花生脂肪及脂肪酸含量的遗传分析

花生脂肪及脂肪酸含量的遗传 AgriculturalScience&Technology,2011,12(7):943一,946,974 Copyright? 2011,InformationInstituteofHAAS.AllrightsresewedAgriculturalBiotechnology InheritanceofFatandFattyAcidCompositionsin Peanut(ArachishypogaeaL_) ZHANGXin— you'',ZHUShui-jin',HANSuo-yi,XUJing.,LIUHua,TANGFeng-shou.,DONGWen. zhao,ZANGXiu—wang,ZHANGZhong—xin 1.CollegeofAgricultureandBiotechnology,ZhejiangUniversity,Hangzhou310029;2IndustrialCropsResearchInstitute.HenanAcademy ofAgriculturalSciences,KeyLaboratoryForGeneticImprovementofOilbearingCropsinHenanProvince, PeanutSub-centeroftheNational CenterforOil-bearingCropsImprovement,Zhengzhou450002 Abstract[0bjective]Theaimwastoexploretheinheritanceoffatandfatlyacidcompositionsjnpeanut.『Method]Geneticandcorrelation analyseswereperformedusingamixedgeneticmodelofmajorgenepluspolygenewith215F9RILsandtheirparentsZHENG8903andYUHUA No.4.[Result]TheoptimalgeneticmodeIofthefatcontentwastwolinkedcomplementarymajorgenepluspolygeneinheritancemode1.The heritabilityofmajorgeneswas22.88%.Oleic,linoleic,andpalmiticacidcontentwerecontrolledbytwomajorgenesandtheheritabilitywas75. 11%,75. ?%and6O.95%,respectivelyStearicandarachidicacidcontentwerecontrolledbytwomajorgenespluspolygenes.theheritability ofmajorgeneswas26.43%and33.17%,respectively.O/Lratioandbehenicacidcontentwere controlledbythreemajorgenes.theheritability was70.65%and30.70%,respectively.』 ConclusionlTheresultinthisstudywasthataccumulatingpolygenesshouldbeemphasizedin the}m provementoffatcontent.whileutilizationofmajorgenesisenhancedinimprovementofoleic andlinoleicacids. Keywo?rdsPeanut;Fat;Fattyacidcomposition;Inheritance Peanutjsoneoftheimportantoilseedcropsjntheworld: itoccupiesakeypositioninedibleoiIprocessing.Thekernel oilratesofpeanutgenerallyrangefrOm46%to57%.ofwhich oleicandIinoleicacidconstitute8O%ofthetota.PeanutoiI jsalsooneofthegoodediblevegetableoils.whichisanother reasonwhypeanutsarepopularwithconsumers-3j.New cultivarsofpeanutsshouldnotonlyhavehighoiIcontent.but alsoareasonablefartyacidcompositionto;ncreasinglyim— provehumanhealth. Understandinathegeneticmechanismoffatandfatty acidsinpeanutisanimportantpremiseandfoundationfor peanutbreeding.Previousstudiesshowedthatthegenetic characteris"csoffatcontentarepresentedascomplexquanti— tativetraits.Currently.analysisofQTLinplantsfoundthat thegeneticeffectsofgenescontrollingquantitativetraitwere different.ClassicaIquantitativegeneticscanuncoverthecom- binedeffectofgenes.butcannotdetectindividuaIeffects. Thus,themixedgeneticmodelofmajorgenepluspolygeneis establishedonthebasisofpreviousstudies:themodelhas beenwidelyusedbecauseitcandetectspecificgene effectsl.-ej.Althoughotherresearchershavecarriedoutfruit— fufstudiesonpeanutgeneticsoffatcontent.uptonow.there isnooptimalmixedgeneticmodelofmajorgenepluspoly— geneforpeanutfatandfattyacidcompositions. Inthisstudy.thesegregationanalysismethodofthema- jorgenepluspolygenemixedgeneticmodelwasusedtostudy thegeneticmechanismoffatandfattyacidcompositions.The relativeimpodanceofallthemajorgenesandpolygeneswas explored.Thisresearchprovidesascientificbasisforthege- neticimprovementofpeanutoil. Received:June28,2011Accepted:July12,2011 ScienceandTechnologyInnovationTale, ntsSchemeinHenanProv- ince(104200510003). $Correspondingauthor.E-mail:shjzhu@zju.edu.cn MaterialsandMethods Plantmaterials Thematerialswererecombinantinbredlinepopulation (F.)with215lines.Crosscombinationthatbetweenahighly yieldIineZHENG8903ffemaleparent)andavariety.YUHUA No.4(maleparent)werebothbredbyHenanAcademyof AgriculturalScience.ZHENG8903wasgoodresistantIine, YUHUANo.4hadsuperioragronomiccharacterswithhigh yieldandearlymatureanditwasreleasedbyHenanProvince In1991.TheRILswasconstructedbysingleseeddescendant (SSD)methods.Therewasgreatdifferencebetweenthetwo parentsandthepopulationwasgeneticallystable. Fieldexperiment ThefieldtrialwascarriedoutattheexperimentalfarmOf theHenanAcademyofAgriculturalSciences(Yuangyang, Henan.China)inMay2008.Theseedswereplantedfollow- jngarandomizedblockdesign:theprocedurewasrepeated twotimes.Plantingdensitywas15cmx40cm.anda1O0 cmwideprotectivebeltwasIaidaroundtheplantedseeds. Methods DeterminationoffatandfattyacidsFatandfattyacids weredeterminedbytheAgriculturafProductsQualitySupervi— sionandTestingCenter(Zhengzhou)oftheMinistryofAgri— culture.Thesamplesusedwerepost—harvestseedsstoredfor 3—4months.1neachsample.fatandfartyacidswereidenti— fledtwice.andtheaveragewastaken. DeterminationoffatcontentThefatcontentofvarious materialswasdeterminedbytheSoxhletextractionmethod.in whichtheperformancestandardwasGB/T14772—1993.The protocolsfordetectionconsistedofsampledryingusinganhy- drousetherextractionandetherremova1.Theremainingresi. dueobtainedwasfat. DeterminationoffattyacidsFattyacidcontentwasdeter. minedbycapillarygaschromatographyanalysisforeachsam— pie.inwhichtheperformancestandardwasGB/T17377. 1998.Thequantitativedeterminationoffattyacidcomposition wascarriedoutusingtheinternaInormalizationmethod.in whichthetotaIpeakarearepresents1OO%ofthecomposi- tion,assumingthatalIcomponentsofthesamplearedis- playedinthechromatogram. Dataprocessing PhenotypicdataanalysisDescriptivestatisticsandanaly. sisofvariance(ANOVA)offatandfattyacidcontentwere performedtotestdifferencesamongRILs.Geneticvariability amongtheRILs,coefficientofvariance(C,kurtosisvalue andskewnessvaluewereestimated.respectively.Correlation coefficientfr1amongthedifferenttraitswasalsoestimaled. Allnecessarycomputationforthetraitswasperformedwith theSPSS(StatisticalProductandServiceSolutions)13.0 software. GeneticdataanalysisSupposethatunderpolygeneand environmentmodifications,theeffectsofmajorgenesinseg— regationgenerationdisplayanindependentnOrmaldistribu- tion.Thus,theentiresegregationgenerationcanbeseenas amixeddistributionmadeupofmanyindependentnormaldis- tributionsTheiointanalysisofP,P,.andRlLswasconduc- ted-sj.Thegeneticmodelwiththebestfitwaschosenac- cordingtoAkaike'sinfOrmationcriterion(AIC).EquaIdistri— bution(,and),aswellasSmirnov(仃)andKol? mogorov(Dn1testswereappliedtodeterminegoodness—of- fit.TheoptimaImode1wasthatforstat;sticaIsignificanceat 30 25 20 l5 10 5 0 40 3O 2O l0 0 _一 A 一IIIlI》252015105 O AgriculturalScience&TechnologyVo1.12,No.7,2011 least.Correspondinggeneticparameterswereestimatedby leastsquares[一. ResultsandAnalysis Descriptivestatisticsshowedvariationoffatandfattyacid contentintheparentsandRILs(Table1).AIltraitsinRIL populationshowedawiderangeofvariation.andagreatnum— berofIinesexceedingtheparentswerefound.T-testindica— tedbetweenmid-parentandIinesexceedingtheparents'sig- nificantdifference.Phenotypicdataon215RILsshowednear normaJdistributionforalItraits.Thecoefficientofvariation (Cestimatedforfatandfattyacidcontentwere3.55%一 39.98%.TheANOVAresultsshowedthattheIinesoffatand fattyacidcontentaresignificantlydifferentandsuitableforge— neticanalysis(Table2).Thefrequencydistributionofpheno- typesshowskewedandmulti-peakphenomena(Fig.1),indi- catingthattheremavbemajorgenespresent.Threealterna- tivemodelsforthegenetictraitswereselected;theselection wasbasedontheIineseparationandanalysismethodoffat andfattyacidcompositionbetweenthevariouspossiblege- neticmodelsforthemaximumlikelihoodratiotest(datanot shown,.Theoptimummodelwasselectedaccordingtothe AICcriteriaandasetoftests(Table3).Geneticparameters wereestimatedbasedonthechosenmodel(Table4). 424446485O52543439444954252933374145n71.01.3lJ6L92.0 \E … 5O 分4O 喜o 星20 1O {.00l_15L301.451.60L75L909.410.4tt.4t2.413.41.92.22.52.83.13.4 A:Fat;B:Oleic;C:Linoleic;D:O/L;E:Stearic;F:Arachidic;G:Palmitic;H:Behenic. Fig.1FrequencydistributionofFatandFattyAcidcontentinRILs Table1ThevariationparametersinparentsandRILpopulationoffatandfattyacidcompositio ns Table2AnalysisofvarianceoffatandfattyacidcompositionsinRIL population Geneticanalysisoffatcontent AccordingtotheAICvalueandthegoodness-of-仇test. theoptimaIgeneticmodeIofthefatcontentwasE-2-7.in whichthereweretwopairsofcomplementaryIinkagesofthe additivegeneticmodelofmajorgenepluspolygene.Thecom- binedmainadditivegeneeffectandadditive×additiveinterac- tioneffectwas一1.82.Theheritabilityofmajorgeneswas 22.88%,whilethepolygenicheritabilitywas47.92%.indica- tingthatthegenetictraitmightbeaffectedbyenvironmentaI factors.Theseresultsweresimilartotheadditivegenetic u0口H 00盯.J0u口0_I ZHANGXin-youeta1.InheritanceofFatandFattyAcidCompositionsinPeanut(Arachishy pogaeaL.) effectsI.andadditivexadditiveinteractioneffectsilo1offat obtainedbyotherresearchers. Geneticanalysisoffattyacids 0leicacidAccordingtotheAICvalueandthegoodness.of- fittest.thebestgeneticmodelofoleicacidcontentwasB-1- 2.inwhicholeicacidcontentwascontrolledbytwopairsofin— dependentadditivemajorgenes.Theadditiveeffectofthefirst majorgenewas-4.21andthatofthesecondmajorgenewas 1.63.TheheritabiIityofthemajorgeneswas75.11%:noeffect wasobservedinmultiplegenes.Thegeneticanalysisconduc- tedbyLopezeta1.alsoshowedthathigholeicacidgenetic contentwascontrolledbytwomajorgenesJ. LinoleicacidAccordingtotheAICvalueandthegoodness. of-fittest.thebestgeneticmodeIoflinoleicacidcontentwas B-1-2.inwhichlinoleicacidcontentwasaffectedbytwopairs ofindependentgeneticadditivemajorgenes.Theadditive effectofthefirstmajorgenewas3.59andthatofthesecond Table3AICvaluesandtestforgoodness-?of??fitofoptimummodels majorgenewas一1.52.Theheritabilofthemajorgenes was75%:nopolygeneeffectwasobserved. O/LratioAccordingtotheAlCvalueandthegoodness—of- fittest,thebestgeneticmodeIofO/LratiowasF-3,that was.themodeIofthethreepairsofindependentequaIaddi- tivemajorgenes.Thethreepairsofmajorgenesshowede— qualadditivegeneticeffects.lnwhichtheadditiveeffectwas O.30.Theheritabilityofthemajorgenewas70.65%:no effectwasobservedfrompolygenes. StearicacidAccordingtotheAICvalueandthegoodness- of-fittest.thebestgeneticmodeJofstearicacidcontentwas E-14.thatwas,themixedmodeIofthetwopairsofmajor geneswithadditive—dominanceepistasispluspolYgenes.The additiveeffectofthefirstmajorgenewas一0.16andthatof thesecondmajorgenewas0.28.Theheritabilityofthemajor genewas26.43%.whereasthatofpolYgeneswas11.46%. ArachidicacidAccordingtotheAJCvalueandthegood. ness-of-fittest.thebestgeneticmodeIofarachidicacidcon- tentwasE-1-8,thatis,themajorgeneplusthepolygenicge- neticmode1withtwopairsofindependentoverlappingadditive roles.Thecombinedmainadditiveeffectandadditivexaddi. tiveinteractioneffectwas一0.10.Theheritabilityofthemaior genewas33.17%.whereasthatofpolygeneswas'9.64%. PalmlUcacidAccordingtothe,Cvalueandthegoodness. 0f-fittest.thebestgeneticmodeIofpalmiticacidcontentwas B-1-3,thatwas,thetwopairsofindependentmajorgeneswith equaladditiveeffect.Theadditiveeffectofthetwomajorgenes was0.43.Theheritabilityofmajorgeneswas60.95%:no effectwasobservedfrompolygenes BehenicacidAccordingtotheAICvalueandthegoodness一 0f—fi【test.thebestgeneticmodeIofbehenicacidwasF一3.in whichtherewerethreepairsofIndependentmajorgeneswith equaIadditivegeneticeffects.Thethreepairsofmaiorgenes showedanequaIadditiveeffectof一0.17.Theheritabilityof maiorgeneswas30.7O%:nOpolygeneeffectwasobserved. UlSCUSSIon Otherresearchershavecarriedoutfruiffulstudieson peanutgeneticsoffatcontent.Previousstudiessuggestthat thetraitsaremainlycontrolledbyadditivegeneeffects一' oracodecisionoftheadditiveandnon—additiveeffects. However.theadditiveeffectplaysadominantrolelJ.Our resultsweresimilarwiththoseofpreviousstudies,exceptfor theIowhereditabilityofmajorgenesandtheabsenceofpoly— geneeffects.indicatingthatfatcontentmighthavemore complexgeneticmechanismsandwasaffectedbyenviron— mentalfactors. OleicandJinoleicacidwerethemostjmportantfattyacids andmakeuD80%ofthetotalfattyacidcontentinpeanut,fOl_ Iowedbypalmiticstearic,andarachidicacid.Thisstudy foundthatoleicandlinoleicacidwerecontrolledbytwopairs ofadditivemajorgenesOtherresearchershavereposedthat oleicacidiscontrolledbytworecessivegenes[.,'一.]The researchofWaneta1.showedthatoleicacidandlinoleicacid werecontrolledbyaconsiderableadditiveeffect,jrlwhichthe dominanteffectwasminimaIandpartofthedominant.The recessiveallelesofoleicacidperformefficiently:thedominant allelesoflinoleicacidperformefficiently.aswelI.Inaddition. Knaufteta1.reportedthecaseofsinglegeneinheritance. Thedifferencejnresultsmaybeduetothedifferentanalysis methodsandgeneticmaterialsused.However,thissuggests thatthegeneticmechanismsofoleicandIinoleicacidarenot complicatedandrelativelyeasytoexploitforfutureinvestiga— tionsonthegeneticimprovementofpeanut. IngeneraI.unsaturatedfattyacidsarebeneficiaIto health.buttheincreaseinunsaturatedbondsalsoincreases lheriskofdeteriorationbecauseoftheoxidationjnpeanutand itsproducts.therebyreducingItsshelfIire.OleicandIinoleic acidareunsaturatedfattyacidswithoneandtwounsaturated bonds,respectively.LinoleicacidisanessentiaIfattyacid. whichcannotbesynthesizedbythebodyandmustbeinclud— edjndiets,丁wounsaturatedbonds.whicharemoreproneto oxidativerancidity.reducetheshelflireofpeanutproducts. 0/Lratioisanimportantbiochemicafindicatorforpeanutstor- age.AhigherO/Lratiomeansbetterstorabilityandlonger shelfIire.Nordeneta1.examined494differenttypesofpea- nutandfoundtwoO/Lratiovaluesof40.indicatingthatthe storabilityofthepeanutsiSsuperiortothatofothergeno— typesl"J.Alpresent.increasedoleicacidcontentorO/Lval— ueshasbecomeoneoftheimportantgoalsfortheimprove— mentofpeanutquality ThegeneticmodeIofO/Lratioisconsistentwiththecon— clusionsoftheO/Lratiotraitforthe"additive—dominance" modeIestablishedbyWaneta1._1.jnwhichnoepistasis Responsibleeditor:LITing-ting AgriculturalScience&TechnologyVo1.12,No.7,2011 effectandcOnsiderableadditiveeffectareobserved.O/Lratio iscontrolledbythethreepairsofmajorgenes,whichdemon— stratethetraitsinfluencedbyothermodifiergenes.exceptthe twomajorgenesl.Inaddition.thestudyofthematerials with{owO/Lshowsthatinheritancemayhavemorecomplex mechanisms. Ourresultsareconsistentwithpreviousstudiesonthein. heritanceofstearicandarachidicacid',Thesestudiesshow thatthesixtraitsofoleicandIinoleicacid.O/Lratiostearic acid.palmiticacid,andbehenicacidcontentareadvantages ofadditiveeffects.whichisbeneficialforqualityimprovement inpeanutthroughcrossbreedingmethods. References n1FANGJH.CHANGRZEconomiccropsjncropsandtheirwiIdrel— ativesinChina[M].Beijing:ChinaAgriculturePress,2007:126— 166. 『21KNAUFTDA,M?REKM,GORBETDW.Fu~herstudiesonthe jnheritaneeoffatryacidcompositionjnpeanutfJJ.PeanutScj, 1993(2O):74—76. 『3lMO0REKM.KNAUF.rDA,Theinhentanceofhigholeicacidin peanutlJ1.Heredity.1989(80):252—2&3 I4;OKEEFESF.WILEYVA,KNAUFTDA.Comparisonofoxidative stabilityofhighandnormaIoleicpeanutoilsIJ].AmOIlChem. Soc,1993(7O):489—4g2. I5IZHANGYM,GAlJYldentificationofmixedmajorgenesandpoly genesinheritaricemodelofquantitativetraitsbyusingDHorRIL populationlJ].ActaGenetSin,2000(27):634—640. I6IGAIJY,ZHANGYM,\/vANGJ?