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The Plight of the American
The primary focus of contemporary North American chestnut research is restoration of the American chestnut, Castanea dentata (Burnham et al., 1986; Miller et al., 1996; Wallace, 1987). The American chestnut was one of the most important timber and nut-producing trees of the eastern United States (Smith, 1950). In the early 20th century, it was completely eliminated from its ecological niche by the introduced fungal pathogen Endothia (Cryphonectria) parasitica (Anagnostakis, 1988, 1993a; Roane et al., 1986). The return of the chestnut to its place in the Appalachian forest canopy requires the concerted efforts of Government, University, and Private Foundation researchers and the work of dedicated volunteer enthusiasts. The two-part plan of action involves research on biological control of the chestnut blight disease (MacDonald and Fulbright, 1991) and breeding the trees for resistance to the fungus. A secondary goal of current research is to provide support for the establishment of a commercial chestnut industry (for nut production) based on improved cultivars. Chestnut research in the United States is coordinated through a single United States Department of Agriculture (USDA) Regional Research Project (Project Number: NE-140): Biological Improvement of Chestnut and Management of Chestnut Pathogens and Pests.
Identification and characterization of native and introduced germplasm are of fundamental importance to the success of the restoration effort. Breeding for resistance to the chestnut blight fungus depends on the use of chestnut trees with well characterized blight-resistants and on the continued availability of locally adapted American chestnut trees to use as parents. Back to Menu (Top of Page).
The American chestnut is not extinct. Populations of C. dentata persist as shrubs - actually suppressed seedlings - across a vast area of eastern North America (Griffin, 1992; Jaynes, 1974; Paillet, 1989,1993; Rutter et al., 1991). Survival of this very shade tolerant species varies greatly from site to site, with past land-use and forest management practices, elevation, latitude, slope, and rainfall patterns (Griffin et al., 1983, 1991; Paillet, 1984,1987). In general, survival appears better in the northern part of the native range and at higher elevations in the Appalachian mountains. The wild populations of surviving American chestnuts represent a very important genetic resource and are the source of much of the germplasm used in the regional breeding programs. Many of the surviving specimens appear as clumps of multiple stems arising from a common root system (clones), although the connections are not always evident. Larger stems may be physiologically mature (they may bloom). It is likely that these trees have survived because they have escaped blight infection to date, not because they are resistant to the fungus. The largest stems are often heavily cankered with blight and usually die before nuts are produced. Castanea dentata has grown well in some areas where it was planted outside of its native range. At some sites, in the upper Midwest and in the Pacific Northwest, it has escaped blight infection, and in Michigan it survives, in part due to the occurrence of natural biocontrol agents (Garrod et al., 1985). There is a naturalized population in Wisconsin (where blight was only recently discovered) that is under investigation as a model site for the study of biological control and American chestnut ecology (Paillet and Rutter, 1989; Tiedeman and Hasselkuss, 1975).
North American Castanea germplasm resources also include native populations of C. pumila var. pumila, and C. pumila var. ozarkensis as well as populations of the introduced species C. crenata, C. henryi, C. sativa, C. seguinii and C. mollissima. (Table 1). Cultivated forms are readily available from many commercial sources (Table 2). The most commonly planted species, especially in the East, is C. mollissima. Although many nut producing cultivars are propagated by nurseries and hobby orchardists, very few varieties have achieved success as an orchard crop (Jaynes, 1975). Of these, the most widely planted are the C. mollissima cultivars ‘Crane’ and ‘Nanking’ and the interspecific hybrids ‘Colossal’, ‘Skookum’, ‘Sleeping Giant’, ‘Eaton’, and the Dunstan hybrids for which US plant patents have been issued (Table 3). European and Euro-Japanese hybrid cultivars, including some of very recent introduction (Craddock and Pellegrino 1992), are grown only in the western states of California, Oregon and Washington. Cultivar evaluations are currently underway in Connecticut, Georgia, Michigan, Oregon, and Tennessee (Anagnostakis, 1996; Craddock et al., 1995).
The most complete collection of Castanea species and hybrids is maintained at the Connecticut Agricultural Experiment Station near New Haven, CT (Anagnostakis, 1993c).
Additional germplasm collections are being made from throughout the native range of C. dentata, and breeding orchards have been established at several locations (Table 4). Threatened populations of C. pumila are also being collected. Each breeding orchard will contain, ideally, a diverse population of local C. dentata types (Hebard, 1994b). This will allow us to choose an American parent for each new breeding line that is well adapted to the local growing conditions and will increase the likelihood that our future hybrids will grow well there. Rather than transplant from the wild, new additions to the breeding orchards are made by planting seed collected from natural populations, as grafts onto seedling rootstocks, or as rooted cuttings, when feasible. Advantages of using grafted clones include the possibility of earlier bloom for breeding, genotypic evaluations of the selections and lastly, from a conservation perspective, the parent clone will not be removed from its place in the woods. For example, scionwood (small twigs with dormant buds) were collected in late winter 1996-97 from labeled clones at Lula Lake, within the Tennessee River Gorge Trust and from several other sites on Lookout Mountain and Walden Ridge in Tennessee. The scionwood was grafted onto rootstocks growing in the propagation greenhouse and then transplanted into the breeding orchard. In this way, the surviving local trees were multiplied without risking loss of the parent clones (Craddock, 1997).
The search for additional sources of blight resistance includes other species of chestnut not well represented in the existing breeding programs as well as diverse cultivars of Japanese and Chinese chestnut. Soon after blight struck, USDA researchers introduced blight-resistant chestnut species from China and Japan into the US in an attempt to replace the American chestnut (Anagnostakis, 1989,1992a; Galloway, 1926). Although no substitute has yet been found for the "timber type" American tree (Burnham, 1986; Diller and Clapper, 1969; Schlarbaum et al., 1992), many Asian and Asian-American hybrid trees survive from the original introductions, some in forest-type plantations (Berry, 1992; Jaynes and Dierauf, 1982; Keys et al., 1975; Little and Diller, 1964) and others in orchard settings. Naturalized populations of C. mollissima are now present in Connecticut (Anagnostakis, 1993b) and of C. sativa in California and Oregon (Bhagwandin, 1994).
The US Department of Agriculture maintains a system of germplasm repositories for many fruit and nut crops. Although Castanea is included in the National Plant Germplasm System (accessions are indexed through the Germplasm Resources Information Network: http://www.ars-grin.gov/npgs/), the site designated for chestnut is unfortunately unable to house the collection. Several Castanea accessions are maintained (on a temporary basis) at the Corvallis site (site records available at: http://www.ars-grin.gov/ars/PacWest/Corvallis/ncgr/ncgr.html) for purposes of post-entry quarantine. Back to Menu (Top of Page).
of North American Castanea germplasm
Little is known about the amount and distribution of variation in North American Castanea species (Huang et al., 1994b). Variable traits include a number of important agronomical (Hebard, 1994a) and physiological (Huang et al., 1994a; Shain et al, 1994; Shain and Spalding, 1995) characters with significant effects on tree form, blight resistance, climatic adaptability, and nut quality (Anagnostakis, 1996; Senter et al., 1994). Genetic models have been proposed for several heritable traits (Table 5).
Much of our understanding of the genetic basis for blight resistance in chestnut comes from research begun in the 1930s by the USDA and continued at the Connecticut Agricultural Experiment Station (Anagnostakis, 1992; Burnham et al., 1986; Clapper, 1952; Crane et al., 1937; Diller and Clapper, 1969; Graves, 1950; Hebard and Shain, 1989; Huang et al., 1996; Jaynes, 1962, 1964; Thor, 1978). Great progress has been made towards a hybrid tree combining the timber form of the American with the blight resistance of the Chinese chestnut. The most promising hybrids to date are the result of the backcross breeding program outlined by the late Charles Burnham and supported by The American Chestnut Foundation (TACF). According to Burnham’s plan, the only desirable character from the Chinese parent is its blight resistance. The backcross method is used to dilute out all the other characteristics of the Chinese parent while preserving the timber form and adaptive characteristics of the American parent (Burnham, 1989).
Many of the parent trees used in the TACF breeding work were first- and second-generation Chinese-American hybrids from the earlier Connecticut and USDA programs. The TACF breeding collection at Meadowview, Virginia includes many hybrids and advanced selections from second and third backcrosses. Third backcrosses (B3s) are nearly indistinguishable from the recurrent parent, in this case American chestnut. When the third backcrosses are intercrossed with each other, the offspring (B3F2s) have a chance of inheriting the genes for blight resistance from both parents. A large, intensively studied population of F2 interspecific hybrids from a second backcross generation (B2F2s) is segregating for blight resistance and for many morphological and molecular markers (Hebard, 1994a; Kubisiak et al., 1997). Molecular markers have been used to map blight resistance in this hybrid population. As breeding tools, molecular markers will greatly facilitate selection of resistant progeny. Identification of molecular markers was done at the Southern Institute of Forest Genetics, Saucier, Mississippi (Kubisiak et al., 1997), at the University of Massachusetts (Bernatzky and Mulcahy, 1992) and at Auburn University (Huang et al., 1994a,b), with plant material supplied by all NE-140 members. Chinese chestnut-specific markers for resistance to chestnut blight disease have been identified, and the work is now being expanded to examine Japanese chestnut markers as well. A genetic linkage map was constructed by using the three-generation pedigree. Molecular markers were also associated with the inheritance of various leaf and stem traits (Table 5).
Resistance to the Oriental Chestnut Gall Wasp (Dryocosmus kuriphilus), a serious insect pest of chestnut, must also be included in all future breeding programs. All three chinquapin species (C. henryi, C. pumila var. ozarkensis and C. pumila var. pumila) seem to have valuable resistance to gall wasp, which is currently found in Georgia, Alabama, Tennessee, and North Carolina. Gall wasp-resistant cultivars of C. crenata and C. mollissima x C. crenata hybrids have been introduced recently from Japan and Korea. Back to Menu (Top of Page).
It was, of course, upon imported chestnut nursery stock that chestnut blight was introduced into North America from Japan (Anagnostakis, 1989, 1993a). More recently, the international exchange of chestnut germplasm has been vital to chestnut breeding and to the development of improved cultivars, and will continue to be important in the future. Movement of Castanea germplasm in and out of the United States is difficult and dangerous - but not impossible (Bassi and Craddock, 1993; Craddock and Pellegrino, 1994). Importation of any and all chestnut material is regulated by the USDA Animal and Plant Health Inspection Service (http://www.aphis.usda.gov/; USDA-APHIS, 4700 River Rd., Riverdale, MD 20737-1228 USA) and will require a Plant Protection Quarantine Permit for entry. The post entry quarantine period for Castanea germplasm is two years and must be at an APHIS approved site. Export requirements may depend on the country of destination regulations. In general, a phytosanitary certificate must accompany the material.
The restoration of the American chestnut will be successful if the current rate of progress is maintained in research on biological control of chestnut blight and chestnut breeding. Efforts are underway to conserve threatened North American Castanea germplasm. New biotechnologies are becoming available for germplasm conservation (Craddock and Lagersteadt 1987), characterization (Kubisiak et al., 1997) and transformation (Carraway et al., 1994). Better characterization and facilitated international exchange of germplasm will benefit the chestnut research and the chestnut industry in North America and throughout the world. Back to Menu (Top of Page).
Table 1. Castanea
germplasm resources in North America: species and distributions.
Distribution in North America
C. dentata Southern Ontario and Eastern USA
C. pumila var. pumila Cumberland Plateau and Southern Appalachian Mts.
C. pumila var. ozarkensis Ozark Plateau
C. pumila var. alnifolia Southeastern uplands and coastal plain
(C. ashei, C. floridana, C. paucispina) Deep South, Gulf Coast and Florida
C. dentata Upper Midwest, California, Oregon
C. crenata Northeastern USA, California
C. sativa California, Oregon (naturalized)
C. mollissima Widespread throughout the USA
C. henryi Rare (in cultivation, only)
C. seguinii Rare (in cultivation, only)
Table 2. Number of commercial
sources* for Castanea germplasm in North America. A list of
chestnut sources is available from S. Anagnostakis at The Connecticut Agricultural
Experiment Station**, and is updated yearly.
Number of Sources Offering:
Species Seed Seedlings Grafted Liners
C. dentata 1 12 0
C. pumila var. pumila 1 5 0
C. pumila var. ozarkensis 0 0 0
C. crenata 0 1 3
C. sativa 0 2 5
C. mollissima 2 48 19
C. seguinii 0 0 0
C. henryi 0 0 0
interspecific hybrids 0 27 10
*Adapted from Whealy et al., 1993
**web page: www.state.ct.us/caes or e-mail: email@example.com or telephone: 203-974-8498
Table 3. US Plant
Patents for Castanea Germplasm.
US Plant Patent #
Carolina 7041 Chestnut Hill Nursery, Alachua FL
Revival 5537 Chestnut Hill Nursery, Alachua FL
Willamette 7195 Chestnut Hill Nursery, Alachua FL
Heritage 6574 Chestnut Hill Nursery, Alachua FL
Table 4. An (incomplete)
listing of North American Castanea Germplasm Repository and
National Clonal Germplasm Repository - Corvallis, (Dr. Kim Hummer, Curator), USDA-ARS, 33447 Peoria Road, Corvallis, OR 97333 USA. Few accessions; temporary site, only. Approved for post-entry quarantine.
National Germplasm Repository - Brownwood, (Dr. L. J. Grauke, Curator), USDA-ARS, Rt. 2, Box 133, Somerville, Texas 77879 USA. No accessions at present time. Approved for post-entry quarantine.
American Chestnut Foundation Research Farm, (Dr. Fred Hebard, Superintendent), Rt. 1 Box 17, Meadowview, VA 24361 USA. Largest breeding program in the USA.
American Chestnut Cooperators Foundation, (Dr. Gary Griffin, Curator), 2667 Forest Service Road 708, Newport, VA 24128 USA. Breeding collection of C. dentata.
Badgersett Research Farm, (Mr. Philip Rutter), RR1 Box 141, Canton, MN 55922-9740 USA. Private breeding collection, northern ecotypes.
University of Tennessee, Dept. of Forestry, Wildlife and Fisheries, (Dr. Scott Schlarbaum), PO Box 1071, Knoxville, TN 37901-1071 USA. Large collection of cultivars, species and hybrids.
University of Tennessee at Chattanooga, Dept. of Biological and Environmental Sciences, (Dr. J .Hill Craddock), 615 McCallie Ave., Chattanooga, TN 37403-2598 USA. Breeding collection of southern C. dentata ecotypes.
Connecticut Agricultural Experiment Station, (Dr. Sandra Anagnostakis), PO Box 1106, New Haven, CT 06504 USA. May be the most complete collection of Castanea species and hybrids in the world.
Beltsville Agricultural Research Center, USDA-ARS, Beltsville MD USA. Remnants of discontinued USDA breeding orchards; records lost.
Burnt Ridge Nursery, (Michael Dolan, private grower), 432 Burnt Ridge Road, Onalaska Washington 98570 USA. More than 50 accessions. Approved for post-entry quarantine.
Empire Chestnut Co., (Dr. Greg Miller, private
grower, president of the U.S. Chestnut Marketing Association), 3276 Empire
Rd. SW, Carrollton, OH 44615 USA. A large number of cultivar and
Table 5. Inheritance
of leaf and stem morphological traits and resistance to chestnut blight
(From: Hebard, 1994a and Kubisiak et al., 1997).
No. genes Dominant parent
Inh1, Inh2 interveinal leaf hair 2 Chinese C
Vnh vein hair density 1 Chinese C
Twh1, Twh2 twig hair density 2 Chinese C
Stp1, Stp2 stipule size 2 American C
Red1 red stem color 1 American C
Red2 red stem color 1 American A
Red3 red stem color 1 American I
Cbr1 blight resistance 1 American B
Cbr2 blight resistance 1 American F
Cbr3 blight resistance 1 American G
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Craddock, J.H. and Lagerstedt, H.B. (1987) Cryopreservation of pollen at the National Clonal Germplasm Repository (abstract). HortScience 23(1): 28-29.
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Craddock, J.H., Sauve, R. and Schlarbaum, S.E. (1995) Chestnut variety testing in Tennessee. Tennessee Nurserymen's Association Newsletter 7(29): 20, 22.
Crane, H.L., Reed, C.A., and Wood, M.N. (1937) Nut Breeding. Pages 827-890 in: USDA Yearbook of Agriculture, Washington, DC.
Diller, J.D. and Clapper, R.B. (1969) Asiatic and hybrid chestnut trees in the eastern United States. J. For. 67: 328-331.
Galloway, B.T. (1926) The search in foreign countries for blight-resistant chestnuts and related tree crops. USDA Department Circular #383, Washington, DC, 16 pp.
Graves, A.H. (1950) Relative blight resistance in species and hybrids of Castanea. Phytopathology 40: 1125-1131.
Griffin, G.J. (1992) American chestnut survival in understory mesic sites following the chestnut blight pandemic. Can. J. Bot. 70:1950-1956.
Griffin, G.J., Hebard, F.V., Wendt, R.W. and Elkins, J.R. (1983) Survival of American chestnut trees: Evaluation of blight resistance and virulence of Endothia parasitica. Phytopathology 73: 1084-1092.
Griffin, G.J., Smith, H.C., Dietz, A, and Elkins, J.R. (1991) Importance of hardwood competition to American chestnut survival, growth, and blight development in forest clearcuts. Can. J. Bot .69:1804-1809.
Hebard, F.V. (1994a) Inheritance of juvenile leaf and stem morphological traits in crosses of Chinese and American chestnut. J. Heredity 85: 440-446.
Hebard, F.V. (1994b) The American Chestnut Foundation breeding plan: Beginning and intermediate steps. J. Amer. Chestnut Found. 8: 21-28.
Hebard, F.V. and Shain, L. (1989) Screening for resistance to chestnut blight. J. Amer. Chestnut Found. 4(1): 37-42.
Huang, H., Carey, W.A., Dane, F., and Norton, J.D. (1996) Evaluation of Chinese chestnut cultivars for resistance to Cryphonectria parasitica. Plant Dis. 80: 45-47.
Huang, H., Dane, F., and Norton, J.D. (1994a) Genetic analysis of 11 polymorphic isozyme loci in chestnut species and characterization of chestnut cultivars by multi-locus allozyme genotypes. J. Amer. Soc. Hort. Soc. 119: 840-849.
Huang, H., Dane, F., and Norton, J.D. (1994b) Allozyme diversity in Chinese, Seguin and American chestnut (Castanea spp.). Theor. Appl. Genet. 88: 840-849.
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Jaynes, R.A. (1964) Interspecific crosses in the genus Castanea. Silvae Genetica 13: 146-154.
Jaynes, R.A. (1974) Germplasm reserves of North American Nut Trees. Fruit Varieties 28(4): 95-99.
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Thor, E. Breeding of American chestnut. Pages 7-10 in: MacDonald, W.L., Cech, F.C., Luchok, J., and Smith, C. (eds). (1978) Proceedings of the American Chestnut Symposium . West Virginia University Books, Morgantown. 122 pp.
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Wallace, R.D. (eds.) Chestnuts and creating a commercial chestnut industry. Proc. 2nd PNW Chestnut Cong., 22-23 August 1987, Corvallis, OR. Chestnut Growers Exchange, Portland, OR. 122 pp.
Whealy, K., Demuth, S., Thuente, J., and Adelmann, A. (1993) Fruit, Berry and Nut Inventory. Second Edition. An inventory of nursery catalogs listing all fruit, berry and nut varieties available by mail order in the United States. Seed Saver Publications, Decorah, Iowa. 518 pp. ISBN 0-882424-51-4 and 0-882424-50-6.
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