Anthocyanin Composition in Carignan and Grenache Grapes and Wines As Affected by Plant Vigor and Bunch Uniformity
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08-edo_05b-tomazic 24/09/14 20:37 Page201 ANTHOCYANIN COMPOSITION IN CARIGNAN AND GRENACHE GRAPES AND WINES AS AFFECTED BY PLANT VIGOR AND BUNCH UNIFORMITY Maite EDO-ROCA *,M ontse NADAL , AntoniSÁNCHEZ-ORTIZ and Míriam LAMPREAVE DepartamentdeBioquímicaiBiotecnologia,Facultatd’Enologia,UniversitatRoviraiVirgili, Marcel·líDomingos/n,43007,Tarragona,Catalonia,Spain Abstract Résumé Aims : TodeterminetheanthocyanincompositioninCarignan Objectifs :Déterminerlesteneursenanthocyanesdesraisins andGrenachegrapesandwinesasaffectedbyvintage,plant etdesvinsdeCarignanetdeGrenachesousl’influencedela vigorandbunchuniformity. climatologiedumillésime,delavigueurdesplantesetdela variabilitédanslesgrappes. Methods and results :AnthocyanincompositionofCarignan andGrenachegrapesandwineswereanalysedby Méthodes et résultats :LesraisinsetlesvinsdeCarignanet chromatographictechniquesconsideringtheinfluenceoftwo deGrenacheontétéétudiéspardestechniques differentvigorlevelsovertwovintages.Theheterogeneityin chromatographiquesàdeuxniveauxdifférentsdevigueur thedistalpartsofthebunchwasalsotakenintoaccount.Warm pendantdeuxannées.Lavariabilitédanslespartiesdistalesde vintagewasbetterfortheaccumulationofanthocyanins. lagrappeaégalementétépriseencompte.Lemillésimechaud However ,eachvarietyresponseddifferentlyaccordingtovine étaitfavorableàl’accumulationdesanthocyanes.Parcontre, vigor. Grenacheanthocyaninsynthesisdecreasedinlowvigor chaquevariétéaréagidifféremmentenfonctiondelavigueur (weak)vines,whereasCarignananthocyanincontentdepended delavigne.LaconcentrationdesanthocyanesenGrenachea onvigor,berrysize,rootstockandvintage. Inbothvarieties, diminuédanslesconditionsdefaiblevigueur,alorsque butmoresignificantlyinCarignan,therewasatendencyto l’accumulationdesanthocyanesenCarignanétaitfonctionde accumulateacylatedanthocyaninsinbottomberries. lavigueur,dupoidsdelabaie,duporte-greffeetdumillésime. Pourlesdeuxcépages,maisdefaçonplussignificativepourle Conclusion : Carignananthocyaninconcentrationwas Carignan,lescomposésacyléss’accumulaient increased inlowvigorplants,where clustersreceivedgreater préférentiellementdanslesbaiesinférieures. sunexposure,unlikeGrenache,where bettercanopy managementinthefruitzoneisnecessary. Avoidingthepoor Conclusion :Lasynthèsed’anthocyanesdansleCarignanest growingconditionsforGrenacheintheregionandimproving favoriséeenconditionsdefaiblevigueur,favorablesàune thecanopy/fruitratiodeservescarefulconsiderationinorderto bonneexpositiondesgrappes,tandisquelafaiblevigueur reachoptimalanthocyanincontent. affectenégativementleGrenache.Chezcedernier,lagestion delasurfacefoliairedanslazonefructifèreestnécessairepour Significance and impact of the study : Knowledgeof maintenirunecouverturevégétaleoptimale(surface anthocyaninaccumulationaccordingtobothplantvigorand foliaire/récolte)etprotégerlesgrappes. bunchripenessisofmajorimportancetodeterminethe optimalharvestdateforeachcultivarandthusimprovethe Importance et impact de l’étude :Connaîtrelacomposition qualityofwine. enanthocyanesdesgrappesdeCarignanetdeGrenacheen fonctiondelavigueuretdelavariabilitédelamaturitédans Key words :ripenessvariability,vigor,vintage,anthocyanins, lesgrappespeutaideràgérerlacouverturevégétaledes Carignan,Grenache plantesafind’obteniruneconcentrationd’anthocyanesplus élevéedanslesraisinsetlesvinsproduits. Mots clés :variabilitédelamaturité,vigueur,vintage, anthocyanes,Carignan,Grenache manuscript received 29th April 2013 - revised manuscript received 6th March 2014 J. Int. Sci. Vigne Vin , 2014, 48 , 201-217 *Correspondingauthor : [email protected] - 201 - ©Vigne et Vin Publications Internationales (Bordeaux, France) 08-edo_05b-tomazic 24/09/14 20:37 Page202 Maite EDO-ROCA et al. INTRODUCTION petunidin-3- O-glucoside,anddelphinidin-3- O- glucoside)anthocyanins.AccordingtoCohen et al. Grapesofredvarietiescontainlargeamountsof (2008),theproportionofdioxygenatedanthocyanins anthocyanincompoundsintheskins,andsometimesin shouldincreaseunderconditionsoflowdaytime thepulptoo.Theseanthocyaninsarepartiallyextracted temperatures.Conversely,Guidoni et al. (2008)referred duringwinemaking.FortheEuropean Vitis vinifera tothesensitivityoftheenzymeF3’Htotemperaturein species,therearefivemonoglucosideanthocyaninsand ordertojustifythehighconcentrationsofdioxygenated theircorrespondingacylatederivatives(acetylandp- anthocyaninsfoundinNebbiolograpesduringawarm coumaryl);malvidin-3- O-glucosideisthemostabundant yearasopposedtoacoolyear. anthocyanin(Pomar et al. ,2005),althoughinsome varietiespeonidin-3- O-glucosidepredominates(Mattivi Thedistributionofanthocyaninsbetweentheacylated et al. ,2006).Theevolutionofanthocyaninsduring andnon-acylatedformsisalteredbythecombinationof veraisonandripeningcorrespondstodifferent temperatureandsolarradiation.Thisisbecause enzymaticactivitiesatthecellularlevel(Castellarín et acyltransferaseactivityincreaseswithtemperature al. ,2011).InHe et al. (2010)’sreview,twosynthetic (Haselgrove et al. ,2000,Spayd et al. ,2002).This pathwaysweredescribedfromnaringeninflavanone : combinationiscomplexbecauseitcanbesynergisticif onethatgeneratesanthocyanidincyanidinfromthe thetemperatureismoderateorantagonisticifthe actionofF3’H(flavonoid3’-hydroxylase),DFR temperatureisextreme(Tarara et al. ,2008).Downey et (dihydroflavonol4-reductase)andANS(anthocyanidin al. (2004)observedthatunderconditionsofmoderate syntase)andanotherthatgeneratesanthocyanidin temperatureandshading,theproportionofp- delphinidinfromtheactionofF3’5’H(flavonoid3’,5’- coumarylatedanthocyaninswasincreasedwithrespect hydroxylase),DFRandANS.TheactionofUFGT tonon-acylatedandacetylatedanthocyanins.However, (UDPglucose :flavonoid-3- O-glucosyltransferase) ifthetemperaturesweretoohigh,theconcentrationof generatescyanidin-3- O-glucosidefromcyanidinand, totalanthocyaninswoulddecreaseduetothe finally,peonidin-3- O-glucosidebytheactionofOMT degradationofnon-acylatedandacetylated (O-methyltransferase).UFGTalsosynthesizes anthocyanins,whicharelessstablethanp-coumarylated delphinidin-3- O-glucosidefromdelphinidinandthe anthocyanins(Rodriguez-Saona et al. ,1999).This actionofOMTgeneratespetunidin-3- O-glucosideand decreasewasmorepronouncedinshadedberries. This malvidin-3- O-glucoside. Attheendoftheripening suggeststhatwhilehightemperaturescaninduce process,thesynthesisandaccumulationofthenon- anthocyanindegradation,solarradiationcaninduce acylatedanthocyaninsissloweddownorevenstopped anthocyaninbiosynthesis. Italsomeansthatthereare whereasthereissomeincreasewiththeacylated probablytwosystemsofregulationandaccumulationof derivatives(p-coumarylatedand/oracetylated) anthocyanins :ononehand,asystemthatgenerates (González-SanJoséandDíez,1992,Jordao et al. ,1998, anthocyaninsprogressivelyduringtheripeningperiod RyanandRevilla,2003). and,ontheotherhand,asysteminducedbyradiation exposure.However,inwarmregions,theprolonged Anthocyanincomposition dependsontheinteraction exposureofbunchtodirectsunlightand,consequently, betweenclimate,soil,viticulturalpracticesandgenotype tohightemperatures,shouldbeavoidedinorderto (JacksonandLombard,1993,Downey et al. ,2006), preventtheslowdownorshutdownofanthocyanin whichinvolvesvariationsintheexpressionofgenes synthesisandmaximizeberrycolor(Spayd et al. ,2002, codingfordifferentenzymes(Yamane et al. ,2006). NadalandLampreave,2007,Chorti et al. ,2010). Canopymanagementmodifiesthegrowthandstructure ofthevine,causingchangesinexposuretosolar Itiswellknownthattheoptimumtemperaturefor radiationandtemperatureinthefruitzone(Smart,1985, anthocyaninsynthesisisaround30ºC,andthathigher Smart,1987,Bergqvist et al. ,2001).Densecanopies valuesupto 35 ºCinhibitit.Therefore,modificationof increasethelevelofshadinginthefruitzone,causinga thevinemicroclimatethroughcanopymanagementcan reductionintheactivityofF3’5’Horanincreaseinthe preventexcessivesunlightandhightemperaturesfrom activityofF3’Handhenceanincreaseinthe reachingthebunchandimproveanthocyanincontent concentrationofdioxygenatedanthocyanins :peonidin- (Downey et al. ,2003).Moreover,densecanopiescan 3- O-glucosideandcyanidin-3- O-glucoside(Downey et causeunevenberryripeningwithinthebunch.This al. ,2004).Someauthors,however,foundquitethe heterogeneitycanleadtovariationsinthefinal oppositeresultincolderweatherconditions,suchasin compositionofthegrapeintermsofsugar,acidity, northernItaly(Chorti et al. ,2010)andnorth-eastern aromaandcolor(Kasimatis et al. ,1975,Tarterand UnitedStates(Tarara et al. ,2008).Temperatureisalsoa Keuter,2005,PagayandCheng,2010).Itisnotaneasy factorthataltersthebiosyntheticpathwayof tasktodeterminetheidealharvestdate,whengrapes dioxygenatedortrioxygenated(malvidin-3- O-glucoside, havereachedtheiroptimumcompositionforaparticular J. Int. Sci. Vigne Vin , 2014, 48 , 201-217 ©Vigne et Vin Publications Internationales (Bordeaux, France) - 202 - 08-edo_05b-tomazic 24/09/14 20:37 Page203 styleofwine.Inthiscontext,itisnecessarytoevaluate describedin Edo-Roca et al. (2013) .Sugars,acidsand theripenessofthepulpandskintoimprovethe phenolicsweredeterminedineachtopandbottomhalf knowledgeofthegeneticpotentialofeachvariety.Inred ofthebunch.Carignanclustersareinverted-cone- varieties,winequalitydependsontheproperripeningof shapedandtheirtoppartsrepresentedupto60 %ofthe theskinandseedsandthenatureoftheirphenolic totalweightofthebunch.However,inGrenache,this compounds.AuthorssuchasAmraniJouteiandGlories percentagewasincreasedupto65 % becausethe (1994)andRibéreau-Gayon et al. (2000)havealso shouldersarelocatedatthetoppositionofthecluster . suggestedusingtheanthocyaninconcentrationasatool Foreachdistalpartofthebunch,asampleof100 forthedeterminationofharvestdates.Theextractionof