoky air. Transplanted at a mature age, trees grow slowly but are long lived, up to 500 years. It is propagated by seeds. In culture, it requires regular irrigation, and is injured by dry air. It is considered both an ornamental and food source.

E. Walnut Species of walnut (Juglans L., Juglandaceae A. Rich. ex Kunth.) are distributed in temperate and subtropical areas of the Northern Hemisphere, mainly in mountain forests. Walnuts are distributed in three separate regions, Mediterranean, East-Asian Himalayas, and North American.

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Juglans regia is one of the most valuable nut crops in the world. Nuts are distinguished by exceptional food value, high calories, and medicinaldietetic properties. Kernels contain 43 percent oil, 20 percent protein, 15 percent carbohydrates, and vitamins A and C. Due to its useful properties, walnut has a long history of cultivation in almost all temperate regions of the world. Walnut forests of Western Tien-Shan are unique in the world. Here walnut populations, along with its companions, apple, pear, alycha, plum and wild grape, are of great importance both as ecological systems and as a source for genetic improvement of cultivars. Ugam center (South Kazakhstan region) is the northern-most extent of the natural area of walnut. Here, this species does not form extensive forests: trees are scattered in small groves of 0.6 to 0.8 ha. It usually grows in moist sites where ground water is available, in hollows, or river alluvial plains. Often it is found on shady slopes, though it is light-demanding for good nut production. Sometimes on dry slopes, scattered trees of walnut present a park-like appearance. Walnut is deciduous, monoecious, and windpollinated. Walnut trees, among other fruit trees, are sharply differentiated by their size and vigor, tree height and crown diameter often reach 30 m and trunk is 2 m. Trees may have single trunks or be multistemmed. Crown form of single trees is globular; in dense, crowded stands, it is elongated and irregular. Low and middle branches often spread at right angles. The bark on older wood that is light-gray and covered with deep cracks is very beautiful. Bark of young shoots in spring is very shiny, olive-gray, and in autumn, greenish-brown, glabrous, often ribbed with numerous lenticels. At tree base, thick sprouts may grow. Leaves are large, long, compound odd-pinnate with a distinctive, pleasant aroma. Leaflets are ovate, entire, glabrous, acuminate, and terminal leaflet is the largest. After bud-break (April–May), leaves are pink, pubescent; in summer, dark-green, shiny above and dull, lighter under sides; in autumn, at the end of September and October, they are yellowish. Staminate flowers open before leaves appear; they are in multiflorous thick green catkins, growing on shoots of the previous year. Simultaneous with leaves, inconspicuous pistillate flowers appear in small clusters formed on the ends of current-year shoots. Bisexual flowers are also found, but very rarely. Walnut is an exceptionally polymorphous species. Nuts are diverse in shape, color, shell thickness, kernel output, and fat content. Fruit of walnut is a false drupe. The fleshy hulls surrounding the true fruit (or nut) consist of a fused perianth amid bracts. The hull surrounding the nut is usually slightly pubescent, first green with white dots, and later, during

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ripening brownish. Average fruit weight is 9.7 g (2.5–20 g), average length is 32.2 mm (18–42 mm), width 28.3 mm (18–39 mm). Fruit shape is rounded, oblong, or strongly extended. Shell surface is from very rough to almost smooth. Shell thickness is extremely diverse, from hard and thick to, usually, thin and delicate. Some shells even have holes where kernels are visible and some are so fragile that they are easily crushed by hands. Color is from whitish to brown. Kernel weight is 3.0 to 5.8 g, with an average of 4.3 g. Ratio of kernel-to-shell weight is 33.5 to 58.2 percent. Average percent oil is 43.3 and, with some selected forms, as high as 76.8 percent. Walnut begins to bear at 8 to 10 years with the highest yields achieved at 30 to 100 years. Maximum yield was 86 kg/tree in Ugam nut stands. Under natural conditions walnut is propagated by both seeds and vegetatively through shoots. Longevity is 200 to 300 years. At present, the natural renewal of walnut in Ugam populations is unsatisfactory. In winter, annual shoots may show freeze damage; in the most severe winters, some trees froze to the base. Walnut is used as a food, medicinal, and ornamental plant. LITERATURE CITED Bakhteev, F. Kh. 1970. The most important fruit plants. Prosveshchenie, Moscow. Burbank, L. 1955. Selected works. Moscow, Leningrad. Dubinin, N. P. 1976. General genetics. Nauka Acad. Sci. Moscow. Dzhangaliev, A. D. 1941. Wild grape of Karatau (Western Tien-Shan). Moscow Agr. Academy, Timiryasev. Dissertation. Dzhangaliev, A. D. 1950. Agrobiological characteristics of wild grape in Karatau mountains (Western Tien-Shan). Trudy Instituta Zemledeliya 2. Alma-Ata. Dzhangaliev, A. D. 1968. Economic-valuable forms of wild apple for introduction in culture. In: Varieties of fruits and small fruits of Kazakhstan. Kainar, Alma-Ata. Dzhangaliev, A. D. 1969. Apple forest of Zailiyskei and Dzhungarskei Alatau, its preservation and use. Bot. Inst. Acad. Sci. USSR, Ph.D. Diss., Leningrad. Dzhangaliev, A. D. 1973. Variety of wild apple forms of Kazakhstan and their national use. Agr. Sci. Kazakhstan, Alma-Ata. Dzhangaliev, A. D. 1975. Fruit forests of Kazakhstan, their importance, preservation and use. Abstr. XII Int. Bot. Congr., Leningrad. Dzhangaliev, A. D. 1975. Fruit forests of Kazakhstan: preservation of their phytogene pool and rational usage. Bul. Main Bot. Gard. Acad. Sci. USSR 100. Dzhangaliev, A. D. 1977. Wild apple of Kazakhstan (biocoenotic role, biological peculiarities, polymorphism, preservation). “Nauka” Kaz SSR, Alma-Ata. Gansen, H. E. 1937. Selection of fruit crops in USSR and USA. Selkhozgiz, Moscow. Red Book of Alma-Ata. 1981. “Nauka” Kaz SSR, Alma-Ata. Salova, T. N. 1985. Wild fruit plants of Kazakhstan. “Vestnik” Agr. Sci. Kazakhstan 2, Alma-Ata. Salova, T. N. 1986. Introduction of fruit and small fruit plants in Botanical Garden of Kazakhstan. In: Problems of rational use of medicinal-technical plants of Kazakhstan. “Nauka,” Kaz SSR, Alma-Ata.

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Salova, T. N. 1988. Apricot stands of Kazakhstan, their importance, use, preservation. Thesis. United Plenum and Republic Committee on the UNESCO programme “Man and Biosphere,” Alma-Ata. Salova, T. N. 1990. Plants of natural flora of Kazakhstan in introduction. Galym, AlmaAta. Timofeev-Resovsky, N. V., A. V. Jablochkov, and N. V. Glotov. 1973. Essay on population study. Moscow. Turekhanova, R. M. 1987. Productivity of sea buckthorn varieties in the premountain zone of Zailiyskei Alatau. Dep. All-Union Sci. Res. Inst. Techn. Inform. Turekhanova, R. M. 1991. Introduction perspectives in culture of the Altai Sea buckthorn varieties under conditions of premountains of Zailiyskei Alatau. Izv. Acad. Sci. Kaz SSR, Ser. Biol. Vavilov, N. I. 1935. Theoretical bases of plant selection. Agr. Literature, Moscow, Leningrad. Vavilov, N. I. 1987. Five continents. “Mysl” Moscow. Vintergoller. B. A. 1976. Rare plants of Kazakhstan. “Nauka” Kaz. SSR, Alma-Ata. Zapryagaeva V. I., and I. V. Muchurin. 1964. Wild fruits of Tajikistan, “Nauka” Moscow, Leningrad. Zhukovsky, P. A. 1971. Cultivated plants and their relatives. Kolos, Leningrad.

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Subject Index Volume 29 bilberry, wild of Kazakhstan, 347–348 blackberry, wild of Kazakhstan, 345 cherry, wild of Kazakhstan, 326–330 cotonaeaster, wild of Kazakhstan, 316–317 cranberry, wild of Kazakhstan, 349 currant, wild of Kazakhstan, 341 elderberry, wild of Kazakhstan, 349–350 gooseberry, wild of Kazakhstan, 341–342 grape, wild of Kazakhstan, 342–343 lingonberry, wild of Kazakhstan, 348–349 mountain ash, wild of Kazakhstan, 322–324 mulberry, wild of Kazakhstan, 350–351 oleaster, wild of Kazakhstan, 351–353 pear, wild of Kazakhstan, 315–316 plum, wild of Kazakhstan, 330–332 raspberry, wild of Kazakhstan, 343–345 rose, wild of Kazakhstan, 353–360 sea buckthorn, wild of Kazakhstan, 361 strawberry, wild of Kazakhstan, 347 vacciniums, wild of Kazakhstan, 347–349 viburnam, wild of Kazakhstan, 361–362

A Almond, wild of Kazakhstan, 262–265 Apple: germplasm acquisition and resources, 1–61 wild of Kazakhstan, 63–303, 305–315 Apricot, wild of Kazakhstan, 325–326 B Barberry, wild of Kazakhstan, 332–336 Bilberry, wild of Kazakhstan, 347–348 Blackberry, wild of Kazakhstan, 345 C Cherry, wild of Kazakhstan, 326–330 Cotoneaster, wild of Kazakhstan, 316–317 Cranberry, wild of Kazakhstan, 349 Currant, wild of Kazakhstan, 341 D Dedication, Sperling, C.E, ix–x E Elderberry, wild of Kazakhstan, 349–350 F Fruit crops: apple germplasm, 1–61, 63–303 apple, wild of Kazakhstan, 63–303, 63–303, 305–315 apricot, wild of Kazakhstan, 325–326 barberry, wild of Kazakhstan, 332–336

G Genetic variation, wild apple, 63–303 Germplasm: acquisition, apple, 1–61 characterization, apple, 45–56 resources, apple 1–61

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374 Gooseberry, wild of Kazakhstan, 341–342 Grape, wild of Kazakhstan, 342–343 H Hawthorne, wild of Kazakhstan, 317–322 Hazelnut, wild of Kazakhstan, 365–366 Honeysuckle, wild of Kazakhstan, 350 K Kazakhstan, See Wild fruits and nuts

SUBJECT INDEX R Raspberry, wild of Kazakhstan, 343–345 Rose, wild of Kazakhstan. 353–360 S Sea buckthorn, wild of Kazakhstan, 361 Strawberry, wild of Kazakhstan, 347 V

L

Vacciniums, wild of Kazakhstan, 347–349 Viburnam, wild of Kazakhstan, 361–362

Lingonberry, wild of Kazakhstan, 348–349

W

M Mountain ash, wild of Kazakhstan, 322–324 Mulberry, wild of Kazakhstan, 350–351 N Nut crops almond, wild of Kazakhstan, 262–265 hazelnut, wild of Kazakhstan, 365–366 pine, wild of Kazakhstan, 368–369 pistachio, wild of Kazakhstan, 366–368 walnut, wild of Kazakhstan, 369–370 O Oleaster, wild of Kazakhstan, 351–353 Ornamental plant cotoneaster, wild of Kazakhstan, 316–317 honeysuckle, wild of Kazakhstan, 350 oleaster, wild of Kazakhstan, 351–353 rose, wild of Kazakhstan, 353–360 viburnam, wild of Kazakhstan, 361–362 P Pear, wild of Kazakhstan, 315–316 Pine, wild of Kazakhstan, 368–369 Pistachio, wild of Kazakhstan, 366–368 Plum, wild of Kazakhstan, 330–332

Walnut, wild of Kazakhstan, 369–370 Wild fruit and nuts of Kazakhstan, 305–371 almond, 262–265 apple, 63–303, 305–315 apricot, 325–326 barberry, 332–336 bilberry, 347–4=348 blackberry, 345 cherry, 326–330 cranberry, 349 currant, 341 cotonaeaster, 316–317 elderberry, 349–350 gooseberry, 341–342 grape, 342–343 hazelnut, 365–366 lingonberry, 348–349 mountain ash, 322–324 mulberry, 350–351 oleaster, 351–353 pear, 315–316 pine, 368–369 pistachio, 366–368 plum, 330–332 raspberry, 343–345 rose, 353–360 sea buckthorn, 361 strawberry, 347 vacciniums, 347–349 viburnam, 361–362 walnut, 369–370

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Cumulative Subject Index (Volumes 1–29)

A Abscisic acid: chilling injury, 15:78–79 cold hardiness, 11:65 dormancy, 7:275–277 genetic regulation, 16:9–14, 20–21 lychee, 28:437–443 mechanical stress, 17:20 rose senescence, 9:66 stress, 4:249–250 Abscission: anatomy and histochemistry, 1:172–203 citrus, 15:145–182, 163–166 flower and petals, 3:104–107 regulation, 7:415–416 rose, 9:63–64 Acclimatization: foliage plants, 6:119–154 herbaceous plants, 6:379–395 micropropagation, 9:278–281, 316–317 Actinidia, 6:4–12 Adzuki bean, genetics, 2:373 Agapanthus, 25:56–57 Agaricus, 6:85–118 Agrobacterium tumefaciens, 3:34 Air pollution, 8:1–42 Alkaloids, steroidal, 25:171–196 Allium phytonutrients, 28:156–159 Almond: bloom delay, 15:100–101 in vitro culture, 9:313

postharvest technology and utilization, 20:267–311 wild of Kazakhstan, 29:262–265 Alocasia, 8:46, 57, see also Aroids Alternate bearing: chemical thinning, 1:285–289 fruit crops, 4:128–173 pistachio, 3:387–388 Aluminum: deficiency and toxicity symptoms in fruits and nuts, 2:154 Ericaceae, 10:195–196 Amarcrinum, 25: 57 Amaryllidaceae, growth, development, flowering, 25:1–70 Amaryllis, 25:4–15 Amorphophallus, 8:46, 57, see also Aroids Anatomy and morphology: apple flower and fruit, 10:273–308 apple tree, 12:265–305 asparagus, 12:71 cassava, 13:106–112 citrus, abscission, 15:147–156 embryogenesis, 1:4–21, 35–40 fig, 12:420–424 fruit abscission, 1:172–203 fruit storage, 1:314 ginseng, 9:198–201 grape flower, 13:315–337 grape seedlessness, 11:160–164 heliconia, 14:5–13 kiwifruit, 6:13–50

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376 magnetic resonance imaging, 20:78–86, 225–266 orchid, 5:281–283 navel orange, 8:132–133 pecan flower, 8:217–255 petal senescence, 1:212–216 pollution injury, 8:15 waxes, 23:1–68 Androgenesis, woody species, 10:171–173 Angiosperms, embryogenesis, 1:1–78 Anthurium: see also Aroids, ornamental fertilization, 5:334–335 Antitranspirants, 7:334 cold hardiness, 11:65 Apical meristem, cryopreservation, 6:357–372 Apple: alternate bearing, 4:136–137 anatomy and morphology of flower and fruit, 10:273–309 bioregulation, 10:309–401 bitter pit, 11:289–355 bloom delay, 15:102–104 CA storage, 1:303–306 chemical thinning, 1:270–300 fertilization, 1:105 fire blight control, 1:423–474 flavor, 16:197–234 flower induction, 4:174–203 fruiting, 11:229–287 fruit cracking and splitting, 19:217–262 functional phytonutrients, 27:304 germplasm acquisition and resources, 29:1–61 in vitro, 5:241–243; 9:319–321 light, 2:240–248 maturity indices, 13:407–432 mealiness, 20:200 nitrogen metabolism, 4:204–246 replant disease, 2:3 root distribution, 2:453–456 scald, 27:227–267 stock-scion relationships, 3:315–375 summer pruning, 9:351–375 tree morphology and anatomy, 12:265–305 vegetative growth, 11:229–287 watercore, 6:189–251 weight loss, 25:197–234 wild of Kazakhstan, 29:63–303, 305–315

CUMULATIVE SUBJECT INDEX yield, 1:397–424 Apricot: bloom delay, 15:101–102 CA storage, 1:309 origin and dissemination, 22:225–266 wild of Kazakhstan, 29:325–326 Arabidopsis: molecular biology of flowering, 27:1–39, 41–77 Aroids: edible, 8:43–99; 12:166–170 ornamental, 10:1–33 Arsenic, deficiency and toxicity symptoms in fruits and nuts, 2:154 Artemisia, 19:319–371 Artemisinin, 19:346–359 Artichoke, CA storage, 1:349–350 Asexual embryogenesis, 1:1–78; 2:268–310; 3:214–314; 7:163–168, 171–173, 176–177, 184, 185–187, 187–188, 189; 10:153–181; 14:258–259, 337–339; 24:6–7; 26:105–110 Asparagus: CA storage, 1:350–351 fluid drilling of seed, 3:21 postharvest biology, 12:69–155 Auxin: abscission, citrus, 15:161, 168–176 bloom delay, 15:114–115 citrus abscission, 15:161, 168–176 dormancy, 7:273–274 flowering, 15:290–291, 315 genetic regulation, 16:5–6, 14, 21–22 geotropism, 15:246–267 mechanical stress, 17:18–19 petal senescence, 11:31 Avocado: CA and MA, 22:135–141 flowering, 8:257–289 fruit development, 10:230–238 fruit ripening, 10:238–259 rootstocks, 17:381–429 Azalea, fertilization, 5:335–337 B Babaco, in vitro culture, 7:178 Bacteria: diseases of fig, 12:447–451 ice nucleating, 7:210–212; 11:69–71

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CUMULATIVE SUBJECT INDEX pathogens of bean, 3:28–58 tree short life, 2:46–47 wilt of bean, 3:46–47 Bacteriocides, fire blight, 1:450–459 Bacteriophage, fire blight control, 1:449–450 Banana: CA and MA, 22:141–146 CA storage, 1:311–312 fertilization, 1:105 in vitro culture, 7:178–180 Banksia, 22:1–25 Barberry, wild of Kazakhstan, 29:332–336 Bean: CA storage, 1:352–353 fluid drilling of seed, 3:21 resistance to bacterial pathogens, 3:28–58 Bedding plants, fertilization, 1:99–100; 5:337–341 Beet: CA storage, 1:353 fluid drilling of seed, 3:18–19 Begonia (Rieger), fertilization, 1:104 Biennial bearing. See Alternate bearing Bilberry, wild of Kazakhstan, 29:347–348 Biochemistry, petal senescence, 11:15–43 Bioreactor technology, 24:1–30 Bioregulation: See also Growth substances apple and pear, 10:309–401 Bird damage, 6:277–278 Bitter pit in apple, 11:289–355 Blackberry: harvesting, 16:282–298 wild of Kazakhstan, 29:345 Black currant, bloom delay, 15:104 Bloom delay, deciduous fruits, 15:97 Blueberry: developmental physiology, 13:339–405 harvesting, 16:257–282 nutrition, 10:183–227 Boron: deficiency and toxicity symptoms in fruits and nuts, 2:151–152 foliar application, 6:328 nutrition, 5:327–328 pine bark media, 9:119–122 Botanic gardens, 15:1–62 Bramble, harvesting, 16:282–298

377 Branching, lateral: apple, 10:328–330 pear, 10:328–330 Brassica classification, 28:27–28 Brassicaceae, in vitro, 5:232–235 Breeding. See Genetics and breeding Broccoli, CA storage, 1:354–355 Brussels sprouts, CA storage, 1:355 Bulb crops: See also Tulip development, 25:1–70 flowering, 25:1–70 genetics and breeding, 18:119–123 growth, 25: 1–70 in vitro, 18:87–169 micropropagation, 18:89–113 root physiology, 14:57–88 virus elimination, 18:113–123 C CA storage. See Controlled-atmosphere storage Cabbage: CA storage, 1:355–359 fertilization, 1:117–118 Cactus: crops, 18:291–320 grafting, 28:106–109 reproductive biology, 18:321–346 Caladium. See Aroids, ornamental Calciole, nutrition, 10:183–227 Calcifuge, nutrition, 10:183–227 Calcium: bitter pit, 11:289–355 cell wall, 5:203–205 container growing, 9:84–85 deficiency and toxicity symptoms in fruits and nuts, 2:148–149 Ericaceae nutrition, 10:196–197 foliar application, 6:328–329 fruit softening, 10:107–152 nutrition, 5:322–323 pine bark media, 9:116–117 tipburn, disorder, 4:50–57 Calmodulin, 10:132–134, 137–138 Caparis, See Caper bush Caper bush, 27:125–188 Carbohydrate: fig, 12:436–437 kiwifruit partitioning, 12:318–324 metabolism, 7:69–108

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378 Carbohydrate (cont.) partitioning, 7:69–108 petal senescence, 11:19–20 reserves in deciduous fruit trees, 10:403–430 Carbon dioxide, enrichment, 7:345–398, 544–545 Carnation, fertilization, 1:100; 5:341–345 Carrot: CA storage, 1:362–366 fluid drilling of seed, 3:13–14 Caryophyllaceae, in vitro, 5:237–239 Cassava, 12:158–166; 13:105–129; 26:85–159 Cauliflower, CA storage, 1:359–362 Celeriac, CA storage, 1:366–367 Celery: CA storage, 1:366–367 fluid drilling of seed, 3:14 Cell culture, 3:214–314 woody legumes, 14:265–332 Cell membrane: calcium, 10:126–140 petal senescence, 11:20–26 Cellular mechanisms, salt tolerance, 16:33–69 Cell wall: calcium, 10:109–122 hydrolases, 5:169–219 ice spread, 13:245–246 tomato, 13:70–71 Chelates, 9:169–171 Cherimoya, CA and MA, 22:146–147 Cherry: bloom delay, 15:105 CA storage, 1:308 origin, 19:263–317 wild of Kazakhstan, 29:326–330 Chestnut: blight, 8:281–336 in vitro culture, 9:311–312 Chicory, CA storage, 1:379 Chilling: injury, 4:260–261; 15:63–95 injury, chlorophyll fluorescence, 23:79–84 pistachio, 3:388–389 Chlorine: deficiency and toxicity symptoms in fruits and nuts, 2:153 nutrition, 5:239

CUMULATIVE SUBJECT INDEX Chlorophyll fluorescence, 23:69–107 Chlorosis, iron deficiency induced, 9:133–186 Chrysanthemum fertilization, 1:100–101; 5:345–352 Citrus: abscission, 15:145–182 alternate bearing, 4:141–144 asexual embryogenesis, 7:163–168 CA storage, 1:312–313 chlorosis, 9:166–168 cold hardiness, 7:201–238 fertilization, 1:105 flowering, 12:349–408 functional phytochemicals, fruit, 27:269–315 honey bee pollination, 9:247–248 in vitro culture, 7:161–170 juice loss, 20:200–201 navel orange, 8:129–179 nitrogen metabolism, 8:181 practices for young trees, 24:319–372 rootstock, 1:237–269 viroid dwarfing, 24:277–317 Classification: Brassica, 28:27–28 lettuce, 28:25–27 potato, 28:23–26 tomato, 28:21–23 Clivia, 25:57 Cloche (tunnel), 7:356–357 Coconut palm: asexual embryogenesis, 7:184 in vitro culture, 7:183–185 Cold hardiness, 2:33–34 apple and pear bioregulation, 10:374–375 citrus, 7:201–238 factors affecting, 11:55–56 herbaceous plants, 6:373–417 injury, 2:26–27 nutrition, 3:144–171 pruning, 8:356–357 Colocasia, 8:45, 55–56, see also Aroids Common blight of bean, 3:45–46 Compositae, in vitro, 5:235–237 Container production, nursery crops, 9:75–101 Controlled-atmosphere (CA) storage: asparagus, 12:76–77, 127–130 chilling injury, 15:74–77

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CUMULATIVE SUBJECT INDEX flowers, 3:98; 10:52–55 fruit quality, 8:101–127 fruits, 1:301–336; 4:259–260 pathogens, 3:412–461 seeds, 2:134–135 tropical fruit, 22:123–183 tulip, 5:105 vegetable quality, 8:101–127 vegetables, 1:337–394; 4:259–260 Controlled environment agriculture, 7:534–545, see also Greenhouse and greenhouse crops; hydroponic culture; protected culture Copper: deficiency and toxicity symptoms in fruits and nuts, 2:153 foliar application, 6:329–330 nutrition, 5:326–327 pine bark media, 9:122–123 Corynebacterium flaccumfaciens, 3:33, 46 Cotoneaster, wild of Kazakhstan, 29:316–317 Cowpea: genetics, 2:317–348 U.S. production, 12:197–222 Cranberry: botany and horticulture, 21:215–249 fertilization, 1:106 harvesting, 16:298–311 wild of Kazakhstan, 29:349 Crinum, 25:58 Crucifers phytochemicals, 28:150–156 Cryopreservation: apical meristems, 6:357–372 cold hardiness, 11:65–66 Cryphonectria parasitica. See Endothia parasitica Crytosperma, 8:47, 58, see also Aroids Cucumber: CA storage, 1:367–368 grafting, 28:91–96 Cucurbita pepo, cultivar groups history, 25:71–170 Currant: harvesting, 16:311–327 wild of Kazakhstan, 29:341 Custard apple, CA and MA, 22:164 Cyrtanthus, 25:15–19 Cytokinin: cold hardiness, 11:65

379 dormancy, 7:272–273 floral promoter, 4:112–113 flowering, 15:294–295, 318 genetic regulation, 16:4–5, 14, 22–23 grape root, 5:150, 153–156 lettuce tipburn, 4:57–58 petal senescence, 11:30–31 rose senescence, 9:66 D Date palm: asexual embryogenesis, 7:185–187 in vitro culture, 7:185–187 Daylength. See Photoperiod Dedication: Bailey, L.H., 1:v–viii Beach, S.A., 1:v–viii Bukovac, M.J., 6:x–xii Campbell, C.W., 19:xiii Cummins, J.N., 15:xii–xv Dennis, F.G., 22:xi–xii De Hertogh, A.A., 26:xi–xii Faust, Miklos, 5:vi–x Hackett, W.P., 12:x–xiii Halevy, A.H., 8:x–xii Hess, C.E., 13:x–xii Kader, A.A., 16:xii–xv Kamemoto, H., 24:x–xiii Looney, N.E., 18:xiii Magness, J.R., 2:vi–viii Moore, J.N., 14:xii–xv Possingham, J.V., 27:xi–xiii Pratt, C., 20:ix–xi Proebsting, Jr., E.L., 9:x–xiv Rick, Jr., C.M., 4:vi–ix Ryugo, K., 25:x–xii Sansavini, S., 17:xii–xiv Sherman, W.B., 21:xi–xiii Smock, R.M., 7:x–xiii Sperling, C.E., 29:ix–x Stevens, M.A., 28:xi–xiii Weiser, C.J., 11:x–xiii Whitaker, T.W., 3:vi–x Wittwer, S.H., 10:x–xiii Yang, S.F., 23:xi Deficit irrigation, 21:105–131 Deficiency symptoms, in fruit and nut crops, 2:145–154 Defoliation, apple and pear bioregulation, 10:326–328

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380 ‘Delicious’ apple, 1:397–424 Desiccation tolerance, 18:171–213 Dieffenbachia. See Aroids, ornamental Dioscorea. See Yam Disease: and air pollution, 8:25 aroids, 8:67–69; 10:18; 12:168–169 bacterial, of bean, 3:28–58 cassava, 12:163–164 control by virus, 3:399–403 controlled-atmosphere storage, 3:412–461 cowpea, 12:210–213 fig, 12:447–479 flooding, 13:288–299 hydroponic crops, 7:530–534 lettuce, 2:187–197 mycorrhizal fungi, 3:182–185 ornamental aroids, 10:18 resistance, acquired, 18:247–289 root, 5:29–31 stress, 4:261–262 sweet potato, 12:173–175 tulip, 5:63, 92 turnip moasic virus, 14:199–238 waxes, 23:1–68 yam (Dioscorea), 12:181–183 Disorder: see also Postharvest physiology: bitterpit, 11:289–355 fig, 12:477–479 pear fruit, 11:357–411 watercore, 6:189–251; 11:385–387 Dormancy, 2:27–30 blueberry, 13:362–370 release in fruit trees, 7:239–300 tulip, 5:93 Drip irrigation, 4:1–48 Drought resistance, 4:250–251 cassava, 13:114–115 Durian, CA and MA, 22:147–148 Dwarfing: apple, 3:315–375 apple mutants, 12:297–298 by virus, 3:404–405 E Easter lily, fertilization, 5:352–355 Eggplant: grafting, 28:103–104

CUMULATIVE SUBJECT INDEX phytochemicals, 28:162–163 Elderberry, wild of Kazakhstan, 29:349–350 Embryogenesis. See Asexual embryogenesis Endothia parasitica, 8:291–336 Energy efficiency, in greenhouses, 1:141–171; 9:1–52 Environment: air pollution, 8:20–22 controlled for agriculture, 7:534–545 controlled for energy efficiency, 1:141–171; 9:1–52 embryogenesis, 1:22, 43–44 fruit set, 1:411–412 ginseng, 9:211–226 greenhouse management, 9:32–38 navel orange, 8:138–140 nutrient film technique, 5:13–26 Epipremnum. See Aroids, ornamental Eriobotrya japonica. See Loquat Erwinia: amylovora, 1:423–474 lathyri, 3:34 Essential elements: foliar nutrition, 6:287–355 pine bark media, 9:103–131 plant nutrition, 5:318–330 soil testing, 7:1–68 Ethylene: abscission, citrus, 15:158–161, 168–176 apple bioregulation, 10:366–369 avocado, 10:239–241 bloom delay, 15:107–111 CA storage, 1:317–319, 348 chilling injury, 15:80 citrus abscission, 15:158–161, 168–176 cut flower storage, 10:44–46 dormancy, 7:277–279 flowering, 15:295–296, 319 flower longevity, 3:66–75 genetic regulation, 16:6–7, 14–15, 19–20 kiwifruit respiration, 6:47–48 mechanical stress, 17:16–17 petal senescence, 11:16–19, 27–30 rose senescence, 9:65–66 Eucharis, 25:19–22 Eucrosia, 25:58

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CUMULATIVE SUBJECT INDEX F Feed crops, cactus, 18:298–300 Feijoa, CA and MA, 22:148 Fertilization and fertilizer: anthurium, 5:334–335 azalea, 5:335–337 bedding plants, 5:337–341 blueberry, 10:183–227 carnation, 5:341–345 chrysanthemum, 5:345–352 controlled release, 1:79–139; 5:347–348 Easter lily, 5:352–355 Ericaceae, 10:183–227 foliage plants, 5:367–380 foliar, 6:287–355 geranium, 5:355–357 greenhouse crops, 5:317–403 lettuce, 2:175 nitrogen, 2:401–404 orchid, 5:357–358 poinsettia, 5:358–360 rose, 5:361–363 snapdragon, 5:363–364 soil testing, 7:1–68 trickle irrigation, 4:28–31 tulip, 5:364–366 Vaccinium, 10:183–227 zinc nutrition, 23:109–128 Fig: industry, 12:409–490 ripening, 4:258–259 Filbert, in vitro culture, 9:313–314 Fire blight, 1:423–474 Flooding, fruit crops, 13:257–313 Floral scents, 24:31–53 Floricultural crops: see also individual crops: Amaryllidaceae, 25:1–70 Banksia, 22:1–25 fertilization, 1:98–104 growth regulation, 7:399–481 heliconia, 14:1–55 Leucospermum, 22:27–90 postharvest physiology and senescence, 1:204–236; 3:59–143; 10:35–62; 11:15–43 Protea, 26:1–48 Florigen, 4:94–98 Flower and flowering:

381 Amaryllidaceae, 25:1–70 apple anatomy and morphology, 10:277–283 apple bioregulation, 10:344–348 Arabidopsis, 27:1–39, 41–77 aroids, ornamental, 10:19–24 avocado, 8:257–289 Banksia, 22:1–25 blueberry development, 13:354–378 cactus, 18:325–335 citrus, 12:349–408 control, 4:159–160; 15:279–334 development (postpollination), 19:1–58 fig, 12:424–429 grape anatomy and morphology, 13:354–378 homeotic gene regulation, 27:41–77 honey bee pollination, 9:239–243 induction, 4:174–203, 254–256 initiation, 4:152–153 in vitro, 4:106–127 kiwifruit, 6:21–35; 12:316–318 Leucospermum, 22:27–90 lychee, 28:397–421 orchid, 5:297–300 pear bioregulation, 10:344–348 pecan, 8:217–255 perennial fruit crops, 12:223–264 phase change, 7:109–155 photoperiod, 4:66–105 pistachio, 3:378–387 postharvest physiology, 1:204–236; 3:59–143; 10:35–62; 11:15–43 postpollination development, 19:1–58 protea leaf blackening, 17:173–201 pruning, 8:359–362 raspberry, 11:187–188 regulation in floriculture, 7:416–424 rhododendron, 12:1–42 rose, 9:60–66 scents, 24:31–53 senescence, 1:204–236; 3:59–143; 10:35–62; 11:15–43; 18:1–85 strawberry, 28:325–349 sugars, 4:114 thin cell layer morphogenesis, 14:239–256 tulip, 5:57–59 water relations, 18:1–85

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382 Fluid drilling, 3:1–58 Foliage plants: acclimatization, 6:119–154 fertilization, 1:102–103; 5:367–380 Foliar nutrition, 6:287–355 Freeze protection. See Frost protection Frost: apple fruit set, 1:407–408 citrus, 7:201–238 protection, 11:45–109 Fruit: abscission, 1:172–203 abscission, citrus, 15:145–182 apple anatomy and morphology, 10:283–297 apple bioregulation, 10:348–374 apple bitter pit, 11:289–355 apple flavor, 16:197–234 apple maturity indices, 13:407–432 apple ripening and quality, 10:361–374 apple scald, 27:227–267 apple weight loss, 25:197–234 avocado development and ripening, 10:229–271 bloom delay, 15:97–144 blueberry development, 13:378–390 cactus physiology, 18:335–341 CA storage and quality, 8:101–127 chilling injury, 15:63–95 coating physiology, 26:161–238 cracking, 19:217–262 diseases in CA storage, 3:412–461 drop, apple functional phytochemicals, 27:269–315 growth measurement, 24:373–431 kiwifruit, 6:35–48; 12:316–318 loquat, 23:233–276 lychee, 28:433–444 maturity indices, 13:407–432 navel orange, 8:129–179 nectarine, postharvest, 11:413–452 nondestructive postharvest quality evaluation, 20:1–119 olive processing, 25:235–260 peach, postharvest, 11:413–452 pear, bioregulation, 10:348–374 pear, fruit disorders, 11:357–411 pear maturity indices, 13:407–432 pear ripening and quality, 10:361–374 pear scald, 27:227–267

CUMULATIVE SUBJECT INDEX pear volatiles, 28:237–324 pistachio, 3:382–391 phytochemicals, 28:125–185 plum, 23:179–231 quality and pruning, 8:365–367 ripening, 5:190–205 set, 1:397–424; 4:153–154 set in navel oranges, 8:140–142 size and thinning, 1:293–294; 4:161 softening, 5:109–219; 10:107–152 splitting, 19:217–262 strawberry growth and ripening, 17:267–297 texture, 20:121–224 thinning, apple and pear, 10:353–359 tomato parthenocarpy, 6:65–84 tomato ripening, 13:67–103 volatiles, pear, 28:237–324 Fruit crops: see also individual crop alternate bearing, 4:128–173 apple bitter pit, 11:289–355 apple flavor, 16:197–234 apple fruit splitting and cracking, 19:217–262 apple germplasm, 29:1–61, 63–303 apple growth, 11:229–287 apple maturity indices, 13:407–432 apple scald, 27:227–267 apple, wild of Kazakhstan, 29:63–303, 305–315 apricot, origin and dissemination, 22:225–266 apricot, wild of Kazakhstan, 29–325–326 avocado flowering, 8:257–289 avocado rootstocks, 17:381–429 barberry, wild of Kazakhstan, 29:332–336 berry crop harvesting, 16:255–382 bilberry, wild of Kazakhstan, 29:347–348 blackberry, wild of Kazakhstan, 29:345 bloom delay, 15:97–144 blueberry developmental physiology, 13:339–405 blueberry harvesting, 16:257–282 blueberry nutrition, 10:183–227 bramble harvesting, 16:282–298 cactus, 18:302–309 carbohydrate reserves, 10:403–430

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CUMULATIVE SUBJECT INDEX CA and MA for tropicals, 22:123–183 CA storage, 1:301–336 CA storage diseases, 3:412–461 cherry origin, 19:263–317 cherry, wild of Kazakhstan, 29:326–330 chilling injury, 15:145–182 chlorosis, 9:161–165 citrus abscission, 15:145–182 citrus cold hardiness, 7:201–238 citrus, culture of young trees, 24:319–372 citrus dwarfing by viroids, 24:277–317 citrus flowering, 12:349–408 cotoneaster, wild of Kazakhstan, 29:316–317 cranberry, 21:215–249 cranberry harvesting, 16:298–311 cranberry, wild of Kazakhstan, 29:349 currant harvesting, 16:311–327 currant, wild of Kazakhstan, 29:341 deficit irrigation, 21:105–131 dormancy release, 7:239–300 elderberry, wild of Kazakhstan, 29:349–350 Ericaceae nutrition, 10:183–227 fertilization, 1:104–106 fig, industry, 12:409–490 fireblight, 11:423–474 flowering, 12:223–264 foliar nutrition, 6:287–355 frost control, 11:45–109 gooseberry, wild of Kazakhstan, 29:341–342 grape flower anatomy and morphology, 13:315–337 grape harvesting, 16:327–348 grape irrigation, 27:189–225 grape nitrogen metabolism, 14:407–452 grape pruning, 16:235–254, 336–340 grape root, 5:127–168 grape seedlessness, 11:164–176 grape, wild of Kazakhstan, 29:342–343 grapevine pruning, 16:235–254, 336–340 honey bee pollination, 9:244–250, 254–256 jojoba, 17:233–266 in vitro culture, 7:157–200; 9:273–349 irrigation, deficit, 21:105–131 kiwifruit, 6:1–64; 12:307–347

383 lingonberry, 27:79–123 lingonberry, wild of Kazakhstan, 29:348–349 longan, 16:143–196 loquat, 23:233–276 lychee, 16:143–196, 28:393–453 mountain ash, wild of Kazakhstan, 29:322–324 mulberry, wild of Kazakhstan, 29:350–351 muscadine grape breeding, 14:357–405 navel orange, 8:129–179 nectarine postharvest, 11:413–452 nondestructive postharvest quality evaluation, 20:1–119 nutritional ranges, 2:143–164 oleaster, wild of Kazakhstan, 29:351–353 olive salinity tolerance, 21:177–214 orange, navel, 8:129–179 orchard floor management, 9:377–430 peach origin, 17:331–379 peach postharvest, 11:413–452 peach thinning, 28:351–392 pear fruit disorders, 11:357–411; 27:227–267 pear maturity indices, 13:407–432 pear scald, 27:227–267 pear volatiles, 28:237–324 pear, wild of Kazakhstan, 29:315–316 pecan flowering, 8:217–255 photosynthesis, 11:111–157 Phytophthora control, 17:299–330 plum origin, 23:179–231 plum, wild of Kazakhstan, 29:330–332 pruning, 8:339–380 rambutan, 16:143–196 raspberry, 11:185–228 raspberry, wild of Kazakhstan, 29:343–345 roots, 2:453–457 rose, wild of Kazakhstan, 29:353–360 sapindaceous fruits, 16:143–196 sea buckthorn, wild of Kazakhstan, 29:361 short life and replant problem, 2:1–116 strawberry fruit growth, 17:267–297 strawberry harvesting, 16:348–365 strawberry, wild of Kazakhstan, 29:347 summer pruning, 9:351–375

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384 Fruit crops (cont.) Vaccinium nutrition, 10:183–227 vacciniums, wild of Kazakhstan, 29:347–349 viburnam, wild of Kazakhstan, 29:361–362 virus elimination, 28:187–236 water status, 7:301–344 Functional phytochemicals, fruit, 27:269–315 Fungi: fig, 12:451–474 mushroom, 6:85–118 mycorrhiza, 3:172–213; 10:211–212 pathogens in postharvest storage, 3:412–461 truffle cultivation, 16:71–107 Fungicide, and apple fruit set, 1:416 G Galanthus, 25:22–25 Garlic, CA storage, 1:375 Genetic variation: alternate bearing, 4:146–150 photoperiodic response, 4:82 pollution injury, 8:16–19 temperature-photoperiod interaction, 17:73–123 wild apple, 29:63–303 Genetics and breeding: aroids (edible), 8:72–75; 12:169 aroids (ornamental), 10:18–25 bean, bacterial resistance, 3:28–58 bloom delay in fruits, 15:98–107 bulbs, flowering, 18:119–123 cassava, 12:164 chestnut blight resistance, 8:313–321 citrus cold hardiness, 7:221–223 cranberry, 21:236–239 embryogenesis, 1:23 fig, 12:432–433 fire blight resistance, 1:435–436 flowering, 15:287–290, 303–305, 306–309, 314–315; 27:1–39, 41–77 flower longevity, 1:208–209 ginseng, 9:197–198 grafting use, 28:109–115 in vitro techniques, 9:318–324; 18:119–123

CUMULATIVE SUBJECT INDEX lettuce, 2:185–187 lingonberry, 27:108–111 loquat, 23:252–257 muscadine grapes, 14:357–405 mushroom, 6:100–111 navel orange, 8:150–156 nitrogen nutrition, 2:410–411 pineapple, 21:138–164 plant regeneration, 3:278–283 pollution insensitivity, 8:18–19 potato tuberization, 14:121–124 rhododendron, 12:54–59 sweet potato, 12:175 sweet sorghum, 21:87–90 tomato parthenocarpy, 6:69–70 tomato ripening, 13:77–98 tree short life, 2:66–70 Vigna, 2:311–394 waxes, 23:50–53 woody legume tissue and cell culture, 14:311–314 yam (Dioscorea), 12:183 Geophyte. See Bulb, tuber Geranium, fertilization, 5:355–357 Germination, seed, 2:117–141, 173–174; 24:229–275 Germplasm: acquisition, apple, 29:1–61 characterization, apple, 29:45–56 cryopreservation, 6:357–372 in vitro, 5:261–264; 9:324–325 pineapple, 21:133–175 resources, apple, 29:1–61 Gibberellin: abscission, citrus, 15:166–167 bloom delay, 15:111–114 citrus, abscission, 15:166–167 cold hardiness, 11:63 dormancy, 7:270–271 floral promoter, 4:114 flowering, 15:219–293, 315–318 genetic regulation, 16:15 grape root, 5:150–151 mechanical stress, 17:19–20 Ginseng, 9:187–236 Girdling, 4:251–252 Glucosinolates, 19:99–215 Gooseberry, wild of Kazakhstan, 29:341–342 Gourd, history, 25:71–171

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CUMULATIVE SUBJECT INDEX Graft and grafting: herbaceous, 28:61–124 incompatibility, 15:183–232 phase change, 7:136–137, 141–142 rose, 9:56–57 Grape: CA storage, 1:308 chlorosis, 9:165–166 flower anatomy and morphology, 13:315–337 functional phytochemicals, 27:291–298 irrigation, 27:189–225 harvesting, 16:327–348 muscadine breeding, 14:357–405 nitrogen metabolism, 14:407–452 pollen morphology, 13:331–332 pruning, 16:235–254, 336–340 root, 5:127–168 seedlessness, 11:159–187 sex determination, 13:329–331 wild of Kazakhstan, 29:342–343 Gravitropism, 15:233–278 Greenhouse and greenhouse crops: carbon dioxide, 7:357–360, 544–545 energy efficiency, 1:141–171; 9:1–52 growth substances, 7:399–481 nutrition and fertilization, 5:317–403 pest management, 13:1–66 vegetables, 21:1–39 Growth regulators. See Growth substances Growth substances, 2:60–66; 24:55–138, see also Abscisic acid, Auxin, Cytokinins, Ethylene, Gibberellins abscission, citrus, 15:157–176 apple bioregulation, 10:309–401 apple dwarfing, 3:315–375 apple fruit set, 1:417 apple thinning, 1:270–300 aroids, ornamental, 10:14–18 avocado fruit development, 10:229–243 bloom delay, 15:107–119 CA storage in vegetables, 1:346–348 cell cultures, 3:214–314 chilling injury, 15:77–83 citrus abscission, 15:157–176 cold hardiness, 7:223–225; 11:58–66 dormancy, 7:270–279 embryogenesis, 1:41–43; 2:277–281

385 floriculture, 7:399–481 flower induction, 4:190–195 flowering, 15:290–296 flower storage, 10:46–51 genetic regulation, 16:1–32 ginseng, 9:226 grape seedlessness, 11:177–180 hormone reception, 26:49–84 in vitro flowering, 4:112–115 mechanical stress, 17:16–21 meristem and shoot-tip culture, 5:221–227 navel oranges, 8:146–147 pear bioregulation, 10:309–401 petal senescence, 3:76–78 phase change, 7:137–138, 142–143 raspberry, 11:196–197 regulation, 11:1–14 rose, 9:53–73 seedlessness in grape, 11:177–180 triazole, 10:63–105 H Haemanthus, 25:25–28 Halo blight of beans, 3:44–45 Hardiness, 4:250–251 Harvest: flower stage, 1:211–212 index, 7:72–74 lettuce, 2:176–181 mechanical of berry crops, 16:255–382 Hawthorne, wild of Kazakhstan, 29:317–322 Hazelnut. See Filbert wild of Kazakhstan, 29:365–366 Health phytochemicals: fruit, 27:269–315 vegetables, 28:125–185 Heat treatment (postharvest), 22:91–121 Heliconia, 14:1–55 Herbaceous plants, subzero stress, 6:373–417 Hippeastrum, 25:29–34 Histochemistry: flower induction, 4:177–179 fruit abscission, 1:172–203 Histology, flower induction, 4:179–184, see also Anatomy and morphology

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386 Honey bee, 9:237–272 Honeysuckle, wild of Kazakhstan, 29:350 Horseradish, CA storage, 1:368 Hydrolases, 5:169–219 Hydroponic culture, 5:1–44; 7:483–558 Hymenocallis, 25:59 Hypovirulence, in Endothia parasitica, 8:299–310 I Ismene, 25:59 Ice, formation and spread in tissues, 13:215–255 Ice-nucleating bacteria, 7:210–212; 13:230–235 Industrial crops, cactus, 18:309–312 Insects and mites: aroids, 8:65–66 avocado pollination, 8:275–277 fig, 12:442–447 hydroponic crops, 7:530–534 integrated pest management, 13:1–66 lettuce, 2:197–198 ornamental aroids, 10:18 tree short life, 2:52 tulip, 5:63, 92 waxes, 23:1–68 Integrated pest management: greenhouse crops, 13:1–66 In vitro: abscission, 15:156–157 apple propagation, 10:325–326 aroids, ornamental, 10:13–14 artemisia, 19:342–345 bioreactor technology, 24:1–30 bulbs, flowering, 18:87–169 cassava propagation, 13:121–123; 26:99–119 cellular salinity tolerance, 16:33–69 cold acclimation, 6:382 cryopreservation, 6:357–372 embryogenesis, 1:1–78; 2:268–310; 7:157–200; 10:153–181 environmental control, 17:123–170 flowering bulbs, 18:87–169 flowering, 4:106–127 pear propagation, 10:325–326 phase change, 7:144–145

CUMULATIVE SUBJECT INDEX propagation, 3:214–314; 5:221–277; 7:157–200; 9:57–58, 273–349; 17:125–172 thin cell layer morphogenesis, 14:239–264 woody legume culture, 14:265–332 Iron: deficiency and toxicity symptoms in fruits and nuts, 2:150 deficiency chlorosis, 9:133–186 Ericaceae nutrition, 10:193–195 foliar application, 6:330 nutrition, 5:324–325 pine bark media, 9:123 Irrigation: deficit, deciduous orchards, 21:105–131 drip or trickle, 4:1–48 frost control, 11:76–82 fruit trees, 7:331–332 grape, 27:189–225 grape root growth, 5:140–141 lettuce industry, 2:175 navel orange, 8:161–162 root growth, 2:464–465 J Jojoba, 17:233–266 Juvenility, 4:111–112 pecan, 8:245–247 tulip, 5:62–63 woody plants, 7:109–155 K Kale, fluid drilling of seed, 3:21 Kazakhstan, see Wild fruits and nuts Kiwifruit: botany, 6:1–64 vine growth, 12:307–347 L Lamps, for plant growth, 2:514–531 Lanzon, CA and MA, 22:149 Leaves: apple morphology, 12:283–288 flower induction, 4:188–189 Leek: CA storage, 1:375 fertilization, 1:118

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CUMULATIVE SUBJECT INDEX Leguminosae, in vitro, 5:227–229; 14:265–332 Lemon, rootstock, 1:244–246, see also Citrus Lettuce: CA storage, 1:369–371 classification, 28:25–27 fertilization, 1:118 fluid drilling of seed, 3:14–17 industry, 2:164–207 seed germination, 24:229–275 tipburn, 4:49–65 Leucojum, 25:34–39 Leucospermum, 22:27–90 Light: fertilization, greenhouse crops, 5:330–331 flowering, 15:282–287, 310–312 fruit set, 1:412–413 lamps, 2:514–531 nitrogen nutrition, 2:406–407 orchards, 2:208–267 ornamental aroids, 10:4–6 photoperiod, 4:66–105 photosynthesis, 11:117–121 plant growth, 2:491–537 tolerance, 18:215–246 Lingonberry, 27:79–123 wild of Kazakhstan, 29:348–349 Longan: See also Sapindaceous fruits CA and MA, 22:150 Loquat: botany and horticulture, 23:233–276 CA and MA, 22:149–150 Lychee: See also Sapindaceous fruits CA and MA, 22:150 flowering, 28:397–421 fruit abscission, 28–437–443 fruit development, 28:433–436 pollination, 28:422–428 reproductive biology, 28:393–453 Lycoris, 25:39–43 M Magnesium: container growing, 9:84–85 deficiency and toxicity symptoms in fruits and nuts, 2:148 Ericaceae nutrition, 10:196–198 foliar application, 6:331

387 nutrition, 5:323 pine bark media, 9:117–119 Magnetic resonance imaging, 20:78–86, 225–266 Male sterility, temperature-photoperiod induction, 17:103–106 Mandarin, rootstock, 1:250–252 Manganese: deficiency and toxicity symptoms in fruits and nuts, 2:150–151 Ericaceae nutrition, 10:189–193 foliar application, 6:331 nutrition, 5:235–326 pine bark media, 9:123–124 Mango: alternate bearing, 4:145–146 asexual embryogenesis, 7:171–173 CA and MA, 22:151–157 CA storage, 1:313 in vitro culture, 7:171–173 Mangosteen, CA and MA, 22:157 Mechanical harvest, berry crops, 16:255–382 Mechanical stress regulation, 17:1–42 Media: fertilization, greenhouse crops, 5:333 pine bark, 9:103–131 Medicinal crops: artemisia, 19:319–371 poppy, 19:373–408 Melon grafting, 28:96–98 Meristem culture, 5:221–277 Metabolism: flower, 1:219–223 nitrogen in citrus, 8:181–215 seed, 2:117–141 Micronutrients: container growing, 9:85–87 pine bark media, 9:119–124 Micropropagation, see also In vitro; propagation: bulbs, flowering, 18:89–113 environmental control, 17:125–172 nuts, 9:273–349 rose, 9:57–58 temperate fruits, 9:273–349 tropical fruits and palms, 7:157–200 Microtus. See Vole Modified atmosphere (MA) for tropical fruits, 22:123–183 Moisture, and seed storage, 2:125–132

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388 Molecular biology: cassava, 26:85–159 floral induction, 27:3–20 flowering, 27:1–39;41–77 hormone reception, 26:49–84 Molybdenum nutrition, 5:328–329 Monocot, in vitro, 5:253–257 Monstera. See Aroids, ornamental Morphology: navel orange, 8:132–133 orchid, 5:283–286 pecan flowering, 8:217–243 Moth bean, genetics, 2:373–374 Mountain ash, wild of Kazakhstan, 29:322–324 Mulberry, wild of Kazakhstan, 29:350–351 Mung bean, genetics, 2:348–364 Mushroom: CA storage, 1:371–372 cultivation, 19:59–97 spawn, 6:85–118 Muskmelon, fertilization, 1:118–119 Mycoplasma-like organisms, tree short life, 2:50–51 Mycorrhizae: container growing, 9:93 Ericaceae, 10:211–212 fungi, 3:172–213 grape root, 5:145–146 N Narcissus, 25:43–48 Navel orange, 8:129–179 Nectarine: bloom delay, 15:105–106 CA storage, 1:309–310 postharvest physiology, 11:413–452 Nematodes: aroids, 8:66 fig, 12:475–477 lettuce, 2:197–198 tree short life, 2:49–50 Nerine, 25:48–56 NFT (nutrient film technique), 5:1–44 Nitrogen: CA storage, 8:116–117 container growing, 9:80–82 deficiency and toxicity symptoms in fruits and nuts, 2:146

CUMULATIVE SUBJECT INDEX Ericaceae nutrition, 10:198–202 fixation in woody legumes, 14:322–323 foliar application, 6:332 in embryogenesis, 2:273–275 metabolism in apple, 4:204–246 metabolism in citrus, 8:181–215 metabolism in grapevine, 14:407–452 nutrition, 2:395, 423; 5:319–320 pine bark media, 9:108–112 trickle irrigation, 4:29–30 vegetable crops, 22:185–223 Nomenclature, 28:1–60 Nondestructive quality evaluation of fruits and vegetables, 20:1–119 Nursery crops: fertilization, 1:106–112 nutrition, 9:75–101 Nut crops: see also individual crop almond postharvest technology and utilization, 20:267–311 almond, wild of Kazakhstan, 29:262–265 chestnut blight, 8:291–336 fertilization, 1:106 hazelnut, wild of Kazakhstan, 29:365–366 honey bee pollination, 9:250–251 in vitro culture, 9:273–349 nutritional ranges, 2:143–164 pine, wild of Kazakhstan, 29:368–369 pistachio culture, 3:376–396 pistachio, wild of Kazakhstan, 29:366–368 walnut, wild of Kazakhstan, 29:369–370 Nutrient: concentration in fruit and nut crops, 2:154–162 film technique, 5:1–44 foliar-applied, 6:287–355 media, for asexual embryogenesis, 2:273–281 media, for organogenesis, 3:214–314 plant and tissue analysis, 7:30–56 solutions, 7:524–530 uptake, in trickle irrigation, 4:30–31 Nutrition (human): aroids, 8:79–84 CA storage, 8:101–127 phytochemicals in fruit, 27:269–315

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Page 389

CUMULATIVE SUBJECT INDEX phytochemicals in vegetables, 28:125–185 steroidal alkaloids, 25:171–196 Nutrition (plant): air pollution, 8:22–23, 26 blueberry, 10:183–227 calcifuge, 10:183–227 cold hardiness, 3:144–171 container nursery crops, 9:75–101 cranberry, 21:234–235 ecologically based, 24:156–172 embryogenesis, 1:40–41 Ericaceae, 10:183–227 fire blight, 1:438–441 foliar, 6:287–355 fruit and nut crops, 2:143–164 ginseng, 9:209–211 greenhouse crops, 5:317–403 kiwifruit, 12:325–332 mycorrhizal fungi, 3:185–191 navel orange, 8:162–166 nitrogen in apple, 4:204–246 nitrogen in vegetable crops, 22:185–223 nutrient film techniques, 5:18–21, 31–53 ornamental aroids, 10:7–14 pine bark media, 9:103–131 raspberry, 11:194–195 slow-release fertilizers, 1:79–139 O Oil palm: asexual embryogenesis, 7:187–188 in vitro culture, 7:187–188 Okra: botany and horticulture, 21:41–72 CA storage, 1:372–373 Oleaster, wild of Kazakhstan, 29:351–353 Olive: alternate bearing, 4:140–141 salinity tolerance, 21:177–214 processing technology, 25:235–260 Onion: CA storage, 1:373–375 fluid drilling of seed, 3:17–18 Opium poppy, 19:373–408 Orange: see also Citrus alternate bearing, 4:143–144 sour, rootstock, 1:242–244

389 sweet, rootstock, 1:252–253 trifoliate, rootstock, 1:247–250 Orchard and orchard systems: floor management, 9:377–430 light, 2:208–267 root growth, 2:469–470 water, 7:301–344 Orchid: fertilization, 5:357–358 pollination regulation of flower development, 19:28–38 physiology, 5:279–315 Organogenesis, 3:214–314, see also In vitro; tissue culture Ornamental plants: see also individual plant Amaryllidaceae Banksia, 22:1–25 cactus grafting, 28–106–109 chlorosis, 9:168–169 cotoneaster, wild of Kazakhstan, 29:316–317 fertilization, 1:98–104, 106–116 flowering bulb roots, 14:57–88 flowering bulbs in vitro, 18:87–169 foliage acclimatization, 6:119–154 heliconia, 14:1–55 honeysuckle, wild of Kazakhstan, 29:350 Leucospermum, 22:27–90 oleaster, wild of Kazakhstan, 29:351–353 orchid pollination regulation, 19:28–38 poppy, 19:373–408 protea leaf blackening, 17:173–201 rhododendron, 12:1–42 rose, wild of Kazakhstan, 29:353–360 viburnam, wild of Kazakhstan, 29:361–362 P Paclobutrazol. See Triazole Papaya: asexual embryogenesis, 7:176–177 CA and MA, 22:157–160 CA storage, 1:314 in vitro culture, 7:175–178 Parsley: CA storage, 1:375 drilling of seed, 3:13–14 Parsnip, fluid drilling of seed, 3:13–14

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390 Parthenocarpy, tomato, 6:65–84 Passion fruit: in vitro culture, 7:180–181 CA and MA, 22:160–161 Pathogen elimination, in vitro, 5:257–261 Peach: bloom delay, 15:105–106 CA storage, 1:309–310 origin, 17:333–379 postharvest physiology, 11:413–452 short life, 2:4 summer pruning, 9:351–375 thinning, 28:351–392 wooliness, 20:198–199 Peach palm (Pejibaye): in vitro culture, 7:187–188 Pear: bioregulation, 10:309–401 bloom delay, 15:106–107 CA storage, 1:306–308 decline, 2:11 fire blight control, 1:423–474 fruit disorders, 11:357–411; 27:227–267 fruit volatiles, 28:237–324 in vitro, 9:321 maturity indices, 13:407–432 root distribution, 2:456 scald, 27:227–267 short life, 2:6 wild of Kazakhstan, 29:315–316 Pecan: alternate bearing, 4:139–140 fertilization, 1:106 flowering, 8:217–255 in vitro culture, 9:314–315 Pejibaye, in vitro culture, 7:189 Pepper (Capsicum): CA storage, 1:375–376 fertilization, 1:119 fluid drilling in seed, 3:20 grafting, 28:104–105 phytochemicals, 28:161–162 Persimmon: CA storage, 1:314 quality, 4:259 Pest control: aroids (edible), 12:168–169 aroids (ornamental), 10:18 cassava, 12:163–164

CUMULATIVE SUBJECT INDEX cowpea, 12:210–213 ecologically based, 24:172–201 fig, 12:442–477 fire blight, 1:423–474 ginseng, 9:227–229 greenhouse management, 13:1–66 hydroponics, 7:530–534 sweet potato, 12:173–175 vertebrate, 6:253–285 yam (Dioscorea), 12:181–183 Petal senescence, 11:15–43 pH: container growing, 9:87–88 fertilization greenhouse crops, 5:332–333 pine bark media, 9:114–117 soil testing, 7:8–12, 19–23 Phase change, 7:109–155 Phenology: apple, 11:231–237 raspberry, 11:186–190 Philodendron. See Aroids, ornamental Phosphonates, Phytophthora control, 17:299–330 Phosphorus: container growing, 9:82–84 deficiency and toxicity symptoms in fruits and nuts, 2:146–147 nutrition, 5:320–321 pine bark media, 9:112–113 trickle irrigation, 4:30 Photoautotrophic micropropagation, 17:125–172 Photoperiod, 4:66–105, 116–117; 17:73–123 flowering, 15:282–284, 310–312 Photosynthesis: cassava, 13:112–114 efficiency, 7:71–72; 10:378 fruit crops, 11:111–157 ginseng, 9:223–226 light, 2:237–238 Physiology: see also Postharvest physiology bitter pit, 11:289–355 blueberry development, 13:339–405 cactus reproductive biology, 18:321–346 calcium, 10:107–152 carbohydrate metabolism, 7:69–108

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CUMULATIVE SUBJECT INDEX cassava, 13:105–129 citrus cold hardiness, 7:201–238 conditioning 13:131–181 cut flower, 1:204–236; 3:59–143; 10:35–62 desiccation tolerance, 18:171–213 disease resistance, 18:247–289 dormancy, 7:239–300 embryogenesis, 1:21–23; 2:268–310 floral scents, 24:31–53 flower development, 19:1–58 flowering, 4:106–127 fruit ripening, 13:67–103 fruit softening, 10:107–152 ginseng, 9:211–213 glucosinolates, 19:99–215 grafting, 28:78–84 heliconia, 14:5–13 hormone reception, 26:49–84 juvenility, 7:109–155 lettuce seed germination, 24:229–275 light tolerance, 18:215–246 loquat, 23:242–252 lychee reproduction, 28:393–453 male sterility, 17:103–106 mechanical stress, 17:1–42 nitrogen metabolism in grapevine, 14:407–452 nutritional quality and CA storage, 8:118–120 olive salinity tolerance, 21:177–214 orchid, 5:279–315 petal senescence, 11:15–43 photoperiodism, 17:73–123 pollution injury, 8:12–16 polyamines, 14:333–356 potato tuberization, 14:89–188 pruning, 8:339–380 raspberry, 11:190–199 regulation, 11:1–14 root pruning, 6:158–171 roots of flowering bulbs, 14:57–88 rose, 9:3–53 salinity hormone action, 16:1–32 salinity tolerance, 16:33–69 seed, 2:117–141 seed priming, 16:109–141 strawberry flowering, 28:28:325–349 subzero stress, 6:373–417 summer pruning, 9:351–375

391 sweet potato, 23:277–338 thin cell layer morphogenesis, 14:239–264 tomato fruit ripening, 13:67–103 tomato parthenocarpy, 6:71–74 triazoles, 10:63–105; 24:55–138 tulip, 5:45–125 vernalization, 17:73–123 volatiles, 17:43–72 watercore, 6:189–251 water relations cut flowers, 18:1–85 waxes, 23:1–68 Phytochemicals, functional: fruits, 27:269–315 vegetables, 28:125–185 Phytohormones. See Growth substances Phytophthora control, 17:299–330 Phytotoxins, 2:53–56 Pigmentation: flower, 1:216–219 rose, 9:64–65 Pinching, by chemicals, 7:453–461 Pine, wild of Kazakhstan, 29:368–369 Pineapple: CA and MA, 22:161–162 CA storage, 1:314 genetic resources, 21:138–141 in vitro culture, 7:181–182 Pine bark, potting media, 9:103–131 Pistachio: alternate bearing, 4:137–139 culture, 3:376–393 in vitro culture, 9:315 wild of Kazakhstan, 29:366–368 Plantain: CA and MA, 22:141–146 in vitro culture, 7:178–180 Plant: classification, 28:1–60 protection, short life, 2:79–84 systematics, 28:1–60 Plastic cover, sod production, 27:317–351 Plum: CA storage, 1:309 origin, 23:179–231 wild of Kazakhstan, 29:330–332 Poinsettia, fertilization, 1:103–104; 5:358–360 Pollen, desiccation tolerance, 18:195

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392 Pollination: apple, 1:402–404 avocado, 8:272–283 cactus, 18:331–335 embryogenesis, 1:21–22 fig, 12:426–429 floral scents, 24:31–53 flower regulation, 19:1–58 fruit crops, 12:223–264 fruit set, 4:153–154 ginseng, 9:201–202 grape, 13:331–332 heliconia, 14:13–15 honey bee, 9:237–272 kiwifruit, 6:32–35 lychee, 28:422–428 navel orange, 8:145–146 orchid, 5:300–302 petal senescence, 11:33–35 protection, 7:463–464 rhododendron, 12:1–67 Pollution, 8:1–42 Polyamines, 14:333–356 chilling injury, 15:80 Polygalacturonase, 13:67–103 Poppy, opium, 19:373–408 Postharvest physiology: almond, 20:267–311 apple bitter pit, 11:289–355 apple maturity indices, 13:407–432 apple scald, 27:227–257 apple weight loss, 25:197–234 aroids, 8:84–86 asparagus, 12:69–155 CA for tropical fruit, 22:123–183 CA storage and quality, 8:101–127 chlorophyll fluorescence, 23:69–107 coated fruits & vegetables, 26:161–238 cut flower, 1:204–236; 3:59–143; 10:35–62 foliage plants, 6:119–154 fruit, 1:301–336 fruit softening, 10:107–152 heat treatment, 22:91–121 lettuce, 2:181–185 low-temperature sweetening, 17:203–231 MA for tropical fruit, 22:123–183 navel orange, 8:166–172 nectarine, 11:413–452

CUMULATIVE SUBJECT INDEX nondestructive quality evaluation, 20:1–119 pathogens, 3:412–461 peach, 11:413–452 pear disorders, 11:357–411; 27:227–267 pear maturity indices, 13:407–432 pear scald, 27:227–257 petal senescence, 11:15–43 protea leaf blackening, 17:173–201 quality evaluation, 20:1–119 scald, 27:227–267 seed, 2:117–141 texture in fresh fruit, 20:121–244 tomato fruit ripening, 13:67–103 vegetables, 1:337–394 watercore, 6:189–251; 11:385–387 Potassium: container growing, 9:84 deficiency and toxicity symptoms in fruits and nuts, 2:147–148 foliar application, 6:331–332 nutrition, 5:321–322 pine bark media, 9:113–114 trickle irrigation, 4:29 Potato: CA storage, 1:376–378 classification, 28:23–26 fertilization, 1:120–121 low temperature sweetening, 17:203–231 phytochemicals, 28:160–161 tuberization, 14:89–198 Processing, table olives, 25:235–260 Propagation: see also In vitro apple, 10:324–326; 12:288–295 aroids, ornamental, 10:12–13 bioreactor technology, 24:1–30 cassava, 13:120–123 floricultural crops, 7:461–462 ginseng, 9:206–209 orchid, 5:291–297 pear, 10:324–326 rose, 9:54–58 tropical fruit, palms 7:157–200 woody legumes in vitro, 14:265–332 Protaceous flower crop: see also Protea Banksia, 22:1–25 Leucospermum, 22:27–90 Protea, leaf blackening, 17:173–201

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CUMULATIVE SUBJECT INDEX Protected crops, carbon dioxide, 7:345–398 Protoplast culture, woody species, 10:173–201 Pruning, 4:161; 8:339–380 apple, 9:351–375 apple training, 1:414 chemical, 7:453–461 cold hardiness, 11:56 fire blight, 1:441–442 grapevines, 16:235–254 light interception, 2:250–251 peach, 9:351–375 phase change, 7:143–144 root, 6:155–188 Prunus: see also Almond; Cherry; Nectarine; Peach; Plum in vitro, 5:243–244; 9:322 root distribution, 2:456 Pseudomonas: phaseolicola, 3:32–33, 39, 44–45 solanacearum, 3:33 syringae, 3:33, 40; 7:210–212 Pumpkin, history, 25:71–170 Q Quality evaluation: fruits and vegetables, 20:1–119, 121–224 nondestructive, 20:1–119 texture in fresh fruit, 20:121–224 R Rabbit, 6:275–276 Radish, fertilization, 1:121 Rambutan. See Sapindaceous fruits CA and MA, 22:163 Raspberry: harvesting, 16:282–298 productivity, 11:185–228 wild of Kazakhstan, 29:343–345 Rejuvenation: rose, 9:59–60 woody plants, 7:109–155 Replant problem, deciduous fruit trees, 2:1–116 Respiration: asparagus postharvest, 12:72–77

393 fruit in CA storage, 1:315–316 kiwifruit, 6:47–48 vegetables in CA storage, 1:341–346 Rhizobium, 3:34, 41 Rhododendron, 12:1–67 Rice bean, genetics, 2:375–376 Root: apple, 12:269–272 cactus, 18:297–298 diseases, 5:29–31 environment, nutrient film technique, 5:13–26 Ericaceae, 10:202–209 grape, 5:127–168 kiwifruit, 12:310–313 physiology of bulbs, 14:57–88 pruning, 6:155–188 raspberry, 11:190 rose, 9:57 tree crops, 2:424–490 Root and tuber crops: Amaryllidaceae, 25:1–79 aroids, 8:43–99; 12:166–170 cassava, 12:158–166; 26:85–159 low-temperature sweetening, 17:203–231 minor crops, 12:184–188 potato tuberization, 14:89–188 sweet potato, 12:170–176 sweet potato physiology, 23:277–338 yam (Dioscorea), 12:177–184 Rootstocks: alternate bearing, 4:148 apple, 1:405–407; 12:295–297 avocado, 17:381–429 citrus, 1:237–269 cold hardiness, 11:57–58 fire blight, 1:432–435 light interception, 2:249–250 navel orange, 8:156–161 root systems, 2:471–474 stress, 4:253–254 tree short life, 2:70–75 Rosaceae, in vitro, 5:239–248 Rose: fertilization, 1:104; 5:361–363 growth substances, 9:3–53 in vitro, 5:244–248 wild of Kazakhstan, 29:353–360

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394 S Salinity: air pollution, 8:25–26 olive, 21:177–214 soils, 4:22–27 tolerance, 16:33–69 Sapindaceous fruits, 16:143–196 Sapodilla, CA and MA, 22:164 Scadoxus, 25:25–28 Scald, apple and pear, 27:227–265 Scoring, and fruit set, 1:416–417 Sea buckthorn, wild of Kazakhstan, 29:361 Seed: abortion, 1:293–294 apple anatomy and morphology, 10:285–286 conditioning, 13:131–181 desiccation tolerance, 18:196–203 environmental influences on size and composition, 13:183–213 flower induction, 4:190–195 fluid drilling, 3:1–58 grape seedlessness, 11:159–184 kiwifruit, 6:48–50 lettuce, 2:166–174 lettuce germination, 24:229–275 priming, 16:109–141 rose propagation, 9:54–55 vegetable, 3:1–58 viability and storage, 2:117–141 Secondary metabolites, woody legumes, 14:314–322 Senescence: chlorophyll senescence, 23:88–93 cut flower, 1:204–236; 3:59–143; 10:35–62; 18:1–85 petal, 11:15–43 pollination-induced, 19:4–25 rose, 9:65–66 whole plant, 15:335–370 Sensory quality: CA storage, 8:101–127 Shoot-tip culture, 5:221–277, see also Micropropagation Short life problem, fruit crops, 2:1–116 Signal transduction, 26:49–84 Small fruit, CA storage, 1:308 Snapdragon fertilization, 5:363–364 Sod production, 27:317–351

CUMULATIVE SUBJECT INDEX Sodium, deficiency and toxicity symptoms in fruits and nuts, 2:153–154 Soil: grape root growth, 5:141–144 management and root growth, 2:465–469 orchard floor management, 9:377–430 plant relations, trickle irrigation, 4:18–21 stress, 4:151–152 testing, 7:1–68; 9:88–90 zinc, 23:109–178 Soilless culture, 5:1–44 Solanaceae: in vitro, 5:229–232 steroidal alkaloids, 25:171–196 Somatic embryogenesis. See Asexual embryogenesis Sorghum, sweet, 21:73–104 Spathiphyllum. See Aroids, ornamental Squash, history, 25:71–170 Stem, apple morphology, 12:272–283 Sternbergia, 25:59 Steroidal alkaloids, solanaceous, 25:171–196 Storage: see also Postharvest physiology, Controlled-atmosphere (CA) storage cut flower, 3:96–100; 10:35–62 rose plants, 9:58–59 seed, 2:117–141 Strawberry: fertilization, 1:106 flowering, 28:325–349 fruit growth and ripening, 17:267–297 functional phytonutrients, 27: 303–304 harvesting, 16:348–365 in vitro, 5:239–241 wild of Kazakhstan, 29:347 Stress: benefits of, 4:247–271 chlorophyll fluorescence, 23:69–107 climatic, 4:150–151 flooding, 13:257–313 mechanical, 17:1–42 petal, 11:32–33 plant, 2:34–37 protectants (triazoles), 24:55–138 protection, 7:463–466

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CUMULATIVE SUBJECT INDEX salinity tolerance in olive, 21:177–214 subzero temperature, 6:373–417 waxes, 23:1–68 Sugar: see also Carbohydrate allocation, 7:74–94 flowering, 4:114 Sugar apple, CA and MA, 22:164 Sugar beet, fluid drilling of seed, 3:18–19 Sulfur: deficiency and toxicity symptoms in fruits and nuts, 2:154 nutrition, 5:323–324 Sweet potato: culture, 12:170–176 fertilization, 1:121 physiology, 23:277–338 Sweet sop, CA and MA, 22:164 Symptoms, deficiency and toxicity symptoms in fruits and nuts, 2:145–154 Syngonium. See Aroids, ornamental Systematics, 28:1–60 T Taro. See Aroids, edible Taxonomy, 28:1–60 Tea, botany and horticulture, 22:267–295 Temperature: apple fruit set, 1:408–411 bloom delay, 15:119–128 CA storage of vegetables, 1:340–341 chilling injury, 15:67–74 cryopreservation, 6:357–372 cut flower storage, 10:40–43 fertilization, greenhouse crops, 5:331–332 fire blight forecasting, 1:456–459 flowering, 15:284–287, 312–313 interaction with photoperiod, 4:80–81 low temperature sweetening, 17:203–231 navel orange, 8:142 nutrient film technique, 5:21–24 photoperiod interaction, 17:73–123 photosynthesis, 11:121–124 plant growth, 2:36–37 seed storage, 2:132–133

395 subzero stress, 6:373–417 Texture in fresh fruit, 20:121–224 Thinning: apple, 1:270–300 peach and Prunus, 28:351–392 Tipburn, in lettuce, 4:49–65 Tissue: see also In vitro culture 1:1–78; 2:268–310; 3:214–314; 4:106–127; 5:221–277; 6:357–372; 7:157–200; 8:75–78; 9:273–349; 10:153–181, 24:1–30 cassava, 26:85–159 dwarfing, 3:347–348 nutrient analysis, 7:52–56; 9:90 Tomato: CA storage, 1:380–386 classification, 28:21–23 chilling injury, 20:199–200 fertilization, 1:121–123 fluid drilling of seed, 3:19–20 fruit ripening, 13:67–103 galacturonase, 13:67–103 grafting, 28:98–103 greenhouse quality, 26:239 parthenocarpy, 6:65–84 phytochemicals, 28:160 Toxicity symptoms in fruit and nut crops, 2:145–154 Transport, cut flowers, 3:100–104 Tree decline, 2:1–116 Triazoles, 10:63–105; 24:55–138 chilling injury, 15:79–80 Trickle irrigation, 4:1–48 Truffle cultivation, 16:71–107 Tuber, potato, 14:89–188 Tuber and root crops. See Root and tuber crops Tulip: See also Bulb fertilization, 5:364–366 in vitro, 18:144–145 physiology, 5:45–125 Tunnel (cloche), 7:356–357 Turfgrass, fertilization, 1:112–117 Turnip, fertilization, 1:123–124 Turnip Mosaic Virus, 14:199–238 U Urd bean, genetics, 2:364–373 Urea, foliar application, 6:332

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396 V Vaccinium, 10:185–187, see also Blueberry; Cranberry; Lingonberry functional phytonutrients, 27:303 wild of Kazakhstan, 29:347–349 Vase solutions, 3:82–95; 10:46–51 Vegetable crops: see also Specific crop Allium phytochemicals, 28:156–159 aroids, 8:43–99; 12:166–170 asparagus postharvest, 12:69–155 cactus, 18:300–302 cassava, 12:158–166; 13:105–129; 26:85–159 CA storage, 1:337–394 CA storage and quality, 8:101–127 CA storage diseases, 3:412–461 Caper bush, 27:125–188 chilling injury, 15:63–95 coating physiology, 26:161–238 crucifer phytochemicals, 28:150–156 cucumber grafting, 28:91–96 ecologically based, 24:139–228 eggplant grafting, 28:103–104 eggplant phytochemicals, 28:162–163 fertilization, 1:117–124 fluid drilling of seeds, 3:1–58 gourd history, 25:71–170 grafting, 28:61–124 greenhouse management, 21:1–39 greenhouse pest management, 13:1–66 honey bee pollination, 9:251–254 hydroponics, 7:483–558 lettuce seed germination, 24:229–275 low-temperature sweetening, 17:203–231 melon grafting, 28:96–98 minor root and tubers, 12:184–188 mushroom cultivation, 19:59–97 mushroom spawn, 6:85–118 N nutrition, 22:185–223 nondestructive postharvest quality evaluation, 20:1–119 okra, 21:41–72 pepper phytochemicals, 28:161–162 phytochemicals, 28:125–185 potato phytochemicals, 28:160–161 potato tuberization, 14:89–188 pumpkin history, 25:71–170 seed conditioing, 13:131–181

CUMULATIVE SUBJECT INDEX seed priming, 16:109–141 squash history, 25:71–170 steroidal alkaloids, Solanaceae, 25:171–196 sweet potato, 12:170–176 sweet potato physiology, 23:277–338 tomato fruit ripening, 13:67–103 tomato (greenhouse) quality: 26:239–319 tomato parthenocarpy, 6:65–84 tomato phytochemicals, 28:160 tropical production, 24:139–228 truffle cultivation, 16:71–107 watermelon grafting, 28:86–91 yam (Dioscorea), 12:177–184 Vegetative tissue, desiccation tolerance, 18:176–195 Vernalization, 4:117; 15:284–287; 17:73–123 Vertebrate pests, 6:253–285 Viburnam, wild of Kazakhstan, 29:361–362 Vigna: see also Cowpea genetics, 2:311–394 U.S. production, 12:197–222 Viroid, dwarfing for citrus, 24:277–317 Virus: benefits in horticulture, 3:394–411 dwarfing for citrus, 24:277–317 elimination, 7:157–200; 9:318; 18:113–123; 28:187–236 fig, 12:474–475 tree short life, 2:50–51 turnip mosaic, 14:199–238 Volatiles, 17:43–72; 24:31–53; 28:237–324 Vole, 6:254–274 W Walnut: in vitro culture, 9:312 wild of Kazakhstan, 29:369–370 Water relations: cut flower, 3:61–66; 18:1–85 deciduous orchards, 21:105–131 desiccation tolerance, 18:171–213 fertilization, grape and grapevine, 27:189–225 kiwifruit, 12:332–339 light in orchards, 2:248–249

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CUMULATIVE SUBJECT INDEX photosynthesis, 11:124–131 trickle irrigation, 4:1–48 Watercore, 6:189–251 apple, 6:189–251 pear, 11:385–387 Watermelon: fertilization, 1:124 grafting, 28:86–91 Wax apple, CA and MA, 22:164 Waxes, 23:1–68 Weed control, ginseng, 9:228–229 Weeds: lettuce research, 2:198 virus, 3:403 Wild fruit and nuts of Kazakhstan, 29:305–371 almond, 29:262–265 apple, 29:63–303, 305–315 apricot, 29:325–326 barberry, 29:332–336 bilberry, 29:347–348 blackberry, 29:345 cherry, 29:326–330 cotoneaster, 29:316–317 cranberry, 29:349 currant, 29:341 elderberry, 29:349–350 gooseberry, 29:341–342 grape, 29:342–343 hazelnut, 29:365–366 lingonberry, 29:348–349 mountain ash, 29:322–324 mulberry, 29:350–351 oleaster, 29:351–353 pear, 29:315–316 pine, 29:368–369 pistachio, 29:366–368 plum, 29:330–332

397 raspberry, 29:343–345 rose, 29:353–360 sea buckthorn, 29:361 strawberry, 29:347 vacciniums, 29:347–349 viburnam, 29:361–362 walnut, 29:369–370 Woodchuck, 6:276–277 Woody species, somatic embryogenesis, 10:153–181 X Xanthomonas phaseoli, 3:29–32, 41, 45–46 Xanthophyll cycle, 18:226–239 Xanthosoma, 8:45–46, 56–57, see also Aroids Sugar: see also Carbohydrate allocation, 7:74–94 flowering, 4:114 Y Yam (Dioscorea), 12:177–184 Yield: determinants, 7:70–74; 97–99 limiting factors, 15:413–452 Z Zantedeschia. See Aroids, ornamental Zephyranthes, 25:60–61 Zinc: deficiency and toxicity symptoms in fruits and nuts, 2:151 foliar application, 6:332, 336 nutrition, 5:326; 23:109–178 pine bark media, 9:124

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Cumulative Contributor Index (Volumes 1–29)

Abbott, J.A., 20:1 Adams III, W.W., 18:215 Aldwinckle, H.S., 1:423; 15:xiii, 29:1 Amarante, C., 28:161 Anderson, I.C., 21:73 Anderson, J.L., 15:97 Anderson, P.C., 13:257 Andrews, P.K., 15:183 Ashworth, E.N., 13:215; 23:1 Asokan, M.P., 8:43 Atkinson, D., 2:424 Aung, L.H., 5:45

Bower, J.P., 10:229 Bradley, G.A., 14:xiii Brandenburg, W., 28:1 Brennan, R., 16:255 Broadbent, P., 24:277 Broschat, T.K., 14:1 Brown, S. 15:xiii Buban, T., 4:174 Bukovac, M.J., 11:1 Burke, M.J., 11:xiii Buwalda, J.G., 12:307 Byers, R.E., 6:253; 28:351

Bailey, W.G., 9:187 Baird, L.A.M., 1:172 Banks, N.H., 19:217; 25:197; 26:161 Barden, J.A., 9:351 Barker, A.V., 2:411 Bass, L.N., 2:117 Bassett, C. L., 26:49 Becker, J.S., 18:247 Beer, S.V., 1:423 Behboudian, M.H., 21:105; 27:189 Bennett, A.B., 13:67 Benschop, M., 5:45 Ben-Ya’acov, A., 17:381 Benzioni, A., 17:233 Bevington, K.B., 24:277 Bewley, J.D., 18:171 Binzel, M.L., 16:33 Blanpied, G.D., 7:xi Bliss, F.A., 16:xiii; 28:xi Boardman, K. 27 xi Borochov, A., 11:15

Caldas, L.S., 2:568 Campbell, L.E., 2:524 Cantliffe, D.J., 16:109; 17:43; 24:229; 28:325 Carter, G., 20:121 Carter, J.V., 3:144 Cathey, H.M., 2:524 Chambers, R.J., 13:1 Chandler, C.K. 28:325 Charron, C.S., 17:43 Chen, Z., 25:171 Chin, C.K., 5:221 Clarke, N.D., 21:1 Coetzee, J. H., 26:1 Cohen, M., 3:394 Collier, G.F., 4:49 Collins, G., 25:235 Collins, W.L., 7:483 Colmagro, S., 25:235 Compton, M.E., 14:239 Conover, C.A., 5:317; 6:119

Horticultural Reviews, Volume 29, Edited by Jules Janick ISBN 0-471-21968-1 © 2003 John Wiley & Sons, Inc. 399

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400 Coppens d’Eeckenbrugge, G., 21:133 Costa, G. 28:351 Coyne, D.P., 3:28 Crane, J.C., 3:376 Criley, R.A., 14:1; 22:27; 24:x Crowly, W., 15:1 Cutting, J.G., 10:229 Daie, J., 7:69 Dale, A., 11:185; 16:255 Darnell, R.L., 13:339, 28:325 Davenport, T.L., 8:257; 12:349 Davies, F.S., 8:129; 24:319 Davies, P.J., 15:335 Davis, T.D., 10:63; 24:55 Decker, H.F., 27:317 DeEll, J.R., 23:69 DeGrandi–Hoffman, G., 9:237 De Hertogh, A.A., 5:45; 14:57; 18:87; 25:1 Deikman, J., 16:1 DellaPenna, D., 13:67 Demmig-Adams, B., 18:215 Dennis, F.G., Jr., 1:395 Dickson, E.E ., 29:1 Dorais, M., 26:239 Doud, S.L., 2:1 Dudareva, N., 24:31 Duke, S.O., 15:371 Dunavent, M.G., 9:103 Duval, M.-F., 21:133 Düzyaman, E., 21:41 Dyer, W.E., 15:371 Dzhangaliev, A.D., 29:63, 305 Early, J.D., 13:339 Eastman, K., 28:125 Elfving, D.C., 4:1; 11:229 El-Goorani, M.A., 3:412 Esan, E.B., 1:1 Evans, D.A., 3:214 Ewing, E.E., 14:89 Faust, M., 2:vii, 142; 4:174; 6:287; 14:333; 17:331; 19:263; 22:225; 23:179 Fenner, M., 13:183 Fenwick, G.R., 19:99 Ferguson, A.R., 6:1 Ferguson, I.B., 11:289 Ferguson, J.J., 24:277

CUMULATIVE CONTRIBUTOR INDEX Ferguson, L., 12:409 Ferree, D.C., 6:155 Ferreira, J.F.S., 19:319 Fery, R.L., 2:311; 12:157 Fischer, R.L., 13:67 Fletcher, R.A., 24:53 Flick, C.E., 3:214 Flore, J.A., 11:111 Forshey, C.G., 11:229 Forsline, P.L., 29:ix; 1 Franks, R. G., 27:41 Fujiwara, K., 17:125 Gazit, S., 28:393 Geisler, D., 6:155 Geneve, R.L., 14:265 George, W.L., Jr., 6:25 Gerrath, J.M., 13:315 Gilley, A., 24:55 Giovannetti, G., 16:71 Giovannoni, J.J., 13:67 Glenn, G.M., 10:107 Goffinet, M.C., 20:ix Goldschmidt, E.E., 4:128 Goldy, R.G., 14:357 Goren, R., 15:145 Gosselin, A., 26:239 Goszczynska, D.M., 10:35 Grace, S.C., 18:215 Graves, C.J., 5:1 Gray, D., 3:1 Grierson, W., 4:247 Griffen, G.J., 8:291 Grodzinski, B., 7:345 Gucci, R., 21:177 Guest, D.I., 17:299 Guiltinan, M.J., 16:1 Hackett, W.P., 7:109 Halevy, A.H., 1:204; 3:59 Hallett, I.C., 20:121 Hammerschmidt, R., 18:247 Hanson, E.J., 16:255 Harker, F.R., 20:121 Heaney, R.K., 19:99 Heath, R.R., 17:43 Helzer, N.L., 13:1 Hendrix, J.W., 3:172 Henny, R.J., 10:1 Hergert, G.B., 16:255

3926 P-04 (index)

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Page 401

CUMULATIVE CONTRIBUTOR INDEX Hess, F.D., 15:371 Hetterscheid, W.L.A., 28:1 Heywood, V., 15:1 Hjalmarsson, I., 27:79–123 Hogue, E.J., 9:377 Hokanson, S.C. 29:1 Holt, J.S., 15:371 Huber, D.J., 5:169 Hunter, E.L., 21:73 Hutchinson, J.F., 9:273 Hutton, R.J., 24:277 Indira, P., 23:277 Ingle, M. 27:227 Isenberg, F.M.R., 1:337 Iwakiri, B.T., 3:376 Jackson, J.E., 2:208 Janick, J., 1:ix; 8:xi; 17:xiii; 19:319; 21:xi; 23:233 Jarvis, W.R., 21:1 Jenks, M.A., 23:1 Jensen, M.H., 7:483 Jeong, B.R., 17:125 Jewett, T.J., 21:1 Joiner, J.N., 5:317 Jones, H.G., 7:301 Jones, J.B., Jr., 7:1 Jones, R.B., 17:173 Kagan-Zur, V., 16:71 Kalt, W. 27:269; 28:125 Kang, S.-M., 4:204 Kato, T., 8:181 Kawa, L., 14:57 Kawada, K., 4:247 Kelly, J.F., 10:ix; 22:xi Kester, D.E., 25:xii Khan, A.A., 13:131 Kierman, J., 3:172 Kim, K.-W., 18:87 Kinet, J.-M., 15:279 King, G.A., 11:413 Kingston, C.M., 13:407–432 Kirschbaum, D.S. 28:325 Kliewer, W.M., 14:407 Knight, R.J., 19:xiii Knox, R.B., 12:1 Kofranek, A.M., 8:xi Korcak, R.F., 9:133; 10:183

401 Kozai, T., 17:125 Krezdorn, A.H., 1:vii Kushad, M.M., 28:125 Lakso, A.N., 7:301; 11:111 Laimer, M., 28:187 Lamb, R.C., 15:xiii Lang, G.A., 13:339 Larsen, R.P., 9:xi Larson, R.A., 7:399 Leal, F., 21:133 Ledbetter, C.A., 11:159 Lee, J.-M., 28:61 Li, P.H., 6:373 Lill, R.E., 11:413 Lin, S., 23:233 Liu, Z., 27:41 Lipton, W.J., 12:69 Littlejohn, G.M., 26:1 Litz, R.E., 7:157 Lockard, R.G., 3:315 Loescher, W.H., 6:198 Lorenz, O.A., 1:79 Lu, R., 20:1 Luby,J.J., 29:1 Lurie, S., 22:91–121 Lyrene, P., 21:xi Maguire, K.M., 25:197 Manivel, L., 22:267 Maraffa, S.B., 2:268 Marangoni, A.G., 17:203 Marini, R.P., 9:351 Marlow, G.C., 6:189 Maronek, D.M., 3:172 Martin, G.G., 13:339 Masiunas, J., 28:125 Mayak, S., 1:204; 3:59 Maynard, D.N., 1:79 McConchie, R., 17:173 McNicol, R.J., 16:255 Merkle, S.A., 14:265 Michailides, T.J., 12:409 Michelson, E., 17:381 Mika, A., 8:339 Miller, A.R., 25:171 Miller, S.S., 10:309 Mills, H.A., 2:411; 9:103 Mills, T.M., 21:105 Mitchell, C.A., 17:1

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Page 402

402 Mizrahi, Y., 18:291, 321 Molnar, J.M., 9:1 Monk, G.J., 9:1 Monselise, S.P., 4:128 Moore, G.A., 7:157 Mor, Y., 9:53 Morris, J.R., 16:255 Murashige, T., 1:1 Murr, D.P., 23:69 Murray, S.H., 20:121 Myers, P.N., 17:1 Nadeau, J.A., 19:1 Nascimento, W.M., 24:229 Neilsen, G.H., 9:377 Nelson, P.V., 26:xi Nerd, A., 18:291, 321 Niemiera, A.X., 9:75 Nobel, P.S., 18:291 Nyujtò, F., 22:225 Oda, M., 28:61 O’Donoghue, E.M., 11:413 Ogden, R.J., 9:103 O’Hair, S.K., 8:43; 12:157 Oliveira, C.M., 10:403 Oliver, M.J., 18:171 O’Neill, S.D., 19:1 Opara, L.U., 19:217; 24:373; 25:197 Ormrod, D.P., 8:1 Ortiz, R., 27:79 Palser, B.F., 12:1 Papadopoulos, A.P., 21:1; 26:239 Pararajasingham, S., 21:1 Parera, C.A., 16:109 Paris, H.S., 25:71 Pegg, K.G., 17:299 Pellett, H.M., 3:144 Perkins-Veazil, P., 17:267 Pichersky, E., 24:31 Piechulla, B., 24:31 Ploetz, R.C., 13:257 Pokorny, F.A., 9:103 Poole, R.T., 5:317; 6:119 Poovaiah, B.W., 10:107 Portas, C.A.M., 19:99 Porter, M.A., 7:345

CUMULATIVE SUBJECT INDEX Possingham, J.V., 16:235 Prange, R.K., 23:69 Pratt, C., 10:273; 12:265 Predieri, S., 28:237 Preece, J.E., 14:265 Priestley, C.A., 10:403 Proctor, J.T.A., 9:187 Puonti–Kaerlas, J., 26:85 Quamme, H., 18:xiii Raese, J.T., 11:357 Ramming, D.W., 11:159 Rapparini, F., 28:237 Ravi, V., 23:277 Reddy, A.S.N., 10:107 Redgwell, R.J., 20:121 Reid, M., 12:xiii; 17:123 Reuveni, M., 16:33 Richards, D., 5:127 Rieger, M., 11:45 Roper, T.R., 21:215 Rosa, E.A.S., 19:99 Roth-Bejerano, N., 16:71 Roubelakis-Angelakis, K.A., 14:407 Rouse, J.L., 12:1 Royse, D.J., 19:59 Rudnicki, R.M., 10:35 Ryder, E.J., 2:164; 3:vii Sachs, R., 12:xiii Sakai, A., 6:357 Salisbury, F.B., 4:66; 15:233 Salova, T. H., 29:305 Saltveit, M.E., 23:x San Antonio, J.P., 6:85 Sankhla, N., 10:63; 24:55 Saure, M.C., 7:239 Schaffer, B., 13:257 Schenk, M.K., 22:185 Schneider, G.W., 3:315 Schuster, M.L., 3:28 Scorza, R., 4:106 Scott, J.W., 6:25 Sedgley, M., 12:223; 22:1; 25:235 Seeley, S.S., 15:97 Serrano Marquez, C., 15:183 Sharp, W.R., 2:268; 3:214

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Page 403

CUMULATIVE SUBJECT INDEX Sharpe, R.H., 23:233 Shattuck, V.I., 14:199 Shear, C.B., 2:142 Sheehan, T.J., 5:279 Shipp, J.L., 21:1 Shirra, M., 20:267 Shorey, H.H., 12:409 Simon, J.E., 19:319 Singh, Z. 27:189 Sklensky, D.E., 15:335 Smith, A.H., Jr., 28:351 Smith, M.A.L., 28:125 Smith, G.S., 12:307 Smock, R.M., 1:301 Sommer, N.F., 3:412 Sondahl, M.R., 2:268 Sopp, P.I., 13:1 Soule, J., 4:247 Sozzi, G. O., 27:125 Sparks, D., 8:217 Splittstoesser, W.E., 6:25; 13:105 Spooner, D.M., 28:1 Srinivasan, C., 7:157 Stang, E.J., 16:255 Steffens, G.L., 10:63 Stern, R.A., 28:393 Stevens, M.A., 4:vii Stroshine, R.L., 20:1 Struik, P.C., 14:89 Studman, C.J., 19:217 Stutte, G.W., 13:339 Styer, D.J., 5:221 Sunderland, K.D., 13:1 Sung, Y., 24:229 Surányi, D., 19:263; 22:225; 23:179 Swanson, B., 12:xiii Swietlik, D., 6:287; 23:109 Syvertsen, J.P., 7:301 Tattini, M., 21:177 Tétényi, P., 19:373 Theron, K.I., 25:1 Tibbitts, T.W., 4:49 Timon, B., 17:331 Tindall, H.D., 16:143 Tisserat, B., 1:1 Titus, J.S., 4:204 Trigiano, R.N., 14:265

403 Tunya, G.O., 13:105 Turekhanova, P.M., 29:305 Upchurch, B.L., 20:1 Valenzuela, H.R., 24:139 van Doorn, W.G., 17:173; 18:1 van den Berg, W.L.A., 28:1 van Kooten, O., 23:69 van Nocker, S. 27:1 Veilleux, R.E., 14:239 Vorsa, N., 21:215 Vizzotto, G., 28: 351 Wallace, A., 15:413 Wallace, D.H., 17:73 Wallace, G.A., 15:413 Wang, C.Y., 15:63 Wang, S.Y., 14:333 Wann, S.R., 10:153 Watkins, C.B., 11:289 Watson, G.W., 15:1 Webster, B.D., 1:172; 13:xi Weichmann, J., 8:101 Wetzstein, H.Y., 8:217 Whiley, A.W., 17:299 Whitaker, T.W., 2:164 White, J.W., 1:141 Williams, E.G., 12:1 Williams, M.W., 1:270 Wismer, W.V., 17:203 Wittwer, S.H., 6:xi Woodson, W.R., 11:15 Wright, R.D., 9:75 Wutscher, H.K., 1:237 Yada, R.Y., 17:203 Yadava, U.L., 2:1 Yahia, E.M., 16:197; 22:123 Yan, W., 17:73 Yarborough, D.E., 16:255 Yelenosky, G., 7:201 Zanini, E., 16:71 Zieslin, N., 9:53 Zimmerman, R.H., 5:vii; 9:273 Ziv, M., 24:1 Zucconi, F., 11:1

3:24 PM

Elite selections of Malus sieversii

2D. Fruit of M. sieversii from grow-out at PGRU of 4-year-old seedlings (from seed of elites among GMAL 3604–3649) from area/sites 9.00–9.05. Photo by P. Forsline, Geneva, NY, September 17, 2001.

2C. Fruit of M. sieversii from random population (GMAL 4308–4317) of 10 trees from area/site 12.00 with seeds only collected. Photo by P. Forsline, Kazakhstan, September 15, 1996.

10/15/02

Plate 1.

1B. Elite M. sieversii (GMAL 4049) from area/site 11.00 with both seeds and scions collected. Photo by P. Forsline, Kazakhstan, September 14, 1996.

1A. Elite M. sieversii (GMAL 3614) from area/site 9.03 with both seeds and scions collected. Photo by J. Luby, Kazakhstan, August 29, 1995.

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1C. Elite M. sieversii (GMAL 4054) from area/site 12.00 with both seeds and scions collected. Photo by P. Forsline, Kazakhstan, September 15, 1996.

3:25 PM

Fruit from populations of Malus sieversii.

1D. Fruit representing 13 elites of M. sieversii from area/sites 9.00–9.05 (from scions among GMAL 3604–3649) that have fruited at PGRU as 3year-old trees propagated on M.9 rootstock with ‘Empire’ and ‘Gala’, lower left/lower right as references. Photo by P. Forsline, Geneva, NY, September 17, 2001.

10/15/02

Plate 2.

2B. Fruit of M. sieversii from random population (GMAL 3970–4003) of 34 trees from area/site 6.01 with seeds only collected. Photo by P. Forsline, Kazakhstan, September 10, 1995.

2A. Fruit of M. sieversii from random population (GMAL 3820–3849) of 30 trees from area/site 9.02 with seeds only collected. Photo by P. Forsline, Kazakhstan, August 31, 1995.

3926 (HR29) color insert Page 2