Reconstructing the Heritages of 'Gruè Ner Veltliner' and 'Sauvignon Blanc' from Crossings with 'Traminer' by SSR Analyses

Mitteilungen Klosterneuburg 59 (2009): 199-208

Reconstructing the heritages of 'GruÈ ner Veltliner' and 'Sauvignon blanc' from crossings with 'Traminer' by SSR analyses

FERDINAND REGNER and ROBERT HACK

Lehr- und Forschungszentrum Wein- und Obstbau A-3400 Klosterneuburg, Wiener Straûe 74 E-mail: [email protected]

'GruÈ ner Veltliner' is the main variety in Austrian viticulture. About 30 % of the national production is obtained from this grapevine. Due to the importance of the variety for the Austrian production it is not satisfying to define its origin only as traditional and unknown. By means of more than 120 SSR markers it was feasible to confirm the parentage of the variety 'Traminer'. Fortuitously we could recognize the second parent of 'GruÈ ner Veltliner'. But this grapevine survived as a single plant without a name and still without reference vine therefore still nameless. Concerning the phenotype this unnamed variety seems to be more similar to 'GruÈ ner Veltliner' than to 'Traminer', the other parent. Huge worldwide importance is attributed to the variety 'Sauvignon blanc'. In France more than 25000 ha are planted with 'Sauvignon', in oversea countries like New Zealand, Australia and others acreages are increasing. The origin of the variety was not clear and even the heritage was unknown. Numerous SSR analyses allo- wed to recognize the parentage as a cross of 'Traminer' x 'Chenin blanc'. So 'Traminer' could be determined once more as parental vine of a high quality grapevine variety. 'Traminer' seems to be one of the key varieties for the development of the Mid European diversity of grapevine. Keywords: heritage, clones of grapevine, genetic analysis, SSR markers, origin, outcrossing

Ermittlung der Abstammung von 'GruÈ ner Veltliner' und 'Sauvignon blanc' als Kreuzungen von 'Traminer' mit SSR-Analysen. 'GruÈ ner Veltliner' ist die wichtigste Sorte des oÈsterreichischen Weinbaus. UÈ ber 30 % der heimi- schen Produktion stammen von dieser Sorte. Bei dieser uÈ berragenden Stellung der Sorte ist als Herkunftsangabe die Verwendung als traditionelle Rebsorte und deren unbekannte Herkunft nicht zufriedenstellend. Mittels Mikrosatel- litenanalyse wurden uÈ ber 120 Genorte untersucht, um die Beziehung der Rebsorte zur vermuteten Herkunft aus einer Traminer-Kreuzung zu uÈ berpruÈ fen. Durch Zufall konnte mittlerweile eine zweite Rebe als Elternsorte identifiziert werden. Diese Rebe konnte bisher keiner bekannten Sorte zugeordnet werden, was eine genaue Bezeichnung mit einem Sortennamen verhindert hat. Ampelographische Untersuchungen zeigten, dass diese Rebe phaÈnotypisch der Sorte 'GruÈ ner Veltliner' aÈhnlicher ist als die andere Elternsorte 'Traminer'. Eine sehr groûe Bedeutung hat fuÈr Frankreich die Sorte 'Sauvignon blanc'. Es sind zur Zeit ca. 25.000 ha mit der Rebe alleine in Frankreich bestockt. Insbesondere Neuseeland, Australien und anderen UÈ berseelaÈndern ist die Tendenz steigend. Die Herkunft und Ab- stammung der Sorte lag ebenfalls im Dunkeln und konnte nunmehr durch zahlreiche Analysen mittels SSR-Mar- kern als eine Kreuzung zwischen 'Traminer' und 'Chenin blanc' identifiziert werden. Damit erweist sich einmal mehr die Sorte 'Traminer' als eine SchluÈ sselsorte zur Entwicklung der heutigen Rebsorten. SchlagwoÈ rter: Abstammung, Rebklone, Genetische Analyse, SSR-Marker, Herkunft, Auskreuzung

DeÂtermination de l'origine du 'GruÈ ner Veltliner' et du 'Sauvignon blanc' en tant que croisements de 'Traminer' aÁ l'aide d'analyses SSR. 'GruÈ ner Veltliner' est le ceÂpage le plus important de la viticulture autrichienne. Plus de 30 % de la production nationale proviennent de ce ceÂpage. Vu le positionnement exceptionnel de ce ceÂpage, il n'est pas satisfaisant que la deÂclaration d'origine mentionne l'utilisation en tant que ceÂpage traditionnel et son origine incon- nue. Plus de 120 loci ont eÂteÂ examineÂs par voie d'analyse des microsatellites afin de veÂrifier si le ceÂpage est reÂellement

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issu d'un croisement du Traminer, son origine preÂsumeÂe. C'est par hasard qu'une deuxieÁme vigne a pu eÃtre identifieÂe comme ceÂpage parent entre-temps. Jusqu'aÁ preÂsent, cette vigne n'a pu eÃtre attribueÂe aÁ aucun ceÂpage connu, ce qui a empeÃcheÂsadeÂsignation exacte par un nom de varieÂteÂ. Les essais d'ampeÂlographie ont montreÂ que, du point de vue du pheÂnotype, cette vigne ressemble plus au ceÂpage 'GruÈ ner Veltliner' que l'autre ceÂpage parent, le 'Traminer'. Pour la France, le ceÂpage 'Sauvignon blanc' est d'une treÁs grande importance. AÁ l'heure actuelle, rien qu'en France, cette vigne est cultiveÂe sur preÁs de 25 000 ha. La tendance est aÁ la hausse, notamment en Nouvelle-ZeÂlande, en Au- stralie et d'autres pays d'outre-mer. L'origine et l'ascendance du ceÂpage eÂtaient eÂgalement incertaines et le ceÂpage a pu eÃtre identifieÂreÂcemment aÁ l'aide de nombreuses analyses au moyen de marqueurs SSR comme un croisement entre le 'Traminer' et le 'Chenin blanc'. Par laÁ, le ceÂpage 'Traminer' se reÂveÁle de nouveau eÃtre un ceÂpage-cleÂ dans le deÂ- veloppement des ceÂpages modernes. Mots cleÂs: Ascendance, clones de vignes, analyse geÂneÂtique, marqueurs SSR, origine, croisement

The grapevine variety 'GruÈ ner Veltliner' contributes could be the wrong definition of this variety as a about 30 % to the national grape production of Austria. green-berried type of the variety 'Roter Veltliner' in Therefore this variety is the most important one for the 19th century. There are also white- and brown-be- Austrian winegrowers (AMBROSI et al., 1994). In some rried types of 'Roter Veltliner'. Especially the brown- of the viticultural areas like Kremstal and Kamptal 75 berried type was widespread and in former times one % of the total area are planted with 'GruÈ ner Veltliner'. of the most important grapes for the local viticulture. (www.weinausoesterreich.at). Besides the 16700 ha in In some regions growers call the variety ªOÈ sterreicherª Austria it should be mentioned that there are about as it was very popular in Austria. The grapevines 'Brau- 4000 ha in Slovakia, 2000 ha in Hungary and 1800 in ner Veltliner' and 'GruÈ ner Veltliner' look very similar the Czech Republic as well as small areas in Italy (REG- under certain circumstances. That was the reason for NER, 2007). These data show that 'GruÈ ner Veltliner' has mixing them up without noticing the mistake. Never- to be accounted one of the important varieties of Eu- theless the variety 'Roter Veltliner' is the original one rope. Cultivation of 'GruÈ ner Veltliner' was even started and is placed at the centre of this family. When in books at Umpqua Valley in the U.S. (www.wine.appellationa- before the 19th century the name 'Veltliner' was men- merica.com). tioned, it always meant 'Roter Veltliner'. BABO and The variety is known as traditional with unknown ori- MACH (1881) declared that the misnaming was already gin. It was confirmed that from the botanical point of spread so far that they didn't want to change anything view the synonym 'Weiûgipfler' would better describe despite their better knowledge. morphologic characteristics of the vine. In former stu- About the heritage no knowledge was available beside dies it could be recognized that the variability within the wrong parentage of 'Roter Veltliner'. We have suffi- the 'GruÈ ner Veltliner' variety is not as high as estimated cient information about the area where 'GruÈ ner Veltli- th before (REGNER et al., 2008). ner' was cultivated in the 19 century. The variety The name 'GruÈ ner Veltliner' is rather young and didn't sometimes was planted despite the ban by the local exist at the beginning of the 19th century. Former de- nobles. signations of this variety were 'Weiûgipfler' or 'GruÈn- BURGER (1837) was convinced that the centre of the cul- muskateller'. BABO and MACH (1881) defined the variety tivation of this variety (he called it Plinia austriaca) was as 'GruÈ ner Veltliner' for the first time. But it seems that located north of the Danube in the area around Retz, the designation was of sufficient quality to recognize especially within the villages Pulkau, Zellerndorf, the variety and to confirm that the former names were Haugsdorf and Stinkenbrunn but also along the roads true synonyms. From the botanical point of view 'GruÈ- to BruÈ nn and Horn. There the vineyards were planted ner Veltliner' is not a genuine Veltliner variety and the with only one variety although it was usual at that better denomination would be the synonym 'Weiûgipf- time to mix different varieties in the same vineyard. It ler'. is amazing that BURGER (1837) included the variety in GOETHE (1887) favoured 'Weiûgipfler' as the correct one family with 'Rotgipfler' and 'Rheinriesling'. Both name for this variety. Comparing it with 'Rotgipfler', varieties show according to today's knowledge a paren- both varieties seem to be closely related as both show tage of 'Traminer'. 'Traminer' as a parental plant. But why is this variety SCHAMS (1832), who reported about the guidelines for called 'Veltliner'? The reasons remain unclear. One viticulture in the city of Pressburg in 1804, mentions

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an appeal of the mayor of the town to avoid cultivation the viticultural regions of the Jura, Loire and Alsace of 'GruÈ nmuskateller' (synonym of 'GruÈ ner Veltliner') are candidates for the origin, but not Bordeaux or as it was considered to be a mass producer with low Sauternes. wine quality. He also mentioned that the name 'Muska- For the Austrian production it can be confirmed that teller' has nothing to do with true 'Muskat' varieties. the variety was firstly planted for an economic wine Moreover the question arose if 'GruÈ nmuskateller' was production to a greater extent during the time of ree- an individual variety and not identical to 'GruÈ ner Velt- stablishing the vineyards to avoid Phylloxera damage. liner'. It also was supposed that 'GruÈ nmuskateller' was TRUMMER (1841 and 1855) did not mention the variety an ancient type of today's variety. Due to the relatively when he created his ampelographic description of the low variability within the variety it can be supposed varieties used for Styrian wine production. He mentio- that the variety is much younger than other traditional ned the variety 'Muscatsylvaner' (synonym of 'Sauvi- varieties (HACK, 2007). Some morphologic differences gnon blanc') for the French production. Even GOETHE are caused by phytosanitary status and occurrence of (1887) did not count 'Sauvignon' as an important va- specific endophytes in plant material. riety for Styria. In a collection of ampelographic water- Firstly 'GruÈ nmuskateller' was mentioned by SPRENGER colour paintings created around 1850 (published recen- (1766) as a variety derived from OÈ denburg but with tly as a reprint) and presenting most of the varieties shallow description, which made identification impos- common in Styria and Slovenia no pictures are available sible. Older designations of the 'Veltliner' variety lack for 'Sauvignon' (KREUZER and KREUZER, 2001). There- quality in description for attributing the name to on of fore the Austrian production of 'Sauvignon' is not older today's varieties. In most cases it is not clear if they than one hundred years, but today it is of increasing meant the variety, the wine or the region Valtellina in importance for the cooler regions. It is supposed that Italy. It was also tried to find varieties related to 'Veltli- canes were brought from France for reestablishing vi- ner' in this Italian region, but without any success. neyards after phylloxera had devastated them. Just alike the origin of 'Sauvignon blanc' is also un- The aim of this work was to verify the heritage of 'GruÈ- known. It is supposed that Central France or Bordeaux ner Veltliner' and 'Sauvignon blanc' from 'Traminer' could be the place where this variety was selected. Ac- and even to define the second parental variety. Nowa- cording to historical literature the naming of the variety days identification of grapevine genotypes is done by was a persistent confusion. The separation from 'Savag- genetic fingerprints (THOMAS et al., 1993). The method nin' (French synonym for 'Traminer') was not perfor- is sensitive for differentiation, but stable enough to re- med in a satisfying manner. produce data under different conditions (THIS et al., Therefore clear definition happened very late in the 2004). Therefore in the meantime the easiest way to 19th century. Probable synonyms of 'Sauvignon' were identify unknown varieties is to compare the genetic 'Sauvagnin' (RENDU, 1857), 'Sauvagnien' (CHEVALIER, profile to existing data of different collections (VOSMAN 1860), 'Sauvoignin' (JULLIEN, 1832), and 'Sauvagneux' et al., 2001; THIS et al., 2004). In some cases it has al- (ODART, 1862) and others. ODART (1862) tried a clear ready been possible to show how identification of clo- differentiation for the first time and concluded that nes could function (TECHERA et al., 2004; REGNER et al., these both varieties were not closely related. VIALA 2006) and VERMOREL (1909) mentioned all types (vert, jaune, petit, fumeÂ, rose and violet). It seems that the special Material and methods flavour of 'Sauvignon' was not appreciated very much, as wines with strong ªsauvigonneª were blen- Plant material of all involved varieties is kept at the De- ded in cuveÂe wines. BOWERS and MEREDITH (1996) at partment for Grapevine Breeding at the Educational the UC Davis recognized by means of RFLP analysis and Research Centre for Viticulture and Pomology in the close relation of 'Traminer' and 'Sauvignon'. In Klosterneuburg. The material originated from different this study these two varieties show highest homology locations of Europe, especially samples of 'Chenin (about 93 %) among all involved varieties. Under blanc' and 'Sauvignon' were of French origin. The vine these circumstances 'Sauvignon' also could be traced from St. Georgen had been found at an unproductive back to mutations of 'Traminer'. Nevertheless only place (used for the cultivation of vines not later than areas where 'Traminer' ('Savagnin') was grown in the the 16th century) on slopes of the Leitha hills. It is a re- past can be assumed as geographical origin. Especially lict of former times and due to several dead trunks it

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seems realistic that this vine has derived from that time. Results Maybe the sandy soil kept off phylloxera and bushes and trees have protected the leaves against mildew dis- The genetic profile of 'Traminer' and 'GruÈ ner Veltliner' eases. share one allele at each locus, and therefore the conclu- The morphology of the vines was evaluated according sion that they have a close genetic relationship can be to the O.I.V descriptors (ANONYMOUS, 1983) and data drawn (Table 1). From the morphologic point of view were compared for their pronounced morphology. they differ more than it can be supposed from closely DNA from vines was extracted from young leaves by related varieties (Table 2). Therefore it was estimated following the protocol described by THOMAS et al. that the second parent of 'GruÈ ner Veltliner' dominated (1993) and modified by REGNER et al. (1998). The varie- the ampelographic character. But the second parent ties involved in this study were analysed at least with 36 could been found neither within the 'Veltliner' family SSR markers. nor within the collection of old and rare varieties kept The VVSmarkers were developed by T HOMAS and within the collection of Klosterneuburg. By luck an an- SCOTT (1993) and the VVMD markers by BOWERS et al. cient vine from St. Georgen near Eisenstadt was geno- (1996) as well as by BOWERS et al. (1999). The VRZAG typed and this variety seems to be the missing part of markers (SEFC et al., 1999) and all other markers were the origin of 'GruÈ ner Veltliner' (Table 1). Nevertheless used from the ªVitis Microsatellite Consortiumª collec- the genetic profile of this variety could not be confir- tion designated with internal coding of the consortium. med from any other of the numerous collections in Eu- Amplification was performed in 20 ml of the buffer so- rope. The profile was source of comparison of un- lution, which consisted of 16 mM (NH4)2SO4,67mM known varieties within the project Grapegen 06 but Tris-HCl pH = 8.8, 1.5 mM MgCl2, 0.01 % Tween 20, none of the participating collections found closer simi- 0.1 mM each dNTP (GenXpress, Maria WoÈ rth, Au- larities to known varieties. At that moment we can de- stria), 0.2 mM primer, 1 Unit SAVADY Taq DNA poly- fine the variety as an unknown variety. 244 alleles were merase (Peqlab, Erlangen, Germany), and 50 ng geno- obtained by genotyping 'GruÈ ner Veltliner' and the un- mic DNA of grapevine. known variety from St. Georgen at 120 loci. Only the A Mastercycler (Eppendorf, Hamburg, Germany) ther- markers not amplified in all involved varieties were ex- mocycler processed 36 cycles for SSRs. The amplifica- cluded from the studies. 'GruÈ ner Veltliner' and the un- tion of the SSR loci was performed by following our known variety from St. Georgen shared 144 identical general protocol but by applying specific annealing alleles. The differences are strong enough to exclude conditions. The general PCR protocol applied for these mutations as the source of 'GruÈ ner Veltliner'. On the studies was 2 min. denaturation at 94 8C and 35 cycles other hand they share almost at each locus one allele with annealing phase for 30 sec. (temperature between what confirms the hypothesis of hybridization with 45 8C and 55 8C) and denaturation for 15 sec. at 92 'Traminer'. It could be shown for all 19 chromosomes 8C. The annealing temperature for each locus was set that the proposed heritage is supported by the segrega- according to the Tm - 10 8C temperature. A final exten- ting alleles. Nevertheless some loci also exist that could sion of the fragments was performed at 72 8C for 5 min. not satisfy the criteria for a perfect heritage (Table 1a) Due to the different size range of the involved loci mul- but do not reach 10 % of the loci. Nevertheless our ex- tiplex PCR was feasible. At least the alleles of three loci periences from documented crossings have shown that were separated on one sequencing gel. deviations from the parental alleles increase in combi- Yield of DNA fragments was estimated by running an nations of genotypes with larger differences. As the un- aliquot of the sample on a 2 % agarose gel stained with known variety from St. Georgen shows more different ethidium bromide. The samples were denaturated by alleles, it could be that adaptation happened during re- heating up with formamide and loaded together with a combination. One of the alleles at VVMD 6 and at size standard (Genescan 350 Tamra, Appl. Biosystems, VVMD 32 are unique and could not be observed within Warrington, Great Britain) to a 6 % polyacrylamid gel. all these thousands of varieties analysed so far. One al- Detection of the SSR fragments labelled with the fluore- lele at the loci VRZag 15 and VRZag 62 is rare and scent dyes 6FAM, TET and HEX was carried out by an usually found in non V. vinifera varieties. automated sequencer (ABI 373, Perkin-Elmer, Vienna). In the meantime we successfully tried to propagate the Labelling with these different fluorescent colouring material because of the risk of losing the whole geno- agents facilitated the application of multiplex PCR. type due to the age of the single remaining vine. Fur-

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thermore we are curious to harvest first grapes and 2). While 'GruÈ ner Veltliner' lacks anthocyanins at the compare them with 'GruÈ ner Veltliner' grapes. The ma- buds and at the shoot tip St. Georgen vines produce a ture leaf of St. Georgen is very similar to the morpho- high amount of them (Table 2). logy of 'GruÈ ner Veltliner' leaves (Fig. 1) but the main The second parentage we currently investigated was the differences could be found at the shoot tip colour (Fig. origin of 'Sauvignon blanc'. Due to the high degree of

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same RFLP (BOWERS and MEREDITH, 1996) fragments it ferent alleles. But it seems that we are still able to ex- was supposed that 'Traminer' is a parental vine of 'Sau- clude mutations of 'Traminer' as the origin of 'Sauvi- vignon'. Furthermore it also could be that 'Sauvignon' gnon'. Besides the convincing number of alleles we is a mutated type from 'Traminer' ('Savagnin'). We de- should find in that case additional types closer to 'Sau- veloped 105 SSR loci (partly shown in Table 3) and ob- vignon'. From all other varieties with chance to be a pa- tained 220 alleles for 'Sauvignon' but 65 differed from rental type of 'Sauvignon' only 'Chenin blanc' remai- alleles from 'Traminer'. Within several 'Traminer' types ned. Especially 'Semillon', 'Muscadelle' and 'Sauternes (also 'Savagnin') we detected with the same loci 12 dif- blanc' were involved in this study but their profiles ex-

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Figure 1: Mature leaves of 'GruÈ ner Veltliner' (right leaf) Figure 2: Shoot tip of 'GruÈ ner Veltliner' (right shoot) and and St. Georgen in comparison St. Georgen in comparison clude a parentage for 'Sauvignon'. It was surprising that ties of the moderate European climate zones 'Traminer' from the 220 alleles 'Chenin blanc' shared 129 with parentage could be verified. That means that 'Traminer' 'Sauvignon' while 'Traminer' only reached 115 identical is one of the key varieties for the development of grape- alleles. But there are also some alleles which could not vine diversity. Besides the here presented varieties also be covered by one of the both parental genotypes, but 'Silvaner', 'Rotgipfler', 'Pinot' and 'Riesling' show 'Tra- they do not exceed 10 %. On the other hand we also miner' outcrossing behaviour (REGNER, 2000). Further found some differences between 'Sauvignon jaune', key varieties for the Mid European viticulture are the 'blanc' and 'rose' at 6 loci. That means in contrary to varieties 'Heunisch' and 'Roter Veltliner'. Both were 'GruÈ ner Veltliner' the variability within 'Sauvignon' used in large dimensions in former times but nowadays seems to be much higher. Finally we postulate for 'Sau- are slowly disappearing. vignon blanc' the parentage of 'Traminer' x 'Chenin' Regarding the parentage of 'GruÈ ner Veltliner' it could but with some minor changes that happened during hy- be concluded that the maintenance of old and rare va- bridization or mutations during former propagation. In rieties is essential. It seems that only a single vine of this case we are not able to offer a perfect heritage but the grapevine from St. Georgen has survived the centu- no other combination shows higher coincidence with ries without someone taking care for a specific name. the allelic profile of 'Sauvignon'. Some of the local growers suppose that the historical 'GruÈ nmuskateller' would be the right name for this va- Discussion riety but in reality neither a precise historical description nor any pictures are available. Identification of The heritage of traditional varieties was the objective of this variety seems impossible at the moment. As the numerous studies. On the one hand it is helpful to ampelographic traits are close to 'GruÈ ner Veltliner' it know the descent of today's varieties, on the other is realistic that the change from 'GruÈ nmuskateller' was hand it is important for future breeding strategies to not noticed. The problem of this name is also the impli- know all about their base. Since grapevine is an ancient cation that it should be a Muskat flavoured variety. At crop and has been used for viticulture for several thou- the moment no experiences about the taste of the gra- sands of years numerous selection steps were necessary pes from the vine from St. Georgen are available. The to reach the level of today's production. Some of the genetic analysis allows also the prediction of Muskat developments now can be followed, others are still un- flavour due to the alleles found at VrZag 79 (REGNER et known or simply generalized. We know that the diver- al., 2001). St. Georgen grapevine lacks these Muskat sity of Vitis vinifera has two reasons. One is the selec- specific alleles. tion out of wild vines in all viticultural regions of Eu- Another obstacle to these parentages are a not 100 % rope and the second is the continuous improvement of coincidence of the analysed alleles. First it could be ob- material by man (MULLINS et al., 1990). In several varie- served that both parents cover one of the alleles from

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the offspring but the second did not exist in the paren- BOWERS, J.E. and MEREDITH, C.P. 1996: Genetic similarities among wine grape cultivars revealed by Restriction Frag- tal vines. If there is only one allele length it can be sup- ment Length Polymorphism (RFLP) Analysis. J. Amer. posed that the second is a null allele and therefore lost Soc. Hort. Sci. 121(4): 620-624 for amplification. This is the case for the loci BOWERS, J.E. and MEREDITH, C.P. 1997: The parentage of a clas- sic wine grape, Cabernet Sauvignon. Nature Genetics 16: VMC3g8.2 and VMC 3d12 (Table 1) in the 'GruÈ ner 84-87 Veltliner' heritage and at the loci VMC 1d10, VRZag BOWERS, J.E., DANGL, G.S., VIGNANI, R. and MEREDITH, C.P. 26 and VMC 2h10 (Table 3a) for 'Sauvignon'. The true 1996: Isolation and characterization of new polymorphic simple sequence repeat loci in grape (Vitis vinifera L.). allele length could not be detected and such loci are Genome 39: 628-633 not helpful to confirm any hypothesis. BOWERS, J.E., DANGL, G.S. and MEREDITH, C.P. 1999: Develop- Secondly it seems possible that one of the alleles of a lo- ment and characterization of additional microsatellite DNA markers for grape. Amer. J. Enol. Vitic. 50(3): cus could not be covered by the parents as it was found 243-246 at VMC 8g8 for 'Sauvignon' (Table 3a) and for 'GruÈ ner BURGER, J. (1837): Systematische Klassifikation und Beschrei- Veltliner' at VVIP17, VMC 2c3 and VMC 1f10 (Table bung der in den oÈ sterreichischen WeingaÈrten vorkommenden Traubenarten. - Wien: Gerold, 1837 1a). In such case we only can suppose that mutations CHEVALIER, C. 1860: La vigne, les jardins et les vers a soie aÁ have changed the profile during hybridization or later Chenonceau, au XVIe sieÁcle. Ann. Soc. Agric. Sci., Arts by mutations during the propagation process. et Belles-Lettres DeÂp. Indre et Loire 39: 117-125 As we have analysed for the parentage of 'GruÈ ner Velt- Collaboration agreement for the Development of Grape Mi- crosatellite Markers (http://www.agrogene.com) liner' more than 120 and for 'Sauvignon blanc' 105 SSR GOETHE, H. (1887): Handbuch der Ampelographie, 2. Aufl. - loci we could find for each chromosome several alleles Berlin: Parey, 1887 supporting our hypothesis about parentages. None of HACK, R. (2007): Charakterisierung von verschiedenen Klonen der Rebsorte 'GruÈ ner Veltliner' mittels SSR Marker und the recently published heritages (BOWERS and MER- Implementierung einer MySQL-Datenbank mit Webin- EDITH, 1997) is based on such a high amount of genetic terface zur VeroÈ ffentlichung der gewonnenen Daten. - data. In both cases more than 90 % of the investigated Diplomarbeit FH Campus Wien, 2007 loci confirm our theory. We also could observe variabi- IMAZIO, S., LABRA, M., GRASSI, F., W INFIELD, M., BARDINI, M. and SCIENZA, A. 2002: Molecular tools for clone identifica- lity within the varieties. While 'Traminer' shows a hig- tion: The case of the grapevine cultivar 'Traminer'. Plant her rate (REGNER and KASERER, 2002) of deviations Breeding 121: 531-535 'Chenin blanc' and 'Sauvignon' differ only in some JULLIEN, A. (1832): Topographie de tous les vignobles connus, 3e eÂd. - Paris: Chez l'auteur, 1832 rare loci. Age and distribution of a variety is responsi- KREUZER, V. in KREUZER, C. (2001): Zbirka ampelografskih ble for the degree of variation and that suggests that upodobitev Vinzenza in Conrada Kreuzerja. - Maribor: Umetniski kabinet PrimozÆ Premzl, 2001 (Reprint) 'Traminer' is much older than the other varieties (IMA- MULLINS, M., BOUQUET, A. and WILLIAMS, L. (1990): Biology of ZIO et al., 2002). Nevertheless 'Traminer' can be confir- the grapevine. - Cambridge: Univ. Press, 1990 med as a key variety for the development of numerous ODART, A.P. (1862): AmpeÂlographie universelle ou traiteÂ des varieties by outcrossing and both 'GruÈ ner Veltliner' ceÂpages les plus estimeÂs dans tout les vignobles de quel- me and 'Sauvignon' were derived from this source. que renom, 5 ed. - Paris: Libraire Agric., 1862 REGNER, F. 2000: Die genetische Analyse von Rebsorten. Acknowledgements Dt.Weinbau-Jahrb. 51: 125-132 REGNER, F. 2007: Herkunft unserer Rebsorten: GruÈ ner Veltli- We are grateful to Mrs. CORNELIA EISENHELD for techni- ner, BlaufraÈnkisch und St. Laurent. Der Winzer (4): 12-15 cal support. Some of the analyses used for this study REGNER, F. and KASERER, H. 2002: Investigations into the genetic variability of 'Traminer' clones. Mitt. Klosterneuburg were gained in the frame of EC project Grapegen 06. 52: 177-186 REGNER, F., S TADLBAUER, A. und EISENHELD, C. 1998: Heunisch x FraÈnkisch, ein wichtiger Genpool europaÈischer Rebsor- References ten (Vitis vinifera L. sativa). Wein-Wiss. 53: 114-118 REGNER, F., S TADLBAUER, A. and EISENHELD, C. 2001: Molecular AMBROSI, H., DETTWEILER, E., RUÈ HL, E., SCHMID, J. und SCHU- markers for genotyping grapevine and for identifying MANN, F. (1994): Farbatlas Rebsorten. 300 Sorten und clones of traditional varieties. Acta Hortic. 546: 331-342 ihre Weine. - Stuttgart: Ulmer, 1994 REGNER, F., H ACK, R. and SANTIAGO, J.L. 2006: Highly variable ANONYMOUS (1983): Descriptor list for grapevine varieties and Vitis microsatellite loci for the identification of Pinot Vitis species. - Paris: OIV, 1983 noir clones. Vitis 45(2): 85-91 BABO, A.W. und MACH, E. (1881): Handbuch des Weinbaues R F., H ACK, R., HANAK, K., SCHREINER, A., BEISERT, B. und der Kellerwirtschaft. - Berlin: Parey, 1881 EGNER, und RAUHUT, D. 2008: VariabilitaÈt innerhalb der Rebsorte BASSERMANN-JORDAN, F. (1975): Geschichte des Weinbaus, 3. GruÈ ner Veltliner. Mitt. Klosterneuburg 58:105-116 Aufl. - Landau in der Pfalz: PfaÈlzische Verlagsanstalt, RENDU, V. (1857): AmpeÂlographie francËaise comprenant la sta- 1975 tistique, la description des meilleurs ceÂpages, l'analyse

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