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Malus Sieversii: a Diverse Central Asian Apple Species in the USDA

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Malus Sieversii: a Diverse Central Asian Apple Species in the USDA HORTSCIENCE 48(12):1440–1444. 2013. M. dasyphylla, and M. kirghisorum. In the Genetic Resources Information Network (GRIN; USDA, ARS, National Genetic Re- Malus sieversii: A Diverse Central sources Program, 2013), these are synony- mous with M. pumila var. niedzwetzkyana, Asian Apple Species in the USDA-ARS M. pumila, and M. sieversii var. kirghisorum, respectively. Dzhangaliev (2003) summa- National Plant Germplasm System rized the Malus species native to Kazakhstan. M. sieversii and M. niedzwetazkyana are Gayle M. Volk2, Adam D. Henk, and Christopher M. Richards most similar to cultivated apple with USDA-ARS National Center for Genetic Resources Preservation, 1111 South M. niedzwetzkyana as a botanical variety of Mason Street, Fort Collins, CO 80521 M. dasyphylla. Furthermore, M. kirghisorum is described as distinct from M. sieversii Philip L. Forsline because it grows adjacent to relic Juglans USDA-ARS Plant Genetic Resources Unit, Geneva, NY 14456 regia forests as part of a mesophilous Turgai forest flora with a limited range in C. Thomas Chao1 Dzhungareskei and Zailijskei Alatau USDA-ARS National Center for Genetic Resources Preservation, 1111 South (Dzhangaliev, 2003). Based on molecular and phenotypic evi- Mason Street, Fort Collins, CO 80521 dence, M. sieversii is likely a progenitor species Additional index words. genepool, genetic diversity, Malus pumila, subspecies, taxonomy of the domesticated apple, Malus ·domestica (Harris et al., 2002; Velasco et al., 2010; Abstract. There are several Central Asian Malus species and varieties in the USDA-ARS Zhou and Li, 2000). In fact, M. sieversii and National Plant Germplasm System (NPGS) apple collection. Malus sieversii is the most M. ·domestica have sometimes been coclas- comprehensively collected species native to Central Asia. Other taxa such as M. sieversii sified as M. pumila (Harris et al., 2002; var. kirghisorum, M. sieversii var. turkmenorum, M. pumila, and M. pumila var. Juniper and Mabberley, 2006; Phipps et al., niedzwetzkyana have primarily been donated to the collection by other institutions and 1990, 1991). Although there are data suggest- arboreta. We sought to determine if genetic and/or phenotypic differences among the ing similarities between these species, ge- individuals that make up the gene pools of these taxa in the NPGS exhibit unique netic evidence suggests that they can be characteristics. Genetic data, based on microsatellite analyses, suggested that the genetically distinguished based on microsa- diversity within each taxa is significantly greater than that among taxa. Trait data also tellite markers (Gross et al., 2012). In fact, revealed very few differences among taxa, the primary characteristic being the dark red putative hybrids between M. sieversii and M. fruit coloration and tinted flesh color of the accessions assigned to M. pumila var. ·domestica can also be identified, thus fur- niedzwetzkyana resulting from a known single-gene mutation in anthocyanin production. ther supporting the differentiation of these We found that M. sieversii is a highly diverse species with a range in genetic and species (Gross et al., 2012). Other species phenotypic trait variation that includes the characteristics of the other Central Asian such as M. orientalis, M. sylvestris, and taxa of interest. We conclude that the gene pools that comprise the accessions within the M. prunifolia likely played a role in Malus NPGS Central Asian Malus collection are highly overlapping with respect to both domestication (Coart et al., 2006; Cornille phenotypic traits and genotypic characters. et al., 2012; Forte et al., 2002; Gross et al., 2012; Luby, 2003; Robinson et al., 2001; Velasco et al., 2010). Single nucleotide poly- The mountains of Central Asia have forest drought tolerance, and disease resistance morphism transferability was 26% from species that include Malus sieversii (Ledeb.) (Dzhangaliev, 2003; Yan et al., 2008; Zhou, M. ·domestica to either M. sieversii or M. Roem., a highly diverse apple crop wild 1999). M. sylvestris, supporting the evidence that relative. The Russian scientist Vavilov ex- The U.S. Department of Agriculture there are significant genomic differences plored the forests of Central Asia in the 1920s (USDA) sponsored four plant exploration among M. ·domestica and its likely pro- and made note of the wide range of M. sieversii trips in Central Asia to collect wild apples genitor species (Micheletti et al., 2011). phenotypes, suggesting that this region, includ- between 1989 and 1996. Participants on those Previously, we assessed the diversity of ing Kazakhstan, Kyrgyzstan, Tajikistan, and trips sought clones with unusual phenotypes the M. sieversii collection in the USDA-ARS western China, is a center of origin for the and collected seeds from trees growing in NPGS using microsatellite markers. Collec- domesticated apple (Luby et al., 2001; Wan diverse habitats, primarily in Kazakhstan. tion sites were significantly differentiated et al., 2011; Yan et al., 2008). Malus sieversii Over 1000 seedlings from the wild-collected and had unique alleles; however, differenti- grows in a wide range of habitats, including seeds have been grown in Geneva, NY, in the ation within individual families was more areas with hot summers and short winters as USDA-ARS NPGS apple collection. Most of than three times the level of that among sites. well as those with long, severe winters in the the seedlots were assigned to the M. sieversii Differentiation was congruent with geo- Tien Shan Mountains (Dzhangaliev, 2003). species, although some have been labeled as graphical location with southwestern collec- Wild populations are found in montane, scrub, Malus sieversii var. kirghisorum (Al. Fed. & tion sites being more admixed and more humid as well as dry continental forests and Fed.) Ponomar. The repository also has a diverse than the northern sites (Richards in diverse stream habitats in xeric areas number of accessions that have been classi- et al., 2009b). These data supported results (Forsline et al., 2003). Malus sieversii trees fied as Central Asian taxa, namely Malus published previously that used isozymes to are phenotypically diverse, offering variation sieversii var. turkmenorum (Juz. & Popov) determine within- and among-population dif- in the time of flowering, quantity and quality Ponomar. and Malus pumila Mill. Malus ferences in M. sieversii collections (Lamboy of fruit, ripening period, fruit biochemical pumila var. niedzwetzkyana Dieck. was for- et al., 1996). composition, tree architecture, winter and merly listed as a variety but in 2010 was Core collections were also proposed using reclassified as M. pumila.Manyofthese available microsatellite and quantitative trait accessions were donated to the repository from data for the M. sieversii trees from Kazakh- Received for publication 18 June 2013. Accepted other gene banks or arboreta. In addition to stan in the NPGS. Three sets of 35 trees, for publication 17 Sept. 2013. these species/varieties, the literature describes representative of Site 6, Site 9, and ‘‘other’’ 1Retired. a number of variations in nomenclature for collection sites, were identified as core col- 2To whom reprint requests should be addressed; Central Asian Malus species, subspecies, lections that capture greater than 95% of the e-mail [email protected]. and varieties including: M. niedzwetzkyana, measured allelic and phenotypic diversity 1440 HORTSCIENCE VOL. 48(12) DECEMBER 2013 (Forsline et al., 2003; Richards et al., 2009a; volatile components (Bai et al., 2012; Fazio 1990s (Forsline et al., 2003; Luby et al., Volk et al., 2005). et al., 2009; Forsline and Aldwinckle, 2004; 2001). Seeds were germinated and planted Malus sieversii accessions in the NPGS Forsline et al., 2003; Luby et al., 2001, 2002). as own-root seedlings within an orchard set- have been evaluated for a number of fruit, Some traits even differed by collection site ting. Clones representing individuals of the biotic, and abiotic stress resistance traits both such as the later harvest time and an increased other Central Asian taxa were provided to the on-site in Geneva, NY, and by collaborators. level of fireblight resistance in Site 6 and NPGS by donors and are maintained as Physiological and phenological traits such as larger, red fruit more common in Site 9 grafted trees within the collection (Table 1). high chilling requirement, early fruit matu- (Forsline and Aldwinckle, 2004). A total of 99 M. sieversii, nine M. sieversii rity, short juvenility, dwarf growth habit, late The large M. sieversii collection in the var. kirghisorum,threeM. sieversii var. flowering, stooling ability, and vigor were of NPGS provided us with an opportunity to turkmenorum,eightM. pumila,andthree particular interest to breeding programs assess the relationship among accessions M. pumila var. niedzwetzkyana trees were (Bassett et al., 2011; Fazio et al., 2009; Forsline collected and labeled as M. sieversii to those available for inclusion in this research. and Aldwinckle, 2004; Forsline et al., 2003; that have been classified as related species/ Genetic comparisons among accessions Luby et al., 2001, 2002). In addition, sources varieties: M. sieversii var. kirghisorum, assigned to the five Malus species/varieties of disease and insect resistance have been M. sieversii var. turkmenorum, M. pumila, were performed using microsatellite markers. identified for apple scab (Venturia inaequalis), and M. pumila var. niedzwetzkyana. We use DNA extraction, polymerase chain reaction fireblight (Erwinia
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