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HORTICULTURAL REVIEWS
Volume 29
edited by
Jules Janick
Purdue University
HORTICULTURAL REVIEWS
Volume 29
HORTICULTURAL REVIEWS
Volume 29
edited by
Jules Janick
Purdue University
Copyright © 2003 by John Wiley & Sons, New York. All rights reserved.
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.
ISBN: 0-471-21968-1
ISSN: 0163-7851
10 9 8 7 6 5 4 3 2 1
3926 P- FM 8/8/02 4:18 PM Page v
Contents
Contributors vii
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
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
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
Contributors
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
ix
3926 P- FM 8/8/02 4:18 PM Page x
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
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
3
(continued)
4
Table 1.1. (continued)
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
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