Transcriptional Regulatory Networks in Cellular Re

ANRV274-PP57-30ARI5April200619:25TranscriptionalRegulatoryNetworksinCellularResponsesandTolerancetoDehydrationandColdStressesKazukoYamaguchi-Shinozaki1,2,4andKazuoShinozaki3,41LaboratoryofPlantMolecularPhysiology,GraduateSchoolofAgriculturalandLifeSciences,UniversityofTokyo,Tokyo113-8657,Japan;email:[e-mail]2BiologicalResourcesDivision,JapanInternationalResearchCenterforAgriculturalSciences(JIRCAS),Ibaraki305-8686,Japan3RIKENPlantScienceCenter,Yokohama203-0045,Japan4CREST,JapanScienceandTechnologyCorporation(JST),JapanAnnu.Rev.PlantBiol.2006.57:781–803TheAnnualReviewofPlantBiologyisonlineatplant.annualreviews.orgdoi:10.1146/annurev.arplant.57.032905.105444Copyrightc2006byAnnualReviews.AllrightsreservedFirstpublishedonlineasaReviewinAdvanceonMarch3,20061543-5008/06/0602-0781$20.00KeyWordsgeneexpression,cis-element,transcriptionfactor,stresstoleranceAbstractPlantgrowthandproductivityaregreatlyaffectedbyenvironmentalstressessuchasdrought,highsalinity,andlowtemperature.Expres-sionofavarietyofgenesisinducedbythesestressesinvariousplants.Theproductsofthesegenesfunctionnotonlyinstresstolerancebutalsoinstressresponse.Inthesignaltransductionnetworkfromper-ceptionofstresssignalstostress-responsivegeneexpression,varioustranscriptionfactorsandcis-actingelementsinthestress-responsivepromotersfunctionforplantadaptationtoenvironmentalstresses.Recentprogresshasbeenmadeinanalyzingthecomplexcascadesofgeneexpressionindroughtandcoldstressresponses,especiallyinidentifyingspecificityandcrosstalkinstresssignaling.Inthisreviewarticle,wehighlighttranscriptionalregulationofgeneex-pressioninresponsetodroughtandcoldstresses,withparticularemphasisontheroleoftranscriptionfactorsandcis-actingelementsinstress-induciblepromoters.781Annu.Rev.PlantBiol.2006.57:781-803.Downloadedfromarjournals.annualreviews.orgbyUniversityofBritishColumbiaLibraryon05/11/10.Forpersonaluseonly.ANRV274-PP57-30ARI5April200619:25ContentsINTRODUCTION.................782FUNCTIONOFOSMOTIC-ANDCOLD-INDUCIBLEGENES...783REGULATIONOFGENEEXPRESSIONBYOSMOTICANDCOLDSTRESSES........783AMAJORABA-INDEPENDENTGENEEXPRESSIONUNDEROSMOTICANDCOLDSTRESSES......................784DREB1/CBFs:MajorTranscriptionFactorsthatRegulateManyCold-InducibleGenesInvolvedinStressTolerance....................785TheDREB/DRERegulonsinPlantsOtherThanArabidopsis..787Cis-ActingRegulatoryElementsandTranscriptionFactorsthatFunctionUpstreamofCBF3/DREB1A...............787TheDREB2ProteinsFunctioninOsmoticStress-ResponsiveGeneExpression..............788OTHERABA-INDEPENDENTGENEEXPRESSIONUNDEROSMOTICANDCOLDSTRESSES.....................788AMAJORABA-RESPONSIVEGENEEXPRESSIONUNDEROSMOTICSTRESSES.........789OTHERTYPESOFABA-DEPENDENTGENEEXPRESSIONUNDERDEHYDRATIONSTRESS.....791REGULATIONOFABABIOSYNTHESISANDDEGRADATIONDURINGDEHYDRATIONANDREHYDRATION................792REGULATIONOFGENEEXPRESSIONDURINGREHYDRATIONAFTERDEHYDRATION...............792GENETICANALYSISOFSIGNALTRANSDUCTIONINRESPONSETODEHYDRATIONANDCOLDSTRESSES......................793CONCLUSIONSANDFUTUREPERSPECTIVES................795INTRODUCTIONDrought,highsalinity,andlowtemperatureareallenvironmentalconditionsthathaveanadverseeffectonthegrowthofplantsandtheproductivityofcrops.Plantshaveadaptedtorespondtothesestressesatthemolecularandcellularlevelsaswellasatthephysiologicalandbiochemicallevels,thusenablingthemtosurvive.Expressionofavarietyofgenesisin-ducedbythesestressesinvariousplants(39,103,114).Theproductsofthesegenesfunc-tionnotonlyinstresstolerancebutalsointheregulationofgeneexpressionandsignaltransductioninstressresponses(5,104,125).Abscisicacid(ABA)isproducedunderwater-deficitconditionsandplaysanimpor-tantroleinthetoleranceresponseofplantstodroughtandhighsalinity.Exogenousapplica-tionofABAalsoinducesanumberofgenesthatrespondtodehydrationandcoldstress(104,136).Nevertheless,theroleofABAincoldstress-responsivegeneexpressionisnotclear.SeveralreportshavedescribedgenesthatareinducedbydehydrationandcoldstressesbutthatdonotrespondtoexogenousABAtreatment(104,132,136).ThissuggeststheexistenceofABA-independent,aswellasABA-dependent,signaltransductioncascadesbetweentheinitialstresssignalandtheex-pressionofspecificgenes.Themolecularmechanismsregulatinggeneexpressioninresponsetodehydrationandcoldstresses782Yamaguchi-Shinozaki·ShinozakiAnnu.Rev.PlantBiol.2006.57:781-803.Downloadedfromarjournals.annualreviews.orgbyUniversityofBritishColumbiaLibraryon05/11/10.Forpersonaluseonly.ANRV274-PP57-30ARI5April200619:25havebeenstudiedbyanalyzingthecis-andtrans-actingelementsthatfunctioninABA-independentandABA-responsivegeneex-pressionduringthestressesinArabidopsis(104,132).Inthisreviewarticle,wefocusontran-scriptionalregulationofgeneexpressioninresponsetodehydrationandcoldstresses,withparticularemphasisontheroleoftran-scriptionfactorsandcis-actingelementsinstress-induciblepromoters.Thesignaltrans-ductionpathwayscontrollingabioticstressre-sponsesareverycomplex,andmanyexcellentreviewarticlesinthisareawererecentlypub-lished(5,9,15,126).FUNCTIONOFOSMOTIC-ANDCOLD-INDUCIBL