The Vitis vinifera background of 'Catawba' and 'Concord' was investigated by using SSR analysis: 'Sémillon' was shown to be an ancestor of 'Catawba', while the wild parent remains unknown. 'Concord' was confirmed to be an offspring of 'Catawba' and another unknown wild parent. Since these two important American varieties most likely resulted from random natural crosses and successful selection, the original, wild growing wild donors remain unknown

Cousins, P.

Eibach, R.

Huber, F.

Maul, E.

Röckel, F.

Schwander, F.

Töpfer, R.

English

JKI Open Journal Systems (Julius Kühn-Institut)

Vitis 55, 53–56 (2016)© The author(s).                               This is an Open Access article distributed under the terms of the Creative Commons Attribution Share-Alike License                               (http://creative-commons.org/licenses/by-sa/4.0/).Correspondence to: F. Huber, Julius Kühn-Institute (JKI), Federal Research Centre for Cultivated Plants, Institute for Grapevine Breed-ing Geilweilerhof, 76833 Siebeldingen, Germany. Fax: +49-(0)6345-41-179. E-mail: franziska.huber@jki.bund.deDOI: 10.5073/vitis.2016.55.53-56A view into American grapevine history: Vitis vinifera cv. 'Sémillon' is an ancestor of 'Catawba' and 'Concord'F. Huber1), F. röckel1), F. ScHwander1), e. Maul1), r. eibacH1), P. couSinS2) and r. TöPFer1)1) Julius Kühn-Institute (JKI), Federal Research Centre for Cultivated Plants, Institute for Grapevine Breeding Geilweilerhof, Siebeldingen, Germany2) E. & J. Gallo Winery, Modesto, California, USASummaryThe Vitis vinifera background of 'Catawba' and 'Concord' was investigated by using SSR analysis: 'Sémillon' was shown to be an ancestor of 'Catawba', while the wild parent remains unknown. 'Concord' was confirmed to be an offspring of 'Catawba' and another unknown wild parent. Since these two important Amer-ican varieties most likely resulted from random natural crosses and successful selection, the original, wild grow-ing wild donors remain unknown.K e y  w o r d s :  Vitis aestivalis; Vitis labrusca; genetic fingerprinting; multiplex PCR; microsatellite; SSR; parentage.IntroductionAccording to MunSon (1909) the initial stock of the later called variety 'Catawba' was discovered in a forest near the Catawba River in North Carolina in 1801. Due to its appealingly dark red berries, very juicy pulp and extraordinary aroma composition, it soon developed to the first well-known American cultivar having commercial im-portance (robinSon et al. 2012). Nowadays, 'Catawba' is still widespread around New York State and mainly used for table grape and juice production. In 2006, 'Catawba' was grown on an area of 522 ha in this region (robinSon et al. 2012). The winegrower John Adlum ("Father of American Viticul-ture") introduced 'Catawba' in the District of Columbia in the year 1823 where it became the first prominent variety planted expansively to produce table grapes, juice and wines (Hedrick 1908). Another commercially important American cultivar, introduced after 'Catawba', is 'Concord'. Believing the historical written records, 'Concord' is a descendant of 'Catawba': Ephraim Bull from Massachusetts dug up a wild V. labrusca next to his fence and planted it on his lot next to other grapevine cultivars, including 'Catawba' (Tukey 1966). E. Bull planted the seed from this V. labrusca accession in 1843 and 'Concord' was selected from among the seedlings. Some characteristics such as the hermaphrodite flowers of 'Concord' and the oval berries in some 'Concord' offspring gave a hint about its V. vinifera portion (Tukey 1966). Both 'Catawba' and 'Concord' wines feature the typical, strong wild foxy flavor (nelSon et al. 1977, raPP et al. 1980, raPP et al. 1993). While the historical origin of 'Catawba' can be retraced rather easily, the actual parents were discussed extensively in the last two centuries (Prince 1830, Tukey 1966, GaleT 2000, Pinney 2007). In any case, the close relatedness to a wild species and the fact that 'Catawba' arose by chance can be stated (robinSon et al. 2012). Two main origin scenarios were discussed on the basis of the ampelography: 'Catawba' to be (1) a true wild V. labrusca variety or (2) an interspecific cross of the wild grapes V. aestivalis or V. labrusca with an unknown V. vinifera cultivar introduced to North America at this time (Hedrick 1908). V. labrusca is a native wild grapevine species in North America and was firstly described by linné in 1763 (aMbroSi 2011). Microsatellite analysis of the genetic resources at the JKI Geilweilerhof resulted in first indications that the European cultivar 'Sémillon' could be the V. vinifera parent.Material and Methods'Sémillon' (GaleT 2000, lacoMbe et al. 2013), 'Catawba' and 'Concord' (Hedrick 1908) were confirmed as ampelo-graphically true to type accessions within the grapevine collection at the Julius Kühn-Institut (JKI), Institute for Grapevine Breeding Geilweilerhof. Analyzing the finger-prints of 600 genotypes with the statistical software FaMoz (Gerber et al. 2003), the one from 'Sémillon' was the only one with a good match with 'Catawba'. Young leaf material from these accessions was lyophilized (Lyocube, Christ, Germany) and DNA was extracted with the aid of a kit (Nu-cleoSpin® 96 Plant II, Macherey-Nagel, Germany). DNA of reference accessions from the germplasm collection in Geneva were kindly provided by T. Chao (U. S. Department of Agriculture, USDA). The analysis didn't include any V. labrusca or V. aestivalis accession.The multiplex PCR was conducted with the KAPA2G Fast Multiplex PCR Kit (KAPABIOSYSTENS, USA) com-prising up to 10 primer pairs with fluorescent labels (forward primer coupled with HEX, ROX/PET, TAMRA or FAM). PCR program: 95 °C for 3 min (initial denaturation), 95 °C for 15 s (denaturation), 60 or 58 °C for 30 s (primer anneal-ing), 72 °C for 30 or 50 s (elongation) and 72 °C for 3 min  54 F. Huber et al.(final elongation) with 30 cycles of denaturation, annealing and elongation. The 137 oligonucleotides used were VMC from the Vitis Microsatellite Consortium in France, UDV (di GaSPero et al. 2005), VVI (MerdinoGlu et al. 2005)VVMD (bowerS et al. 1999), VCHR (ciPriani et al. 2008), VrZAG (SeFc et al. 1999), in-house GF (ZHanG et al. 2009, FecHTer et al. 2012, Schwander et al. 2012, rex et al. 2014) and un-published GF from JKI Geilweilerhof. The fragment length analysis was done on a 3130xl Genetic Analyzer (Applied Biosystems, Germany) and the corresponding Genemapper 4.0. software. To confirm the berry color locus alleles, 'Sé-millon', 'Catawba' and 'Concord' were investigated by the in-house SSR marker GF02-55 for the berry color locus on chromosome 2 (aZuMa et al. 2011). The GF02-55 forward primer is AAAATTGAAGGACAGGAGGAGG, reverse primer is GCAAGGCTGGTCTACTCAGAAA.Results and DiscussionSystematic SSR analysis of genotypes of the grape-vine repository provided first evidence of a parent-child relation of 'Catawba' and 'Sémillon'. As a follow-up study a detailed genotyping included the putative parent of 'Ca-tawba' ('Sémillon') and the presumed offspring 'Concord'. The 'Catawba' and 'Concord' accessions of JKI Geilweiler-hof showed the identical genetic fingerprint to the USDA references, confirming their identity (lacoMbe et al. 2013, VIVC Vitis International Variety Catalogue). Tab. 1 shows 38 SSR markers as an example of in total 166 SSR mark-ers, reasonably equally distributed throughout the genome which were applied in a mapping study. In larger parentage analysis studies was shown that 20 SSRs can be sufficient to confirm parent-child relationships (lacoMbe et al. 2013). T a b l e  1 Subset of SSR markers informative for the relationship of 'Sémillon', 'Catawba' and 'Concord', on the 19 grape chromosomes (Chr). Fragment lengths in [bp] are given for the two alleles of each cultivar. Consistent fragment lengths for 'Sémillon' and 'Catawba' are bold. Same fragment lengths for 'Catawba' and 'Concord' are in italicChr SSR Marker Sémillon alleles [bp] Catawba alleles [bp] Concord alleles [bp]1 UDV-035 148 162 136 162 132 136VMC9D3 199 205 205 217 205 2212 GF02-11 276 276 276 281 276 283GF02-17 327 327 327 333 327 3313 VMC2E7 156 160 154 160 154 160GF03-01 118 122 108 122 108 1224 UDV-034 178 180 178 202 200 202GF04-16 316 316 286 316 280 3165 VRZAG79 247 251 247 251 247 259GF05-11 304 304 295 304 295 2956 GF06-16 257 265 239 265 239 241GF06-08 140 162 150 162 140 1627 GF07-05 157 157 157 163 163 163GF07-14 209 221 211 221 209 2118 VCHR08A 200 200 200 0 163 0UDV-026 127 133 127 0 157 09 GF09-14 356 356 356 370 370 370VMC6E4 141 141 121 141 121 12110 VVIH01 244 250 240 244 240 250GF10-09 298 298 284 298 284 28411 GF11-03 122 132 122 130 122 130UDV-028 140 146 140 150 140 14612 GF12-05 167 171 163 171 163 169VMC4f3 164 170 164 204 168 20413 VCHR13A 139 149 139 139 134 139GF13-08 348 358 348 350 350 35814 GF14-02 221 227 221 245 221 245VMC6C10 110 138 122 138 126 13815 GF15-06 166 178 170 178 170 170GF15-02 132 140 140 144 142 14416 GF16-31 226 230 220 230 228 230GF16-25 320 320 315 320 312 32017 VCHR17A 184 184 156 184 164 184GF17-07 109 109 90 109 90 10918 SCU10 205 211 205 211 205 214VMC8F4.2 94 108 94 106 94 10219 UDV-023 179 201 201 227 223 227GF19-10 153 155 155 159 159 159 Vitis vinifera cv. 'Sémillon' is an ancestor of 'Catawba' and 'Concord' 55T a b l e  2 Results of the analysis using the SSR marker GF02-55 with the product lengths of the cultivars 'Sémillon', 'Catawba' and 'Concord'. This marker is used for investigation of the berry color locus of grapes located on chromosome 2            GF02-55 productsCultivar name        Allele # 1 [bp]Allele # 2 [bp] Genotype Berry colorSémillon 215 217 white/white WhiteCatawba 168 215 red/white RedConcord 172 215 black/white Black137 markers proofed to be heterozygous for 'Catawba'. For these markers, 'Catawba' shared one allele with 'Sémillon' in any case. 'Concord' inherited always one 'Catawba' allele, either the 'Sémillon' allele or the allele of the wild ancestor confirming their parent-child relatedness (Tab. 1). A consideration about the Mendelian heredity of the berry color supports our result on a different level: In grapes, the black berry color is dominant over red and the red color is dominant over white (barriTT and einSeT 1969). The putative 'Catawba' ancestor 'Sémillon' is a white berry cul-tivar. Consequently it carries two recessive alleles for white color at the berry color locus. As 'Catawba' has red grapes, it could have either two alleles for red color or one white allele plus one dominant red allele. To support 'Sémillon' as 'Catawba's ancestor, the offspring must have inherited the white allele and thus being heterozygous at the berry color locus. The JKI Geilweilerhof in Siebeldingen maintains a population, which derived from a cross of 'Blaufraenkisch' and 'Catawba'. The black berried 'Blaufraenkisch' has the allele combinations black/white receiving the white allele from its white berried ancestor 'Heunisch Weiss' (lacoMbe et al. 2013). The 'Blaufraenkisch' x 'Catawba' population segregates concerning the berry color in the following way: in year 2014 out of 112 descendants, 59 were black, 8 red and 25 white. According to this, the ratio of the berry color (black : red : white) was approximately 2 : 1 : 1. As a conclusion, 'Catawba' must be heterozygous at the berry color locus carrying the allele combination red/white which is in accordance with the given 'Sémillon' descent. In case of 'Concord' having black colored berries, it could have the alleles black/white, black/red or black/black. The identified parent-child relationship between 'Catawba' and 'Concord' is also in agreement with the results that 'Concord' has inherited either its recessive red or its recessive white allele besides the black one. The SSR analysis indicated one black allele for 'Concord' and one white allele of 'Catawba', originated from 'Sémillon' (Tab. 2, Figure).To deduce something more about the ancestry of 'Concord' regarding the wild type portion, the SSR marker information around the flower sex locus was used. This is, like the berry color locus, located on chromosome 2 (FecHTer et al. 2012), thus the heredity of the alleles for hermaph-rodite, male and female flowers can be traced. 'Sémillon', 'Catawba' and 'Concord' exhibit hermaphrodite flower sex having one allele for hermaphrodity and one for female flowers. According to FecHTer et al. (2012), 'Sémillon' and 'Concord' differ in their female allele: 'Concord' has the “wild female allele” (HFw) in contrast to 'Sémillon' having the allele typical for elite European cultivars (HFk). 'Catawba' was analyzed with the diagnostic InDel marker for the ade-nine-phosphoribosyl transferase (APT) and was found out also to have a recessive "wild female allele" (HFw) found in wild species. According to this, 'Catawba' inherited the female flower sex allele from the wild parent and the her-maphrodite allele from 'Sémillon' probably being more yield stable than female plants. 'Concord' inherited the 'Sémillon' part of chromosome 2 completely, as indicated in Tab. 1. This was verified by seven SSR markers equally distributed over chromosome 2 for all three cultivars (data not shown). According to this, 'Concord' inherited the hermaphrodite flower sex allele (H) from 'Sémillon' through 'Catawba' and the "wild Fw allele" came from the wild parent. Wild Vitis species are usually dioecious (Tukey 1966) and the male allele is dominant over the female allele (FecHTer et al. 2012). If the other ancestor of 'Concord' would have been a pure wild species, it would show the female flower sex. Since 'Concord' is a product of coincidence, the wild parent is unknown. But a principal components analysis for an ances-try study from Sawler et al. (2013) issued a wild portion of 49 % for 'Catawba' and 69 % in 'Concord'. Considering this Figure: The Pedigree of 'Concord' could be retraced by using phenotypic data combined with genotypic data received from SSR markers (berry color locus) and an APT InDel marker (flower sex locus) as well as former genome-wide genetic analysis from Sawler et al. (2013) concerning the wild portions (dark grey) and V. vinifera portions (light grey). B = black berry color allele, R = red berry color allele, w = white berry color allele. H = hermaphrodite allele, Fk = female allele, Fw = "wild female allele". ♀ = female flowers, ♂ = hermaphrodite flowers.                  + 56 F. Huber et al.information combined with our results, it can be deduced that the unknown ancestors were two different, pure wild species that inherited their "wild female alleles" respectively. The summary of the results are depicted in the Figure.Besides the validation of the genetic background, the marker information reveals genomic regions with wild species content in the 'Catawba' offspring. The accordingly informative SSR markers can be used for background se-lection in future backcross breeding programs as proposed by HerZoG et al. (2013).AcknowledgementSpecial thanks to T. cHao from USDA in Geneva for providing DNA of the reference accessions as well as for deep discussions. Also special thanks to FDW and BLE for funding. ReferencesaMbroSi, H.; 2011: Farbatlas Rebsorten. 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A view into American grapevine history: Vitis vinifera cv. 'Sémillon' is an ancestor of 'Catawba' and 'Concord'

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A view into American grapevine history: Vitis vinifera cv. 'Sémillon' is an ancestor of 'Catawba' and 'Concord'

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