Intra-varietal diversity of 31 accessions named as Tempranillo or described as synonyms of this representative Spanish grapevine variety was assessed using 48 morphological descriptors and 95 AFLP markers obtained using 2 primer combinations. We could identify homonyms as well as accessions belonging to varieties which, depending on their regional origin, are incorrectly cited as Tempranillo synonyms. Genetic analysis revealed a high level of genetic similarity between Tempranillo accessions (SG &gt;0.97), which could correspond to three highly related genotypes. This narrow genetic variation supports a monophyletic origin of Tempranillo, and could be the result of rapid multiplication and distribution of a very low number of clones.

Cabello, F.

Cabezas, J. A.

Cervera, M. T.

Chavez, J.

Martínez-Zapater, J. M.

Rodríguez-Torres, I.

English

JKI Open Journal Systems (Julius Kühn-Institut)

Vitis 41 (1), 3336 (2002)Varietal diversity within grapevine accessions of cv. TempranilloM.T. CERVERA 1), J.A. CABEZAS 1), I. RODRÍGUEZ-TORRES 2), J. CHAVEZ 2), F. CABELLO 2), and J.M. MARTÍNEZ-ZAPATER 1)1) Departamento de Genética Molecular de Plantas, Centro Nacional de Biotecnología, CSIC, Cantoblanco, Madrid, Españaand Departamento de Mejora Genética y Biotecnología, SGIT, INIA, Madrid, España2) Sección de Viticultura y Enología, Finca El Encín, IMIA, Alcalá de Henares, Madrid, EspañaCorrespondence to: Dr. J. M. MARTÍNEZ-ZAPATER, Departamento de Genética Molecular e Plantas, Centro Nacional de Biotecnología,CSIC, Campus de la Universidad Autónoma de Madrid, Cantoblanco, 28049 Madrid, Spain. Fax: +34-91-585-4506; E-mail:zapater@cnb.uam.esSummaryIntra-varietal diversity of 31 accessions named asTempranillo or described as synonyms of this representa-tive Spanish grapevine variety was assessed using 48 mor-phological descriptors and 95 AFLP markers obtained us-ing 2 primer combinations. We could identify homonymsas well as accessions belonging to varieties which, depend-ing on their regional origin, are incorrectly cited asTempranillo synonyms. Genetic analysis revealed a highlevel of genetic similarity between Tempranillo accessions(SG >0.97), which could correspond to three highly relatedgenotypes. This narrow genetic variation supports a mono-phyletic origin of Tempranillo, and could be the result ofrapid multiplication and distribution of a very low numberof clones.K e y   w o r d s :  ampelography, grapevine genetic identifica-tion, monophyletic origin, intra-varietal variability, Tempranillo.IntroductionThe Spanish word temprano means early, and there-fore the name Tempranillo is logically related to the earlyripening of this Spanish red wine variety. Contrary to mostother Mediterranean varieties, Tempranillo has a short veg-etative cycle, a characteristic of Northern varieties, and largehairy hepta-lobullated leaves as well as uniform fruit set anddark-blue berries with thick skin. Tempranillo is widely culti-vated in northern and central Spain where it is the mainvariety in half of the Denominations of Origin (DOs). In vari-ous regions of Spain Tempranillo has different names e.g.Cencibel, Tinto Fino, Tinto del País, Tinta de Toro, Ull deLlebre, etc. Nowadays, original local names associated withTempranillo have been given to other cultivars (homonymies)or different names have been given to clones of Tempranilloin different Spanish regions (synonymies) (RODRÍGUEZ-TORRES 2001). We have used ampelographic and AFLP mo-lecular markers (VOS et al. 1995) to characterize the differentgenotypes sharing this common name as well as the pro-posed synonyms. The results show narrow intra-varietalgenetic variation and allow to identify synonyms and tocharacterize false synonyms and homonyms.Material and MethodsThirty-one accessions of Vitis vinifera described asTempranillo or related synonyms were analyzed (Table). Inaddition the rootstock variety R-110 (Vitis berlandieri xV. rupestris) as a reference in the AFLP analysis is included(CERVERA et al. 1998, 2000, 2001). Representative individualsof these varieties are cultivated in the collection of El Encín,(Instituto Madrileño de Investigación Agraria y Alimentaria(IMIA) de la Comunidad de Madrid, Alcalá de Henares,Spain). Morphological characterization was performed fol-lowing the descriptors of the O.I.V. (1984). This descriptionwas performed in two consecutive years by three  ampelo-graphers and modal values of the 6 observations were con-sidered. Statistical analysis was carried out as described byCERVERA et al. (2001). Total genomic DNA extraction, AFLPand statistical analyses were performed according toCERVERA et al. (1998). Dendrograms were constructed usingthe UPGMA clustering method to the ampelographic datadistance matrix calculated using the correlation distance in-dex and Dice estimates of genetic similarities based on AFLPanalysis, respectively.ResultsA total of 48 descriptors (O.I.V. 1984) were used to char-acterize 31 accessions known as Tempranillo or Tempranillosynonyms. Phenetic distances between each pair of acces-sions are presented in Figure, a. The co-phenetic correlationwas 0.98 (p=0.002) indicating a good fit of the cluster analy-sis. Three different clusters were identified. The first cluster,from Tempranillo (VI) to Tempranillo (NA), included 26 ac-cessions organized in 5 sub-clusters and grouped at a ge-netic similarity level (GS) of 0.89. The second cluster, fromTinta de Madrid (TO) to Tinto del Pais (BU), grouped 4 un-related accessions with GS <0.65 and with Tempranillo ac-cessions in cluster I. The third cluster, Tempranillo Temprano(LO), was even less related with the other accessions (GS<0.32). The results showed phenotypic heterogeneity amongthe samples, which may indicate the presence of more thanone variety (clusters II and III). To unequivocally discrimi-nate Tempranillo accessions and associated synonyms weanalyzed these samples using high throughput molecularmarkers (AFLPs).34 M.T. CERVERA et al.The analysis of 31 accessions of Tempranillo, proposedsynonyms and the rootstock R-110 using two AFLP primercombinations yielded a total of 206 bands of which 95 poly-morphic bands were scored. A dendrogram illustrates ge-netic similarity values obtained from AFLP data (Figure, b).The co-phenetic correlation was 0.99 (p=0.002) indicating agood fit of the cluster analysis. The accessions showed GSlevels between 0.53 and 1. Based on previous results(CERVERA et al. 1998), accessions showing similarities >0.90can be considered as belonging to the same variety whiledifferent varieties show similarities from 0.60 to 0.90. Thedendrogram shows the existence of one main group of ac-cessions including all Tempranillo samples with GS values>0.97 and excluding Tempranillo Temprano (LO), Tinto Ma-drid (S), Tinta de Madrid (TO), Tinto del País (BU) and TintoMadrid (VA). No polymorphism at the DNA level was de-tected between Tempranillo (VI, LO, B, BU), Tinta de Toro(ZA), Tinto Fino (AB), Cencibel (GU), Tinto del País (VA-1,VA-2), and Tinto Madrid (M, SA, TO), as well as betweenCencibel (CR-1, CR-2, CU, M), Tempranillo (CC, ZA),Chinchillana (BA), Tinta de Madrid (ZA), Tinta Madrid (LE),Juan García (ZA), Tinto del País (SO), Escobera (BA), andTinto Madrid (SO). These two sub-clusters were grouped atGS >0.98, which is due to the existence of one polymorphismbetween them. Tempranillo (NA) showed GS >0.98 (one poly-morphism) and 0.97 (two polymorphisms) respectively withthe first and second group of Tempranillo.The low genetic similarity observed when compared toother Tempranillo accessions (GS <0.63) identified Tem-pranillo Temprano (LO) as a different homonym variety. Thisanalysis also allowed the identification of false synonymssuch as Tinto Madrid (S), Tinta de Madrid (TO) as well asthe related accessions Tinto del País (BU) and Tinto Madrid(VA), which should correspond to unrelated varieties.DiscussionTempranillo has long been considered as an example ofso-called variety-populations, consisting of plants of differ-T a b l eGrapevine accessions analyzedCode Local name Author of synonym description OriginA09 Tempranillo (VI) Vitoria (VI)A17 Tempranillo Temprano (LO) Logroño (LO)A23 Tempranillo (LO) Logroño (LO)A40 Tempranillo (NA) Navarra (NA)F42 Tinto Madrid (S) PEÑÍN et al. (1997) Santander (S)G18 Tempranillo (B) Barcelona (B)H37 Tinto Fino (AB) GARCÍA DE LOS SALMONES (1914) Albacete (AB)I25 Cencibel (CR-1) GARCÍA DE LOS SALMONES (1914) Ciudad Real (CR)I29 Cencibel (CR-2) GARCÍA DE LOS SALMONES (1914) Ciudad Real (CR)I41 Cencibel (CU) GARCÍA DE LOS SALMONES (1914) Cuenca (CU)I55 Cencibel (GU) GARCÍA DE LOS SALMONES (1914) Guadalajara (GU)J02 Tinto Madrid (M) PEÑÍN et al. (1997) Madrid (M)J18 Cencibel (M) GARCÍA DE LOS SALMONES (1914) Madrid (M)J37 Tinta de Madrid (TO) CLEMENTE (1807) Toledo (TO)J49 Tinto Madrid (TO) PEÑÍN et al. (1997) Toledo (TO)J52 Tempranillo de Rioja (BU) Burgos (BU)J53 Tinto del País (BU) BORREGO et al. (1990) Burgos (BU)K32 Tinta Madrid (LE) CLEMENTE (1807) León (LE)K47 Tinto Madrid (SA) PEÑÍN et al. (1997) Salamanca (SA)L11 Juan García (ZA) CLEMENTE (1807) Zamora (ZA)L12 Tinto del País (SO) BORREGO et al. (1990) Soria (SO)L15 Tinto Madrid (SO) PEÑÍN et al. (1997) Soria (SO)L16 Tinto del País (VA-1) BORREGO et al. (1990) Valladolid (VA)L18 Tinto Madrid (VA) PEÑÍN et al. (1997) Valladolid (VA)L29 Tinto del País (VA-2) BORREGO et al. (1990) Valladolid (VA)L56 Tinta de Madrid (ZA) CLEMENTE (1807) Zamora (ZA)L57 Tempranillo (ZA) Zamora (ZA)M04 Tinta de Toro (ZA) GARCÍA DE LOS SALMONES (1914) Zamora (ZA)M10 Chinchillana (BA) PEÑÍN et al. (1997) Badajoz (BA)M11 Escobera (BA) PEÑÍN et al. (1997) Badajoz (BA)M31 Tempranillo (CC) Cáceres (CC)ent genetic origin and sharing morphological characteris-tics (CABELLO and DE LA FUENTE 1993). However, geneticcharacterization of accessions based on AFLP markers indi-cated that the variety Tempranillo consists of various cloneswith a higher level of genetic similarity (SG >0.97) than ob-served within other Spanish varieties such as Albillo (0.92),Turruntés (0.87) and Malvasía del Bierzo (0.91), whenanalyzed using the same AFLP primer combinations (CERVERAet al. 2001). This result strongly supports a monophyleticorigin for Tempranillo. The narrow intra-varietal genetic di-versity of Tempranillo may be explained either by a lowersomatic variation rate as compared to other varieties, or areduction of the genetic variability caused by vegetativepropagation of low numbers of genotypes. The26 Tempranillo accessions analyzed may derive from threeclosely related genotypes. The first group of Tempranillosgrouped 12 accessions (from Tempranillo (VI) to Tempranillode Rioja (BU), Figure, b), including a pre-phylloxeric acces-sion collected in 1893 (Tempranillo (LO)). The second group,with 13 accessions (from Cencibel (CR-1) to Cencibel (M)),includes three identical accessions collected from post-phylloxeric plantations in the first half of the 20th century(Cencibel (M), Chinchillana (BA) and Escobera (BA)). Fi-nally, the third group is represented by a single pre-phylloxeric accession collected in Navarra and recentlyadded to the grapevine collection (Tempranillo (NA)). If theold accessions of the three groups are representatives ofthe original clones which were used to re-plant youngervineyards, then clones grown nowadays under the nameTempranillo, as well as related synonyms, may only repre-sent a part of the original variability of this ancient Spanishvariety, while other local ancient clones may have been wipedout and with them most of the intra-varietal diversity.In contrast, ampelographic diversity within Tempranillowas apparently higher than the genetic variability estimatedby AFLP. This result however  may be due to the combineduse of unstable or subjective descriptors (such as densityof petiole hairs) and descriptors, such as berry shape, whichare not measured as quantitative traits. In this way, slightvariation may turn into highly variable values overestimatedin the statistical analysis. In any case, no statistically sig-nificant differences were found if ampelographic descrip-tions of accessions belonging to the three Tempranillo geno-types were compared and thus, no discriminatory descriptorswere identified.The ampelographic and genetic analysis also allowedthe identification of Tempranillo synonyms. In addition toCencibel, Tinto Fino, Tinta de Madrid, Tinta Madrid, TintoMadrid, Tinto del País, Tinta de Toro, Chinchillana, andEscobera we also identified Juan García as a new synonym(Table). It is important to note that all clones belonging toTinta de Madrid, Tinta Madrid, Tinto Madrid and Tinto delPaís can not be considered to be Tempranillo. These varie-ties are heterogeneous at the genetic level grouping Tem-pranillo accessions (Tinto Madrid (M, SA, TO, SO), TintaMadrid (LE), Tinta de Madrid (ZA) and Tinto del País (VA-1,VA-2, SO)) together with clones belonging to other varieties(Tinta de Madrid (TO), Tinto Madrid (S, VA) and Tinto delPaís (BU)) of the same regions. Based on ampelographicdescriptions and AFLP analysis, some of the false syno-nyms and the homonym were assigned to other varieties.The homonym Tempranillo Temprano (LO), was identifiedFigure: Dendrograms representing morphological (a) and genetic (b) relationships among Tempranillo accessions and proposedsynonyms.Varietal diversity of cv. Tempranillo 3536 M.T. CERVERA et al.as an accession of Moristel (GS > 0.93), an autochthonousand ancient variety of northeastern Spain. The false syno-nyms Tinta de Madrid (TO) and Tinto del País (BU) wereidentified as Garnacha (GS > 0.96) and Bobal (GS > 0.93),respectively.AcknowledgementsThe authors would like to thank to CARLOS A. MALPICA forcritical reading of the manuscript, and INÉS POVEDA and ANGEL SANZfor the photographic artwork. M.-T.C. was funded by a MECcontract; J.A.C., I.R. and J.C. were funded by predoctoral fellow-ships from Instituto Nacional de Investigación y Tecnología Agrariay Alimentaria, European Union, and Agencia Española deCooperación Internacional, respectively. This research was fundedby projects from INIA SC00-033 and CM 07B-0033-1999. Sup-port for research activity at the Centro Nacional de Biotecnologíais provided by a CSIC-INIA specific agreement.ReferencesCABELLO, F.; DE LA FUENTE, J. M.; 1993: Caracterización de la variedadTempranillo y sus sinonimias Españolas. XV Jornadas deViticultura y Enología de Tierra de Barros, Dirección General deComercio e Industrias Agrarias de la Consejería de Agricultura yComercio de la Junta de Extremadura.CERVERA, M. T.; CABEZAS, J. A.; SANCHA, J. C.; MARTÍNEZ DE TODA, F.;MARTÍNEZ-ZAPATER, J. M.; 1998: Application of AFLPs to the char-acterization of grapevine Vitis vinifera L. genetic resources. Acase study with accessions from Rioja (Spain). Theor. Appl. Genet.97, 51-59.- -; - -; SÁNCHEZ-ESCRIBANO, E.; CENIS, J. L.; MARTÍNEZ- ZAPATER, J. M.;2000: Characterization of genetic variation within table grapevarieties (Vitis vinifera L.) based on AFLP markers. Vitis 39,109-114.- -; RODRÍGEZ, I.; CABEZAS, J. A.; CHÁVEZ, J.; MARTÍNEZ-ZAPATER, J. M.;CABELLO, F.; 2001: Morphological and molecular characteriza-tion of grapevine accessions known as Albillo, one of the oldestvarieties grown in Spain. Am. J. Enol. Vitic. 52, 127-135.CLEMENTE, S.; 1807: Ensayo Sobre las Variedades de la Vid Común queVegetan en Andalucía, con un Indice Etimológico y tres Listas dePlantas en que se Caracterizan Varias Especies Nuevas. Poulet,Madrid.GARCÍA DE LOS SALMONES, N.; 1914: Colección de Plantas de los CamposOficiales. Recopilación. Instituto Nacional de InvestigacionesAgronómicas, Madrid.O.I.V.; 1984: Codes des Caractères Descriptifs des Variétés et Espècesde Vitis. A. Dedon, Paris.PEÑÍN, J.; CERVERA, A.; CABELLO, F; DIEZ, R.; PEÑÍN, P.; 1997: Cepas delMundo. Ediciones PI y Erre, Madrid.RODRÍGUEZ-TORRES, I.; 2001: Caracterización de variedades de vid pormétodos ampelográficos y bioquímicos. Resolución dehomonimias y sinonimias. Thesis E.T.S.I. Agrónomos.Universidad Politécnica de Madrid.VOS, P.; HOGERS, R.; BLEEKER, M.; REIJANS, M.; VAN DE LEE, T.; HORNES, M.;FRIJTERS, A.; POT, J.; PELEMAN, J.; KUIPER, M.; ZABEAU, M.; 1995:AFLP: A new technique for DNA fingerprinting. Nucl. 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Varietal diversity within grapevine accessions of cv. Tempranillo

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Varietal diversity within grapevine accessions of cv. Tempranillo

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