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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.


3926 P-02 8/8/02 4:20 PM Page 304
3926 P- FM 8/8/02 4:18 PM Page i

HORTICULTURAL REVIEWS
Volume 29

WILD APPLE AND FRUIT


TREES OF CENTRAL ASIA
3926 P- FM 8/8/02 4:18 PM Page ii

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
3926 P- FM 8/8/02 4:18 PM Page iii

HORTICULTURAL REVIEWS
Volume 29

WILD APPLE AND FRUIT


TREES OF CENTRAL ASIA

edited by
Jules Janick
Purdue University

John Wiley & Sons, Inc.


3926 P- FM 8/8/02 4:18 PM Page iv

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,
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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 9 8 7 6 5 4 3 2 1
3926 P- FM 8/8/02 4:18 PM Page v

Contents

Contributors vii

Dedication: Calvin R. Sperling 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. Introduction 2
II. Germplasm Acquisition 12
III. Central Asian Collections 24
IV. Maintenance 37
V. Distribution 40
VI. Characterization and Evalution 45
VII. Utilization 56
VIII. Conclusion 57
Literature Cited 58

2. The Wild Apple Tree of Kazakhstan 63


A. D. Dzhangaliev

I. Introduction 65
II. Historical Review 69
III. The Role of Wild Apple in the Vegetative Cover
of Mountain Regions in Kazakhstan 83
IV. The Influence of Wild Apples on the Structure
of the Environment 121
V. Characteristics of Wild Apple Growth and
Development 156
VI. Intraspecific Polymorphism of Wild Apple 207

v
3926 P- FM 8/8/02 4:18 PM Page vi

vi CONTENTS

VII. Utility and Biochemical Characterization of Wild


Apple Fruit 246
VIII. Preservation of Wild Apples 272
IX. Conclusion 280
Literature Cited 285

3. The Wild Fruit and Nut Plants of Kazakhstan 305


A. D. Dzhangaliev, T. N. Salova, and P. M. Turekhanova

I. Introduction 308
II. Pome Fruits 311
III. Stone Fruits 324
IV. Small and Vine Fruits 332
V. Other Fruits 349
VI. Nuts 362
Literature Cited 370

Subject Index 373

Cumulative Subject Index 375

Cumulative Contributor Index 399


3926 P- FM 8/8/02 4:18 PM Page vii

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 Ser-
vice, Plant Genetic Resources Unit, Cornell University, Geneva, NY, 14456-
0462
Stan C. Hokanson, University of Minnesota, Department of Horticultural Sci-
ence, 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

vii
3926 P- FM 8/8/02 4:18 PM Page viii

Calvin R. Sperling
3926 P- FM 8/8/02 4:18 PM Page ix

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 preser-
vation 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 pur-
sued advanced degrees, he made four trips of two to five months dura-
tion 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 rel-
atives 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 coor-
dinated, 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 com-
mittees. These priorities resulted in explorations to acquire genetic

ix
3926 P- FM 8/8/02 4:18 PM Page x

x DEDICATION: CALVIN R. SPERLING

resources to fill in the gaps identified in the existing NPGS collections.


His responsibilities included implementation and coordination of explo-
ration for the NPGS, research on wild relatives and progenitors of cul-
tivated plants, and research on in situ preservation of crop genetic
resources. Dr. Sperling oversaw the management of nearly 100 success-
ful 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 Cen-
tral 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 intro-
duced 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 ethno-
botany in the NPGS agricultural explorations have been widely recog-
nized. 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 Con-
servation 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
2 P. FORSLINE, H. ALDWINCKLE, E. DICKSON, J. LUBY, AND S. HOKANSON

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 tempera-
tures 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 siever-
sii Lebed., a wild apple species native to Central Asia, has been recog-
nized 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
Table 1.1. Malus sections, series and primary species.

Sections Series Primary Species

Malus Langenf. Sieversinae Langenf. 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.
Baccatus Jiang Baccatae (Rehd.) Rehd. M. baccata (L.) Borkh.
var. mandshurica (Komorov.) Likh
var. sachalinensis (Juz.) Ponom.
var. himalaica (M.) Vass.
Hupehenses Langenf. M. hupehensis (Pampan.) Rehd.
M. halliana (Anon.) Koehne
Sikkimenses Jing M. sikkimensis (Wenzig) Koehne
Sorbomalus Zabel. Sieboldiane (Rehd.) M. sieboldii (Regel) Rehd.
var. sargenti (Rehd.)
Kansuenses (Rehd.) 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.

3
(continued)
4
Table 1.1. (continued)

Sections Series Primary Species

Yunnanenses Rehd. M. yunnanensis (French) Schneid.


M. prattii (Hemsl.) Schneid.
M. honanensis Rehd.
M. ombrophilla Hand.-Mazz.
Florentinae Rehd. M. florentina (Zuccagni) Schneid.
Chloromeles (Decne.) Rehd. M. ioensis (Wood.) Brit.
M. coronaria (L.) Mill.
M. angustifolia (Ait.) Michx.
Docyniopsis Schneid. M. doumeri (Bois.) Chev.
M. melliana (Hand.-Mazz.) Rehd.
M. tschonoskii (Maxim.) Schneid.
M. laosensis Chev.
Eriolobus (D.C.) Schneid. M. trilobata (Poiret) Schneid.

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. ×domestica Borkh. M. floribunda Siebold
M. ×hartwigii Koehne (halliana × baccata) M. ×magdeburgensis Schoch. (spectabilis × domestica)
M. ×micromalus Mak. (baccata × spectabilis) M. ×platycarpa Rehd. (cornonaria × domestica)
M. pumila Miller M. prunifolia (Willd.) Borkh.
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)

Adapted from Way et al. (1990); Langenfelds (1991); Ponomarenko (1992); Li (1996); Li (pers. com.).
1. COLLECTION AND UTILIZATION OF WILD APPLES OF CENTRAL ASIA 5

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 cen-
turies 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 sug-
gested 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 pre-
sent 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 ali-
mentary canals. Thus seedlings would have been randomly established
along the length of the trade routes and hybridization between previ-
ously 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
6 P. FORSLINE, H. ALDWINCKLE, E. DICKSON, J. LUBY, AND S. HOKANSON

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 con-
tributing parents include, Malus sylvestris the European crabapple bear-
ing small astringent, greenish-yellow fruits, native to an area from Britain
across Europe to the Balkans, and Malus baccata and some of its sub-
species or natural hybrids (M. mandshurica, M. prunifolia, and M. asi-
atica) on the eastern side of the trade routes. Recent evidence has shown
M. sylvestris is an unlikely contributor to the genetic makeup of the com-
mercial 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 Theophras-
tus (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 Amer-
ica 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 cul-
tivars of unknown parentage arose such as ‘Delicious’ (‘Hawkeye’),
‘Golden Delicious’, ‘McIntosh’, ‘Jonathan’, ‘Rome Beauty’, and ‘North-
ern 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 inae-
qualis (Cke.) Wint.) with the incorporation of the Vf gene from M. flori-
bunda 821 indicated the potential of interspecific crosses to introduce
3926 P-01 8/8/02 4:19 PM Page 7

1. COLLECTION AND UTILIZATION OF WILD APPLES OF CENTRAL ASIA 7

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 vol-
ume 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 exten-
sive 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 spe-
cific genetic traits. Approximately 3000 accessions are distributed annu-
ally. 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
3926 P-01 8/8/02 4:19 PM Page 8

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

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 coun-
terparts 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 sci-
entists from other institutions, contacts with scientists and government
officials in Central Asian countries were initiated and collaborative col-
lection 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 con-
tacts 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 Coop-
erative 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 expe-
dition 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 Sci-
ences 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.
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