Gene networks involved in drought stress response and tolerance

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JournalofExperimentalBotany,Vol.58,No.2,pp.221–227,2007IntegratedApproachestoSustainandImprovePlantProductionunderDroughtStressSpecialIssuedoi:10.1093/jxb/erl164AdvanceAccesspublication30October,2006

SPECIALISSUEPAPERGenenetworksinvolvedindroughtstressresponseand

tolerance

KazuoShinozaki1,*andKazukoYamaguchi-Shinozaki2,31RIKENPlantScienceCenter,1-7-22Suehiro-cho,Tsurumi-ku,Yokohama,203-0045Japan

2LaboratoryofPlantMolecularPhysiology,GraduateSchoolofAgriculturalandLifeSciences,TheUniversityof

Tokyo,1-1Yayoi,Bunkyo-ku,Tokyo,113-8657Japan3BiologicalResourcesDivision,JapanInternationalResearchCenterforAgriculturalSciences(JIRCAS),

1-1Ohwashi,Tsukuba,Ibaraki,305-8686Japan

Received10December2005;Accepted22August2006

AbstractPlantsrespondtosurviveunderwater-deficitconditionsviaaseriesofphysiological,cellular,andmolecularprocessesculminatinginstresstolerance.Manydrought-induciblegeneswithvariousfunctionshavebeenidentifiedbymolecularandgenomicanalysesinArabidopsis,rice,andotherplants,includinganumberoftranscriptionfactorsthatregulatestress-induciblegeneexpression.Theproductsofstress-induciblegenesfunctionbothintheinitialstressresponseandinestablishingplantstresstolerance.Inthisshortreview,recentprogressresultingfromanalysisofgeneexpressionduringthedrought-stressresponseinplantsaswellasinelucidatingthefunctionsofgenesimplicatedinthestressresponseand/orstresstoler-ancearesummarized.AdescriptionisalsoprovidedofhowvariousgenesinvolvedinstresstolerancewereappliedingeneticengineeringofdehydrationstresstoleranceintransgenicArabidopsisplants.

Keywords:Droughtstress,geneexpression,microarray,stresstolerance,molecularbreeding.

IntroductionDroughtstressinducesarangeofphysiologicalandbio-chemicalresponsesinplants.Theseresponsesincludestomatalclosure,repressionofcellgrowthandphotosynthe-

sis,andactivationofrespiration.Plantsalsorespondandadapttowaterdeficitatboththecellularandmolecularlevels,forinstancebytheaccumulationofosmolytesandproteinsspecificallyinvolvedinstresstolerance.Anassortmentofgeneswithdiversefunctionsareinducedorrepressedbythesestresses(Shinozakietal.,2003;BartelsandSunkar,2005;Yamaguchi-ShinozakiandShinozaki,2005).Mostoftheirgeneproductsmayfunctioninstressresponseandtoleranceatthecellularlevel.Significantly,theintroductionofmanystress-induciblegenesviagenetransferresultedinimprovedplantstresstolerance(Zhangetal.,2004;Umezawaetal.,2006a).Recently,anumberofstress-induciblegeneshavebeenidentifiedusingmicroarrayanalysisinvariousplantspecies,suchasArabidopsisandrice.Now,analysingthefunctionsofthesegenesiscriticaltofurtherourunder-standingofthemolecularmechanismsgoverningplantstressresponseandtolerance,ultimatelyleadingtoenhancementofstresstoleranceincropsthroughgeneticmanipulation.Droughttriggerstheproductionofthephytohormoneabscisicacid(ABA),whichinturncausesstomatalclosureandinducesexpressionofstress-relatedgenes.Severaldrought-induciblegenesareinducedbyexogenousABAtreatment,whereasothersarenotaffected.Indeed,evidenceexistsdemonstratingthepresenceofbothABA-indepen-dentandABA-dependentregulatorysystemsgoverningdrought-induciblegeneexpression(Yamaguchi-ShinozakiandShinozaki,2005).Bothcis-actingandtrans-actingre-gulatoryelementsfunctioninginABA-independentand/or

*Towhomcorrespondenceshouldbeaddressed.E-mail:sinozaki@rtc.riken.jpAbbreviations:ABA,abscisicacid;ABRE,abscisicacid-responsiveelement;DRE,dehydration-responsiveelement;EST,expressedsequencetag;JA,jasmonicacid;LEA,lateembryogenesisabundant.

ªTheAuthor[2006].PublishedbyOxfordUniversityPress[onbehalfoftheSocietyforExperimentalBiology].Allrightsreserved.ForPermissions,pleasee-mail:journals.permissions@oxfordjournals.org

by guest on October 25, 2010jxb.oxfordjournals.orgDownloaded from ABA-responsivegeneexpressioninducedbydroughtstresshavebeenpreciselyanalysedatthemolecularlevel(Yamaguchi-ShinozakiandShinozaki,2005).Thisshortarticlesummarizesrecentresultsobtainedthroughtran-scriptomeanalysisofdrought-induciblegeneexpressioninArabidopsisandriceusingmicroarrays,includinginforma-tionsupportingpotentialfunctionsofdrought-induciblegenesinstressresponseandtolerance.Examplesarealsocitedofhowdrought-induciblegenesareutilizedinimprovementofdrought-stresstolerancebygenetransfer.Thespecificfunctionsofthesegenesinstresstolerancearealsodiscussed,mainlyinArabidopsis.

Transcriptomeanalysisofdrought-induciblegenesinArabidopsisandriceusingmicroarrays

MicroarraytechnologyemployingcDNAsoroligonucleo-tidesisapowerfultoolforanalysinggeneexpressionprofilesofplantsexposedtoabioticstressessuchasdrought,highsalinity,orcold,ortoABAtreatment(Sekietal.,2001,2002a,b;Krebsetal.,2002).Therearetwopredominantvarietiesofmicroarraytechnologyavailable,thecDNAmicroarray(Sekietal.,2001,2002a,b)andtheoligonucleo-tidemicroarray,themostprominentbeingtheAffimetrixGeneChip(Krepsetal.,2002).A7000full-lengthcDNAmicroarraywasutilizedtoidentify299drought-induciblegenes,54cold-induciblegenes,213highsalinity-induciblegenes,and245ABA-induciblegenesinArabidopsis(Sekietal.,2002a,b).Morethanhalfofthesedrought-induciblegeneswerealsoinducedbyhighsalinityand/orABAtreatments,implicatingsignificantcross-talkbetweenthedrought,highsalinity,andABAresponsepathways.Incontrast,only10%ofthedrought-induciblegeneswerealsoinducedbycoldstress.Thousandsofstress-induciblegeneswereidentifiedusingtheAffimetrixGeneChiparraycontainingoligonucleotidesrepresenting~8000independentArabidopsisgenes(Krepsetal.,2002).Thestress-induciblegenesidentifiedfromthecDNAmicroarrayanalysisdidnotcoincidewiththoserecognizedthroughGeneChipanalyses.Thisdiscrepancyinresultsisprimarilyduetodifferencesbetweenthesetsofgenesarrayedinthetwosystems(only1919geneswerearrayedinbothsystems),aswellasdisparitiesinplantgrowthandstresstreatmentconditions(Maruyamaetal.,2004).Recently,theAtGenExpressprojectassayedtheArabidopsistranscriptomeusingtheAffymetrix23000ATH1GeneChip,generatingthousandsoftran-scriptomedatapointsidentifyinggenesexpressedinvarioustissuesandunderdefinedgrowthconditions,stressinduction,andphytohormonetreatment(Schmidetal.,2005).RicegeneexpressioninresponsetohighsalinitystresswasfirstanalysedusingcDNA[expressedsequencetag(EST)]microarraytechnology(Kawasakietal.,2001).Recently,stress-induciblegenesinricewereanalysedusingamicroarrayincluding~1700independentricecDNAs(ESTs).ThesecDNAswereharvestedfromriceplants