HoR[ScicNc[ 41(5):1188-1194. 2006. Over the next several years, this unit was followed by the construction of 7 additional repositories at specifically chosen sites Fruit and Nut Genebanks in the U.S. throughout the country (CAST, 1985; West- wood, 1986, 1989). National Germplasm System The USDA Agricultural Research Service (ARS) has operated 4 regional P1 stations in Joseph Postman and Kim Hummer Ames, Iowa (established 1947), Pullman, U.S. Department ofAgriculture (USDA)—Agricultural Research Service (ARS), Wash. (established 1947), Geneva, N.Y. National Clonal Germplasm Repository (NCGR), Corvallis, OR 97333-2521 (established 1948), and Griffin, Ga. (estab- lished 1949) to receive, preserve, and distrib- Ed Stover ute seed of vegetable and agronomic crops USDA -ARS, NCGR, Davis, CA 95616 (Fig. I). Several additional repositories were later added for special crops such as cotton, Robert Krueger potatoes, and soybeans. In 1958, the National USDA -ARS, NCGR, Riverside, CA 92507 Seed Storage Laboratory (now the National Center for Genetic Resource Preservation) Philip Forsline opened in Fort Collins, Cob., to ensure the USDA -ARS, Plant Genetic Resource Unit, Geneva, NY 14456 long-term "base, backup" preservation of seeds from each of the "working" or actively L.J. Grauke distributing collections (White et al., 1989). USDA -ARS, NCGR, Somerville, TX 77879 This network of federal genebanks came to be known as the National Plant Germplasrn Francis Zee System (NPGS). Clonal repositories were U.S. Department of Agriculture—Agricultural Research Service, National added to the NPGS as a result of the efforts Clonal Germplasm Repository, Hilo, HI 96720 of ARS pear breeder Howard Brooks, who chaired a visionary group of researchers Tomas Ayala-Silva who forwarded this cause. Dr. Brooks formed USDA -ARS, NCGR, Miami, FL 33158 an ad hoc clonal committee after a workshop at the 1974 meeting of the American Society Brian Irish for Horticultural Science (White etal., 1989). USDA -ARS, Tropical Agriculture Research Station, Mayaguez, PR 00680 Brooks asked each committee member in 1977 to develop a plan for a national re- Additional index words, genetic resources, clonal repository, genetics, conservation, disease pository for crops important in their region of resistance, environmental adaptation the country. He requested that the plans specifically itemize the necessary space for Abstract. The year 2005 marked the 25th anniversary of the establishment of the U.S. land, greenhouses, laboratories, and offices Department of Agriculture (USDA) National Plant Germplasm System (NPGS), re- needed to safeguard the germplasm of the positories devoted to clonally propagated, horticultural fruit and nut crops. During this regional crops (Brooks and Barton, 1977). quarter century, facilities in Hilo, Hawaii; Ma y aguez, PR.; Miami, Fla.; and Riverside, During the next winter, armed with these Calif. were developed to preserve collections of tropical and subtropical fruit and nut plans, Brooks appeared before the U.S. Con- crops; facilities in Brownwood, Texas; Corvallis, Ore.; Davis, Calif. and Geneva, N.Y. gressional Committee on Agriculture in preserve the temperate crops. Each of these facilities now has internationally recognized, Washington, DC, to request federal funding globally diverse collections of genetic resources for their assigned genera. Germplasm of to establish a system of National Clonal unique genotypes are maintained as growing , evaluated for phenot ypic and Repositories. The congressmen approved genotypic traits, documented in a national public germplasm database, and freely the concept and asked Brooks to develop distributed as clonal propaggules to researchers and other germplasm users around the a more specific plan to be presented to them world. Seed collections represent wild populations for some crop relatives. These 8 in the next year. They were very impressed genebanks maintain 30,000 accessions representing 1600 species of fruit and nut crops when he opened his briefcase and pulled out and their wild relatives. The genebanks distribute more than 15,000 accessions annually the detailed plans that his committee had to international researchers. Although originally conceived as working collections for already assembled for 12 repository sites crop improvement, NPGS genebanks have also become invaluable in providing the raw (M.N. Westwood, Nov. 2005, personal com- materials for basic plant genetic research, reservoirs for rare or endangered species or munication). During that very same session vulnerable landraces, archives of historic cultivars, and field classrooms for educating of Congress, a $2 million federal appropria- the public. These collections preserve botanical treasures as well as the American tion was approved to begin building a system horticultural heritage for now and for future generations. of National Clonal Germplasm Repositories. This request was to be a joint effort matched History of U.S. National Fruit and developed and maintained by individual through mutual agreement with State Agri- Nut Repositories plant breeders at universities and were often cultural Experiment Stations at Land Grant lost when scientists retired, changed the focus Universities. The Corvallis, Ore. Repository Before 1980, fruit and nut germplasm of their research, or encountered funding was selected as the first to be constructed collections in the United States were largely shortfalls (Brooks and Barton, 1977). As because of the large collections of pears, a result, collections composed of irreplace- hazelnuts, and small fruits already present Received for publication 3 Mar. 2006. Accepted able material and assembled through years at that site. The very active involvement of for publication 7 Apr. 2006. of effort were routinely lost, producing an Dr. Wilson Foote, Associate Director of the We thank Mel Westwood for his oral history and for reviewing the manuscript. We thank the tech- erratic and insecure network for fruit and nut Oregon State University Agricultural Exper- nical staff at each of the NPGS Clonal Repositories germplasm conservation. That germp!asm iment Station, pomobogy Professor, Dr. Mel- for their dedicated stewardship of these invaluable insecurity lessened in 1980 with the estab- vin Westwood, and hazelnut breeder, Dr. collections. lishment of the first U.S. National Clonal Maxine Thompson, encouraged this choice. To whom correspondence should be addressed; Germplasm Repository in Corvallis (Jahn Dr. Westwood was later appointed as e-mail jpostn1anars-grin.gov . and Westwood, 1982; Westwood, 1982). National Technical Advisor for Clonal

1188 HORTSCIENCE VOL. 41(5) Auousr 2006 Table 1. Fruit and nut collections at the 8 USDA NPGS Clonal Repositories (NPGS, 2006). Pullman 1851, Genus Site Accessions5 Species Corvallis,$ Geneva0 1° Acilnidia CUR 198 27 Actiniclia DAV 68 14 Aegle RIV 1 Ames Aeglopsis RIV I Fort Collins Afraegle RIV I Davis Aleurites MAY 2 Ame/anchier CUR 226 15 Anipelopsis DAV 21 6 A,npe/opsis GEN 5 3 N. A nacardiu,n MAY 1 Riverside Griffin ,4nana.s I-Il 160 10 l3rownwood Annaniocania BRW 11 1 n. 5,5 Anno,ia MAY 16 6 Anno,ia MIA 70 10 Antideona MAY 3 Archontophoenis MIA 3 3 Miami Aronia CUR 7 Artocarpirs HI 39 5 4nijar7n1s MAY 10 7 Mayaguez Antocarpu.s MIA 4 • National Clonal Germplasm Repository Asi,nina CUR 70 7 4 Regional Plant Introduction Station RIV 6 Ateri/wa HI 25 • National Center for Genetic Resource Preservation Aierrhoa MAY 19 1 (National Seed Storage Laboratory) Aierrhoa MIA 38 Bactris HI 15 Fig. 1. Locations of USDA National Plant Germplasni System clonal genebanks for fruit and nut crops Bact,is MAY 3 Bacteis MIA 16 3 Ba/sa,nocit,iis RIV I I Bert ho/etia MAY I in the order that they became established. Gernlplasrn during the construction of the Bisniarckia MIA I clonal repositories and developed an oper- Contact information and crop responsibilities Bixa MAY 1 ations manual to guide new staff in establish- for each site are provided (Table 2), and Blighia MIA 1 ing their facilities (Westwood, 1986). number of accessions and collection locations Brosinnon MIA 1 Funding was ultimately approved for 8 of by genus are itemized (Table 1). Biitea MIA 1 the 12 originally planned clonal repositories But/a MIA II I Camellia HI 18 (Fig. 1). Two repositories, one in Corvallis, Ore., established 1980 Orlando, Florida, and one in Riverside, Cal- Canariienu HI 16 8 NCGR—Corvallis is operated in collabo- 143 1 ifornia, were originally established as a result Ca,ka HI ration with Oregon State University and is Canissa MIA 1 of quarantine restrictions between the 2 states located on the grounds of their Horticultural Carta BRW 1340 23 but were later consolidated at Riverside. In Research Farm at 44.55° N latitude. NCGR Canuota MIA 3 1987, the National Arboretum in Washing- Corvallis maintains germplasm collections of Casinuroa MAY 2 ton, DC, was designated as a clonal reposi- more than 26 genera, including several agro- Casinuiroa MIA 3 1 tory for woody landscape crops. nomic and specialty crops. Actinidia, Cor- Caiendishia CUR 7 4 y/us, Ctdonia, Fragaria, Pines, Ribes, Ceanothus CUR 37 9 NPGS Clonal Repositories Chaenomne/es CUR 13 3 Rubies, Vacciniiern are the main assigned and Clunisohalan its MAY 2 The mission of NPGS genebanks includes fruit and nut genera (Table 2). The Reposi- Chnusophv//um MAY 2 1 the acquisition, documentation, preservation, tory maintains more than 8300 clones and Cinna,nonnenu MAY 5 3 evaluation, enhancement, and distribution of 3300 seedlots. The largest clonal collections Cinna,noniu,n MIA 2 1 plant genetic resources. This conservation are Pines with 1900 and Fragaria with 1300 Citrenuocitrus RIV 1 1 activity improves the quality and production accessions. Because Rubies and Vaccinium Citro/brtienel/a RIV 8 of economic crops important to U.S. and are highly diverse, each having between 600 C/from icrocitrues RIV 1 1 world agriculture (NPGS, 2005a). The U.S. and 800 species worldwide, much variability Citroncirus RIV 43 4 4 clonal genebanks house wild relatives as well for these genera is represented with seedlots. Citropsis RIV Citrus MAY 4 Rubies and 570 Vacciniurn as improved genotypes for their assigned Currently, 970 Citrus RIV 896 86 crops and currently maintain 30,000 acces- seedlots are maintained in addition to C/ausena RIV 6 4 sions representing 1600 species (Table I). For a greater number of clones (Table I). C/v,nenia RIV I I some temperate crops, seed collections are Primary clonal collections of Fragania, Coccothrinax MIA 16 7 used to represent wild species populations in Rubies, and Vacciniurn are maintained as Cocos MAY I addition to the genotypes that must be pre- potted plants in screenhouses for protection Cocos MIA 20 1 served clonally. Many nut crops and tropical from virus vectors. Primary clonal collec- Coffea MAY 1 or subtropical species have recalcitrant seeds tions for Actinidia, Conlus, Cvdonia, Pines, cola MAY 4 Coleara RIV I that do not retain viability when stored so that and Ribes are maintained in field collections Copernicia MIA 9 9 on 26 ha of land available to the Repository clonal living plant conservation is required. Cornus CUR 9 1 The original plan for fruit and nut gene banks east of Corvallis. Much of the Rihes and Con/us CUR 795 22 suggested maintenance and preservation as Vaccinitem collections are duplicated in both Corupha MIA 2 primary objectives followed by research and screenhouse and field. In vitro culture of Crataegomnespi/us CUR 3 evaluation (Brooks and Barton, 1977; West- growing shoots provides backup for approx- Crataegues CUR 18 8 wood, 1989). Preservation remains the pri- imately 10% of the collection with priority Crataeniespilus CUR 1 mary focus of the NPGS fruit and nut given to core accessions, frequently re- Cyosop/u/a MIA 4 1 repositories, which are described subsequently quested accessions, and accessions that may (Continued)

HORI SCIENCE VOL. 41(5) AUGUST 2006 1189 Table 1. (Continued) Fruit and nut collections at Table 1. Continued. Table I. Continued. the 8 USDA NPGS Clonal Repositories Genus Site Accessions Species Genus Site Accessions Species (NPGS, 2006). Manilkara MAY 26 Syzygium HI 3 Genus Site Accessionsy Species Manilkara MIA 12 3 Syzygium MAY 3 1 Cudrania DAV 1 Mentha CUR 533 40 Syzygiuin MIA 5 3 Cvdonia CUR 114 Merrillia RIV Tamarindus MAY 1 Dictvosperma MIA 5 Mespilus CUR 48 3 Tamarindus MIA 35 1 Dimocarpus HI 19 Microcitronella RIV Term/na/ia MAY 6 4 Dimocarpus MAY 2 tyuicrocitrus RIV 19 6 Theobroma MAY 156 Dimocarpus MIA 4 Micromelum RIV 2 Theohro,na MIA 72 Diospyros DAV 103 4 Monstera MAY Thrinax MIA 17 3 Diospyros MAY 3 Morus DAV 58 8 Triphasia RIV I I Diospvros MIA 9 5 Murrava RIV 3 Vaccinium CUR 1579 68 Dovyalis MAY 2 Musa MAY 112 4 Vanilla MAY I Duchesnea CUR 3 Musa MIA 73 3 Vasconcellea HI 24 9 Dario HI 3 3 Naringi RIV Fit/s DAV 2913 49 Durio MAY I Nepheliun HI 38 6 Fit/s GEN 1203 26 Dvpsis MIA II 6 Nepheliuni MIA Washington/a MIA I Dyschoriste MIA 1 Olea DAV 139 4 Wenzelia RIV 1 1 Ehretia MIA 2 Oreoniunnea DAV Zingi her MAY 1 1 Elaeis MIA 16 Oxanthera RIV Ziziphus MIA 4 Elaeocarpus MIA 2 Pachira MAY Site, Accessionsv Species Empetrum CUR 8 Pamburus RIV 2 Total for BRW 1351 24 Eremocitrus RIV I Paramignya RIV 2 Total for CUR 11,682 722 Eriobotiya DAV 29 1 Parmentiera MAY 2 Total for DAV 5664 223 Esen beck/a RIV 1 Parthenocissus DAV 2 Total for GEN 5309 99 Eugenia HI 1 Parthenocissus GEN Total for HI 690 77 Eugenia MAY 2 1 Passifiora HI 34 9 Total for MAY 475 104 Eugenia MIA 2 Peraphvllum CUR 8 Total for MIA 3649 224 Euterpe MIA 8 3 Persea MIA 252 3 Total for RlV 1168 151 Feroniella RIV 1 Phoenix MIA 27 7 Grand Total 29,988 1624 Ficus DAV 196 4 Phoenix RIV 69 BRW, Brownwood, Texas; CUR, Corvallis, Ficus MIA 87 47 Pimenta MIA Oregon; DA y , Davis, California; GEN, Geneva, Flacourtia MAY 3 3 Piper MAY 2 New York; HI, Hilo, Hawaii; MAY, Mayaguez, Flacourtia MIA 1 P/star/a DAV 218 12 Pureto Rico; MIA, Miami, Florida; RIV, Riverside, Fortucitrocirus RIV 3 Pleiospermium RIV 3 California. Fort unella MAY 1 Poncirus RIV 54 Approximately 2700 Corvallis accessions are Fortune/la RIV 11 7 Polentilla CUR 3 seedlots; approximately 1640 Geneva accessions Fragaria CUR 1734 37 Pouteria MAY 27 are seedlots. Fragotentilla CUR 3 1 Pouteria MIA 40 5 Garcinia MAY 16 12 Pritchardia MIA Garcinia MIA 22 9 Prunus DAV 1330 87 Gastrococos MIA I Prunus GEN 106 14 be at risk in the field resulting from lack of cold Gaultheria CUR 42 17 Pseudananas HI hardiness. Nonhardy genotypes and species Gaylussacia 5 CUR 15 Pseudocydonia CUR originating in subtropical or tropical locations Genipa MIA 1 Pseudophoenix MIA 3 are also maintained year-round in a green- Glycosmis RIV 4 4 Psidium HI 44 5 Hevea MIA 44 Psidium MAY 4 3 house. Seed collections are stored at —20 °C. Hicksbeachia HI 2 Pterocarya DAV 11 6 Research at Corvallis supported by base Hippophae CUR 2 Punica DAV 149 funding focuses on improving germplasm 1-Jyophorbe MIA 3 1 Pyracomeles CUR storage and maintenance, including in vitro Hyphaene MIA 1 1 Pyronia CUR 7 culture and cryopreservation and develop- Inga MAY 1 1 Pyrus CUR 2180 37 ment of molecular fingerprints (Bassil et al., Inga MIA 2 Ribes CUR 1324 109 2005; Gupta and Reed, 2006; Lewers et al., .Jacaratia HI 1 Rollinia MAY 2005; Mehlenbacher et al., 2005; Reed, 2001; Jug/ans CUR 70 3 Roystonea MIA 7 4 Reed et al., 2006). Other important research Juglans DAV 425 23 Rubus CUR 2078 192 Lansium MAY I I Rubus MAY 2 activities include virus and disease testing, Lecythis MAY 2 Ruscus CUR identity verification, phenotype evaluation, Leucaena HI 7 Ruta RTV propagation techniques, and strategies for Licania MAY 2 Sabal MIA 23 5 seed regeneration. Limonia RIV 1 Saccharum MIA 2421 18 Litchi HI 63 Salacca MAY Litchi MAY 2 1 Sambucus CUR 183 21 Davis, Calif., established 1981 Litchi MIA 18 Sandoricum MAY NCGR—Davis is operated in coordination Livistona MIA 23 10 Sapindus MAY with the University of California and is Lonicera CUR 53 7 Sauropus HI located at 38.54° N latitude. Fruit and nut Macadamia HI 26 5 Schisandra CUR 7 crop genera which are represented by more Macadamia MAY 1 Sever/nm RIV 11 than 100 clonal accessions include Actinidia, Macadamia MIA I Sibbaldia CUR Diospyros, Ficus, Juglans, Olea, Pistacia, Madhuca MIA 1 Sorbaronia CUR 5 4 Malpighia HI 7 Sorbocotoneaster CUR Prunus, Punica, and Vitis (Table 1). The Malpighia MIA 14 3 Sorbocrataegus CUR Davis Repository currently holds over 5400 Malus GEN 3994 55 Sorbopyrus CUR 9 accessions representing approximately 15 Mammea MAY 1 Sorb us CUR 282 60 genera and 175 species of fruit and nut crops. Mangfera MAY 9 Spondias MIA 3 The two genera Vitis and Prunus account for Mangfera MIA 166 Swing/ca RIV approximately 75% of the collection with 2800 and 1300 accessions, respectively.

1190 H0ItTSCIENcF VoL. 41(5) ALI GUST 2006 Table 2. Curators, latitudes, contact information, and crop responsibilities for the fruit and nut repositories in the USDA National Plant Germplasrn System. Site, latitude, curator Contact info Crop responsibilities Brownwood, Texas USDA-ARS-NCGR Carta (pecan, hickory) 31.71°N 10200FM50 L. J. Grauke Somerville, TX 77879 phone (979) 272-1402 fax (979) 272-1401 e-mail: brwlgars-grin.gov

Corvallis, Oregon USDA-ARS-NCGR Actinidia (hardy kiwifruit) A,ne/anchier (service 44.55° N 33447 Peoria Road berry), Asiniina (paw paw), Corvlus (hazelnut), Kim Hummer, berries and specialty crops Corvallis, OR 97333-2521 Ctdonia (quince), Fragaria (strawberry), Joseph Postman, pome fruits phone (541) 738-4200 Hutnulus (hop), Juglans cinerea (butternut) fax (541) 738-4205 Mentha (mint), !i4espilus (medlar), Pvrus (pear), e-mail: cor:ars-grin.gov Ribes (currant, gooseberry) Ruhus (blackberry, raspberry), Sorbus (mountain ash), Vacciniuni (blueberry, cranberry)

Davis, California USDA-ARS-NCGR ,4ctinidia (kiwifruit), Diospsros (persimmon), 38.55° N One Shield Ave, UCD Erioboirva (loquat), Ficus (fig), Juglans (walnut), Ed Stover Davis. CA 95616-8607 Morus (mulberry), 0/ca (olive), Pistacia phone (530) 752-7009 (pistachio), Prunus (stone fruit, almond), Punica fax (530) 752-9659 (pomegranate), Fit/s (grape) e-mail: [email protected]

Geneva, New York Plant Genetic Resource Unit A/ta/us (apple). Prunus cerasus (tart cherries), 42.88° N Cornell University Exp. Station Fit/s (hardy grape) Phil Forsline 630 W. North Street Geneva, NY 14456-0462 phone (315) 787-2390 fax (315) 787-2339 e-mail: Philip.Forsline/q?ars.usda.gov

Hilo, Hawaii USDA-ARS-NCGR Ananas (pineapple), Artocarpus (breadfruit) 19.57° N P0 Box 4487 Averrhoa (star fruit), Bactris (peach palm). Francis Zee 928 Stainback Hwy. Carica (papaya), Canarium (pilinut), Dimocarpus Hilo, HI 96720 (longan), Litchi (lychee), Macadamia phone (808) 959-5833 (macadamia). Ma/pighia (acerola), Nephelium fax (808) 959-3539 (rambutan, pulasan), Passiflora (passion fruit), e-mail: fzee/dpbarc.ars.usda.gov Psidium (guava)

Miami, Florida USDA-ARS-NCGR .4nnona (eherimoya), Cocos nucifera (coconut), 25.64° N 13601 Old Cutler Rd. Dimocarpus (longan), Diospvros digyna (black Tomas Ayala-Silva and Raymond Schnell, fruits, Miami, FL 33158 sapote), Ficus (fig), Litchi (lychee). Mang(,fra nuts phone (305) 254-3635 (mango), Mani/kara zapota (sapodilla), Musa spp. Alan Meerow, ornamentals fax (305) 969-6410 (banana). Persea (avocado), Pouter/a sapota e-mail: [email protected] (mamey sapote), Saccaruoi (sugarcane), Theohroma cacao (cocoa), Zizvphus (Jujube), subtropical and tropical ornamentals, Tripsacum and related grasses.

Mayaguez. Puerto Rico USDA-ARS-TARS Annona (cherimoya). Ga,cinia. Manilkara zapota 18.20° N 2200 Pedro Albizu Campos (nispero), Musa spp. (plantain, banana), Brian Irish and Ricardo Goenaga Ave. Suite 201 Pouter/a sapota (mamey sapote), Tlieohroma Mayaguez, PR 00680 cacao (cocoa), temperate/tropical bamboo, other phone (787) 831-3435 tropical ornamentals. fax (787) 831-3386 e-mail: [email protected]

Riverside, California USDA-ARS-NCGR Citrus and Citrus relatives, Phoenix (date) 33.96° N 1060 Martin Luther King, Jr. Robert Krueger Blvd. Riverside, CA 92507-5437 phone (951) 827-4399 fax (951) 827-4398 e-mail: rivrk(dars-grin.gov

All collections are maintained with 2 trees mine the distribution of genetic variability of tropical and subtropical crops (Ayala-Silva or vines per accession on a 30-ha field site in within and between taxa, and to assist in et al., 2004). The Tropical Agriculture Re- Winters, Calif. Collections of Vitis and Pru- collection management (Aradhya et al., 2003, search Station (TARS) at Mayaguez is located nus are also maintained as potted plants in 2004, 2005). Characterization of accession at 18.201 N latitude and dates back to 1908. screenhouses for protection from disease phenotypes, to assist clients in selecting With the establishment of NPGS Clonal Re- vectors. Nonhardy genotypes and species material for their research programs, is also positories, TARS was designated as a site originating in subtropical or tropical locations an important and ongoing research activity. for the preservation of a number of tropical are maintained year-round in a greenhouse. fruit and nut crops (Table 2). NPGS clonal Research at Davis supported by base Mayaguez, P.R., established 1984 germplasm activities in Mayaguez were ini- funding focuses primarily on use of molecu- The Repository at Mayaguez is one of 4 tially administered by the curator in Miami, lar markers to fingerprint accessions, deter- NPGS genebanks devoted to the preservation but this site now has its own curator and is no

HoslScIs\ t: Voi . 41(5) Ai,:si 2006 1191 longer considered a substation of the Miami from throughout the range of the species (Forsline et al., 2004). Seed collections are Repository. The Mayaguez genebank main- (provenance collections) (Grauke et al., stored at —20 °C with equal amounts at PGRU tains more than 900 accessions of tropical 1989), and by collections of other hickory and NCGRP. In addition, over 5000 seedlings fruit, nut, and ornamental landscape acces- (Carva) species, to provide the broadest base are grown out and characterized at PGRU sions representing 256 genera and over 400 of genetically compatible germplasm. The 22 with some of these to be selected and main- species. The 450 clonal fruit and nut acces- ha of Repository collections in Brownwood tained as clones. Others are used to produce sions maintained in field plots and green- were supplemented in 1987 by addition of 21 seed backups (Volk et al., 2005). houses include primary collections of ha at a worksite in Burleson County, Texas The tart cherry collection includes ap- Manilkara (sapodilla), Musa, Pouteria (mamey (30.52° N latitude). There are over 400 pecan proximately 100 accessions and represents or mammee sapote), Theohronw, several cultivars in the collection represented by the tetraploid cherries of Prunus cerasus and species of Annona and Garcinia, as well as grafted trees at each worksite. Provenance Prunus fruticosa. These are maintained as temperate and tropical bamboos (Table 1). collections are maintained as self-rooted duplicate trees on MXM 2 rootstock. Over Some accessions are duplicated as backups trees, primarily at the Burleson County work- 50% are also backed up at NCGRP as for the Miami, Hilo, or Riverside Reposito- site. Over 300 provenance accessions are dormant buds in cryogenic storage. ries. Research goals are directed toward represented by up to 8 seedling trees each. The grape collection consists of 1200 maintenance of clonal germplasm collections Small collections of other hickory species accessions, most of which are duplicated in and evaluation of accessions for phenotypic from the United States, Mexico, and Asia the field as own-rooted plants. These acces- and genotypic descriptors. Genotypic charac- (Grauke et al., 1991) are also maintained, sions represent mostly North American na- terization is focused on molecular approaches with some accessions being represented only tive species and hybrids that are cold hardy to determine genetic diversity baselines and by herbarium or nut vouchers. Accessions are and supplement the 2800 cold-tender Vitis clonal fingerprints. Development of in vitro characterized for nut quality, phenology of accessions maintained in Davis, Calif. There tissue culture propagation for some collections growth and flowering, and for traits such as is no cryogenic storage system for grapes, but will improve efficiency of propagation, ease leaf and tree form. Research is focused on the maintaining them as own-rooted vines allows of distribution, and lower the risk of pathogen refinement of descriptors for characterizing for reestablishment in the ease of severe dissemination. genetic diversity in Ca,-va and the develop- winter cold damage. ment of molecular genetic methods for fin- Hilo, Hawaii, established 1986 Miami, Fla., established 1984 gerprinting accessions and characterizing populations (Grauke et al., 2003). A long- The National Clonal Gerniplasm Reposi- The Miami Repository is located at 25.64° tory for Tropical Fruit and Nut Crops was N latitude at the Subtropical Horticultural term goal of this Repository is the designa- tion of appropriate and useful in situ reserves initiated in 1986 as ajoint cooperative project Research Station (SHRS). Approximately between the NPGS and the University of 5000 clonal accessions are maintained at to more adequately represent this important native North American genus. Hawaii at Manoa, College of Tropical Agri- SHRS in greenhouses, lathhouses, and field culture and Human Resources. The Reposi- plots, representing sugar cane, Tripsacum, and tory is located at 19.57° N latitude. The unit subtropical woody ornamentals in addition to Geneva, N.Y., established 1984 The National Clonal Germplasm Reposi- became one of the 4 research units at the U.S. fruit and nut crops. Approximately 81 ha are tory for apple, grape, and tart cherry was Pacific Basin Agricultural Research Center available for research and germplasrn preser- (PBARC) in 1999 and was renamed the vation (Ayala-Silva et al., 2004). Important established in 1984 on the campus of Cornell University in Geneva at 42.88° N latitude. In Tropical Plant Genetic Resource Manage- fruit and nut crops preserved at Miami include ment Unit. Collections include approxi- cacao, fig, lychee, mango, and other tropical 1986, the Repository merged with the North- east Regional Plant Introduction Station mately 1000 accessions maintained as and subtropical fruit species (Table 2). Acces- living plants on 13 ha of field plantings at 3 sions are maintained in field plantings, but (NERPIS) and has been known since then as the Plant Genetic Resources Unit (PGRU). locations with selected germplasm grown in certain new plants must be quarantined and tissue culture and in greenhouses. The unit is tested for pathogens before moving to the field. Currently, around 20,000 accessions are held, representing nearly 200 species and encom- responsible for 14 important tropical fruit and Backup core collections have been established Ananas, Artocar- passing both seed-propagated and vegetatively nut collections, including in Mayaguez or Hilo. Research at Miami is pus, A verrhoa, Bactris, Camellia, Canarium, propagated germplasm. The Ma/us (apple), focused on improved horticultural practices to Carica, Dimocarpus, Litchi, Macadamia, Prunus cerasus (tart cherry), and Vitis (grape) ensure the longevity of collections and on the Malpighia, Nepheliuni, Passiflora, collection comprises 5300 accessions. The and Psi- genetic basis of important horticultural traits dium PGRU also maintains a number of seed- (Table 2). Research at Hilo focuses on such as disease resistance. Molecular markers germplasm maintenance, verification, and have been developed for genetic diversity propagated crops, including onion, tomato, and vegetable brassicas. Research activities storage. Examples of research include in vitro analysis and for the production of molecular storage, molecular fingerprinting, flower in- genetic linkage maps. Families of avocado, focus on problems related to germplasm ac- quisition, collection management, and germ- duction mechanism, fungal and bacterial mango, and jackfruit have been produced that diseases of tropical crops, and new crop should allow the mapping of the genes in- plasm utilization, including application of whole plant phenotyping, cellular, and molec- evaluations (Keith et al., 2006; Matsumoto volved with disease resistance, fruit quality, Brower 2006; Steiger et al., 2003). and yield. A candidate gene approach is also ular markers to assess diversity of germplasm being used to find genes involved with disease collections; development of improved apple Riverside, Calif., established 1987 resistance and for control of flowering (Ayala- rootstocks; and in vitro and cryopreservation The goal of the Riverside Repository is to Silva et al., 2004; Schnell et al., 2001. 2003). of clonal and seed propagules. provide the genetic diversity necessary to im- The Ma/us collection includes 4000 ac- prove the productivity of citrus, date palms, and cessions with 2500 of these maintained as related taxa and to reduce genetic vulnerability Brownwood, Texas, established 1984 trees grafted to EMLA 7 semidwarfing root- of these crops for the future. This genebank is The Brownwood Repository is located at stock. Nearly all accessions are backed up in located at the University of California Riverside 31.71° N latitude and has been the home of cryogenic storage at the National Center for at 33.971 N latitude. Primary objectives are to the USDA ARS Pecan Research program Genetic Resource Preservation, Fort Collins, collect, maintain, evaluate, preserve, and distrib- since 1933, when Louis Romberg began Cob. (NCGRP) as dormant buds; therefore, ute pathogen-tested clonal germplasm of CitrILs, collecting pecan cultivars for use in breeding. only one tree of each accession is maintained 32 related genera, and date palms On the recommendation of an active Crop in the field. This provides a major cost and related Phoenix species. The secondary Germplasm Committee, Rombergs grafted savings by reducing acreage nearly 50%. objectives are to conduct and encourage research cultivar collections have been augmented by The other 1500 accessions are seed lots or on improved methods of maintenance, evalua- collections from native populations of pecan seedling populations from wild collections tion, preservation, and distribution.

1192 HORTSCIENCE VOL. 41(5) AUGUST 2006 The Riverside Repository is somewhat others are using cryogenic storage of either able methods to identify and "fingerprint" unique among the clonal repositories in that apical meristems or dormant buds for long- genotypes. Identification and elimination of it is truly a cooperative program with the term backup (Corvallis, Geneva). However, duplicate genotypes will allow us to make University of California (UC), Riverside. for many tropical and subtropical crops, de- more efficient use of resources. DNA techni- The largest number of Citrus genotypes is velopment of cryogenic or in vitro storage ques have become useful tools for both iden- maintained in a university field collection that protocols is difficult and has not yet been tifying clones and for locating the genes is shared between ARS and UC. However, as implemented. associated with useful traits. Our germplasm a result of the unknown pathogen status of is only as useful as the information that plants in the field, ARS maintains an addi- Distribution Policies and Procedures accompanies it. GRIN provides a place to tional protected collection of pathogen-tested store geographic locality coordinates that trees in screenhouses. Screenhouse trees are Distribution of clonal germplasm is more allow plant origin maps to be generated and the primary source for clonal distributions. In complicated than distribution of seeds. that will associate accessions with climate and addition, the Repository is accredited by Clonal repositories do not generally distrib- environment data. Each Repository, in con- USDA-APHIS and the California Department ute finished rooted plants, but rather distrib- sultation with a team of crop specialists, has of Food & Agriculture for receiving foreign ute scions or cuttings that must be propaguled developed a list of "descriptors" or key traits citrus introductions, which are used by both by the recipient. Shipping of requested ac- that are of interest to plant breeders. Germ- ARS and UC. Date palms are maintained by cessions is therefore dependent on the season plasm users can query GRIN for plants with ARS at a UC facility in Thermal, Calif. when appropriate propagables are available. traits that meet their specialized needs whether Several satellite plantings of cold-sensitive This might be early summer for strawberry as parents for breeding or representatives of genotypes are also maintained at other UC runners or blueberry softwood cuttings or a unique geographic area. field stations. Research supporting the objec- midwinter for grape cuttings or fruit tree tives of the Repository has focused on phyto- scions. Propagation material of subtropical Technical Advisory and Crop pathologic issues, including development of and tropical species is often available Germplasm Committees improved pathogen assays, investigation of throughout the year. Germplasm is freely abnormalities in field trees, and cooperative available from all NPGS Repositories in Brooks and Barton (1977) in their original work with various entities toward establishing small quantities for research purposes. The proposal for national fruit and nut reposito- certification programs (Fang et al.,1997; National Plant Germplasm System distrib- ries proposed Technical Advisory Commit- Krueger & Roose, 2003). Research on molec- utes germplasm to foreign requesters in tees to guide the development and operation ular characterization has permitted the desig- compliance with federal quarantine regula- of each genebank and Crop Advisory Com- nation of a "core" collection. tions and restrictions of the United States and mittees, later renamed "Crop Germplasm the recipient country. Committees" or CGCs to guide curators in the management of specific collections. CGCs are national working groups made up Documentation of Collections—GRIN Pathogen Testing of specialists representing various disciplines The Germplasm Resources Information Clonal repositories seek to distribute and include federal, state, and private in- Network or GRIN is the national public germplasm without disseminating germ- dustry members. These committees help database that provides both genebank per- plasm-borne pathogens, including viruses, curators identify and fill gaps in U.S. collec- sonnel and germplasm users access to pass- viroids, and phytoplasrnas. Although not tions, prioritize traits for evaluation, appraise port, characterization, evaluation, inventory, certified to be free of all known pathogens, research proposals, and help ensure the ac- and distribution data from the national germ- much of the clonal germplasm in NPGS has curacy of data entered to the GRIN. CGC plasm collections. In addition to plant germ- been tested for the presence of common or members also assist in the designation of core plasm, GRIN also supports the National important viruses and virus-like pathogens. subsets (NPGS, 2005c). Fruit and nut crops Animal Germplasm System, the National Various testing methods are used depending represented by CGCs are: Microbial Germplasm Program, and the Na- on information available for each pathogen. • Carva tional Invertebrate Germplasm Program Some pathogens can only be detected by • Citrus and Phoenix (NPGS. 2005b). Each repository is responsi- inoculating indicator plants and observing • Jtiglans ble for entering and maintaining data for symptoms, whereas other pathogens can be • Ma/us collections at their site. Several database query detected by modern laboratory methods • Prunus options are available to the public at http:// (Jelkman, 2004: Martin, 2004: Postman et • Pvrus www.ars-grin.gov/npgs/searchgrin.html. al., 2005; Schnell et al., 1997). Pathogen test o Vitis results are often required before a phytosani- • Small fruits (berries) Maintenance and Backup Strategies tary certificate can be obtained for safe in- • Tropical fruits and nuts ternational germplasm exchange. Repositories must find a balance between Future Challenges maximizing diversity and minimizing the risk of loss. Some crops like apples, pears, Characterization Twenty-five years after their inception, pecans, and walnuts are economically main- the NPGS Clonal Germplasm Repositories tained in long-lived orchard collections. Witt(1985),inhis "BriefBook" on genetic have established valuable national collec- Crops such as citrus, stone fruits, or straw- diversity, quoted from a plant breeding report tions representing world diversity of not only berries are prone to insect-borne viruses and that compared germplasm collections with commercially important fruit and nut culti- therefore must be protected in screenhouses "pharmacies filled with miracle drugs without vars, but also their wild relatives, which may to prevent plants from becoming infected. labels." We need to uncover the genes lurking have genetic traits of unanticipated impor- Maintaining multiple plants of each acces- in our germplasm that will allow us to develop tance. Cohen et al. ( 199 1 ) defined 4 eras in ex sion as in-house backup increases the secu- cultivars able to overcome emerging diseases, situ crop germplasm conservation: rity of the collection but reduces the space adapt to changing climates, improve flavor, 1. A period of exploration and acquisition available for additional accessions. This is enhance nutritional value, and expand pro- from the early 1800s to approximately 1950; particularly true for potted plants maintained duction into new environments. We must not 2. A period of establishing conservation in screenhouses or greenhouses. Some re- only preserve our vanishing agricultural her- facilities from the 1950s to the 1980s; positories are using in vitro culture as a me- itage, but also discover new sources of essen- 3. A period focused on regeneration and dium-term backup strategy for growing tial human nutrients, satisfy consumer desires research from 1980 to the second decade of plants (Corvallis, Hilo, Mayaguez), and for novel fruits and nuts, and implement reli- the 21st century; and

HORTSCIENCE VOL. 41(5) AUGUST 2006 1193 4. A future era of enhanced utilization. Bassil, NV., R. Botta, and S.A. Mehlenbacher. Matsumoto Brower, T.K. 2006. Genes uniquely Workers at NPGS clonal genebanks had 2005. Microsatellite markers in hazelnut: iso- expressed in vegetative and potassium chlorate a late start on this roadmap, but have now lation, characterization and cross-species ampli- induced floral buds of Dimocarpus Longan. entered the era where regeneration and evalu- fication. J. Am. Soc. Hortic. Sci. 130:543-549. Plant Sci. 170:500-5 16. Brooks, H.J. and D.W. Barton. 1977. A plan for ation of collections are priorities. Crop germ- Martin, R.R. 2004. Recommended procedures for national fruit and nut gennplasm repositories. detection of viruses of small fruit crops. Aeta plasm collections were originally conceived as HoriScience 12:298-300. Hortie. 656:199-207. a "hedge against starvation" (Loewer, 1991), CAST, 1985. Plant germplasm preservation and Mehlenbacher, S.A., R.N. Brown, E.R. Nouhra, and but as collections become more complete, they utilization in U.S. agriculture. Council for N.V. Bassil. 2005. A genetic linkage map for become valuable for more than just crop Agricultural Science and Technology. Ames, hazelnut (Corylus avellana L.) based on RAPD improvement (Greene and Moms, 2001). Iowa. Report No. 106., 35 pp. and SSR markers. Genome 49:122-133. Some species in NPGS collections are rare or CBD. 2006. Convention on Biological Diversity. NPGS. 2005a. The National Plant Germplasm endangered in the wild, making these gene- United Nations Environmental Programme. System. Available at: http://www.ars-grin.gov/ banks conservators of plants that may become Quebec, Canada. Available at: hitp://www.bio- npgs/. Accessed November 22, 2005. extinct in nature. Todays plant explorers build div.orgdcfault.shtni!. Accessed April 6, 2006. NPGS. 2005b. The Gemplasm Resources Infor- on knowledge of plant distributions that were Cohen, J.I., J.T. Williams, D.L. Pluckneti. and H. mation Network. Available at: hiip:/www.ars Shards. 1991. Ex situ conservation of plant grin.govaboutgrin.html. Accessed November often established by NPGS collectors. Sec- genetic resources: global development and en- 22. 2005. ondary and tertiary gene pools captured in vironmental concerns. Science 253:866-872. NPGS. 2005c. Crop Germplasm Committees. our genebanks are being used for basic re- Fang. D.Q., M.L. Roosc, R.R. Krueger. and C.T. Available at: http://www.ars-grin.gov/npgs/ search on evolution and genetic relationships. Federici. 1997. Fingerprinting trifoliate orange abouicgc.html. Accessed November 22, 2005. Patent and trademark attorneys consult the germplasm accessions with isozymes, RFLPs, NPGS. 2006. Summary Statistics for GRIN. Avail- GRIN database and NPGS curators as impor- and inter-simple sequence repeat markers. able at: littp://www.ars-gi-iii.gov/iipgs/stats/ . tant references on cultivar names or traits. Theor. AppI. Genet. 95:211-219. Accessed January 12:2006. Artists and photographers visit our collec- FAO. 2006. International Treaty on Plant Genetic Postman, JD., J.S. DeNoma. and B.M. Reed. 2005. tions to document accessions in other ways. Resources for Food and Agriculture. Commis- Detection and elimination of viruses in USDA Future challenges include identifying gaps in sion on Plant Genetic Resources for Food and hop (Humulus lupulus) gemplasm collection. Agriculture. UNFAO Rome. Available at: Acta Hortic. 668:143 - 148. collections and acquiring plants to fill those htip://www.fao.orgAG/cgrfa/itpgr.htm. Ac- Reed, B.M. 2001. Implementing cryogenic storage gaps. Recent international agreements such cessed April 6 2006. of clonally propagated plants. Cryo Letters as the Convention on Biological Diversity Forsline, P.L.H.S. Aldwinckle, E.E. Dickson. J.J. 22:97-104. (CBD. 2006) and the International Treaty on Luby and S.C. Hokanson. 2004. Collection, Reed, B.M., L. Schumacher, D. Dumet, and E.E. Plant Genetic Resources for Food and Agri- Maintenance, Characterization and Utilization Benson. 2006. Evaluation of a modified culture (FAO, 2006) have redefined the pro- of Wild Apples of Central Asia. pp. 1-61. In: encapsulation-dehydration procedure in- tocols for exchange of plant genetic resources J. Janick, P. Forslinc, F. Dickson. R. Way, and corporating high sucrose preireatments for the between countries. National quarantine regu- M. Thompson (eds.). Horticultural Reviews. eryopreservation of Ribes gerrnplasm, In Vitro lations that prevent the movement of insect and vol. 29 Wild Apple and Fruit Trees of Central Cell. Dcv. Biol. Plant 41:431 436. disease pests also impact international germ- Asia. Wiley, New York. Schnell, Ri., D.M. Kuhn, C.M. Ronning. and D. Grauke, L.J., M.J. Iqbal, A.S. Reddy. and T.E. Harkins. 1997. Application of RT-PCR for plasm exchange. Genebank personnel will be Thompson. 2003. Development of microsatel- indexing avocado sunblotch viroid. Plant Dis. challenged to maintain expanding collections lite DNA markers in pecan. J. Am. Soc. Hortic. 81:1023-1026. with declining resources and transition from the Set. 128:374-380. Schnell, Ri., C.T. Olano, R.J. Cambell, and J.S. acquisition phase to the evaluation phase as Grauke, L.J., J.A. Payne, and B.W. Wood. 1989. North Brown. 2001. AFLP analysis of genetic di- collections mature. Eliminating unintended re- American pecans: a provenance study. Annu. Rpt. versity within ajackfruit gemplasm collection. dundancy, filling genetic gaps, and expanding North. Nut Growers Assn. 80:124-131. Scientia Hort. 1661:1-14. phenotype and genotype data in GRIN will Grauke, L.J., B.W. Wood, and J.A. Payne. 1991. Schnell, R.J.. J.S. Brown, C.T. Olano, E.J. Power, continue to enhance the value of the NPGS fruit Genetic resources of Carya in Vietnam and C.A. Krol, D.N. Kuhn, and J.C. Motamayor. and nut collections. Cohens era of enhanced China. Anna. Rpt. N. Nut Growers Assn. 2003. Evaluation of avocado germplasm using utilization (Cohen et al., 1991), when mature 82-80-87. microsatellite markers. J. Am. Soc. Hortic. Sci. Greene. S.L. and J.B. Morris. 2001. Crop Science 128:881-889. genebanks expand their partnerships with 41:886-892. Steiger, D., P. Moore, F.T. Zee, Z. Liu, and R. breeders, researchers, and stakeholders, has Gupta, S. and B.M. Reed. 2006. Cryopreservation Ming. 2003. Genetic relationships of macad- arrived for the NPGS clonal repositories. of shoot tips of blackberry and raspberry by amia cultivars and species revealed by Afip encapsulation-dehydration and vitrification. markers. Euphytica 132:269-277. Literature Cited CryoLetters 27(l):29-42. Volk, G.M., A.A. Reilley, A.D. Henk, C.M. Ri- Aradhya, K.M., G.S. Dangl, B.H. Prins, J.M. Jahn, O.L. and M.N. Westwood. 1982. Mainte- chards. P.L. Forsline, and H.S. Aldwinekle. Boursiquot, A.M. Walker, C.P. Meredith, and nance of clonal plant germplasm. HortScience 2005. Ex situ conservation of vegetatively C.J. Simon. 2003. Genetic structure and differ- 17:122. propagated species: development of a seed- entiation in cultivated grapes, Vitis vinifera. Jelkman, W. 2004. Detection of virus and virus- based core collection of Malus sieversii. J. Am. L. Genet. Res. 81:179-192. like diseases of fruit trees. Acta Hortic. 657: Soc. Hortic. Sci. 130:203-210. Aradhya, KM., C. Weeks, and C.J. Simon. 2004. 575-596. Westwood, M.N. 1982. Pear germplasm of the new Molecular characterization of diversity and Keith, L.M., M. Velasquez, and F.T. Zee. 2006. National Clonal Repository: its evaluation and relationships within and among seven culti- Identification and characterization of Pestalo- uses. Acta Hortic. 124:57-66. vated species of Prunus. Sci. Hortie. (Amster- tiopsis Spp. causing scab disease of guava. Westwood, M.N. 1986. U.S. establishes national dam) 103:131-144. Psidium Guajava L. in Hawaii. Plant Dis. clonal repository system for fruit and nut crops. Aradhya. KM., D. Potter, F. Gao. and C.J. Simon. 90:16-23. Diversity 9:19-21. 2005. Cladistic biogeography of Juglans (Ju- Krueger. R. and M.L. Roose. 2003. Use of molec- Westwood, M.N. 1989. Maintenance and Stor- glandaceae) based on chioroplast DNA inter- ular markers in the management of citrus age: Clonal Germplasm. Plant Breeding Re- genie spacer sequences. pp. 143-170. In: germplasm resources. J. Am. Soc. Hortic. Sci. views, vol. 7. Timber Press, Portland, OR, pp. Motley, T.J., Zerega, and H. Cross (eds). 128:827-837. 111-128. Darwins Harvest-New Approaches to the Lewers, K.S., N.V. Bassil, S.M. Styan, and S.C. White, G.A., H.L. Shands, and G.R. Lovell. 1989. Origin, Evolution, and Conservation of Crops. Hokanson. 2005. Strawberry GenBank-derived History and Operations of the National Columbia University Press, New York. and genomic simple sequence repeat (SSR) Plant Germplasm System. Plant Breeding Ayala-Silva, T., R.J. Schnell, A.M. Meerow, R. markers and their utility with strawberry, Reviews, vol. 7. Timber Press, Portland, OR, Goenaga, and F. Zee. 2004. Current status of blackberry, and red and black raspberry. J. pp. 5-56. the subtropical and tropical germplasm repos- Am. Soc. Hortic. Sci. 130:102-115. Witt, S.C. 1985. Biotechnology and Genetic Di- itories of the National Plant Germplasm Sys- Loewer, P. 1991. A hedge against starvation. versity. California Agricultural Lands Project, tem. Proc. Fla. State Hort. Soc. 117:182-187. American Horticulturist June: 38-44. San Francisco. 145 pp.

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