Nitric Oxide in Plant Physiology Edited by Shamsul Hayat, Masaki Mori, John Pichtel, and Aqil Ahmad Related Titles Hirt,H. (ed.) Plant Stress Biology FromGenomicstoSystemsBiology 2010 ISBN:978-3-527-32290-9 Kahl, G.,Meksem,K.(eds.) The Handbook of Plant Functional Genomics ConceptsandProtocols 2008 ISBN:978-3-527-31885-8 Ahmad, A.,Pichtel, J.,Hayat, S.(eds.) Plant-Bacteria Interactions StrategiesandTechniquestoPromotePlantGrowth 2008 ISBN:978-3-527-31901-5 Scott,P. Physiology and Behaviour of Plants 2008 ISBN:978-0-470-85025-1 Buchanan, B.,Gruissem, W.,Jones,R. L. (eds.) Biochemistry & Molecular Biology of Plants 2002 ISBN:978-0-943088-39-6 Nitric Oxide in Plant Physiology Edited by Shamsul Hayat, Masaki Mori, John Pichtel, and Aqil Ahmad TheEditors AllbookspublishedbyWiley-VCHarecarefully produced.Nevertheless,authors,editors,and Dr.ShamsulHayat publisherdonotwarranttheinformationcontained AligarhMuslimUniversity inthesebooks,includingthisbook,tobefreeof DepartmentofBotany errors.Readersareadvisedtokeepinmindthat Aligarh202002 statements,data,illustrations,proceduraldetailsor India otheritemsmayinadvertentlybeinaccurate. Dr.MasakiMori LibraryofCongressCardNo.: appliedfor NationalInstituteofAgrobiologicalSciences DivisionofPlantSciences BritishLibraryCataloguing-in-PublicationData 2-1-2Kannondai,Tskuba Acataloguerecordforthisbookisavailablefromthe Ibaraki305-8602 BritishLibrary. Japan Bibliographicinformationpublishedby Prof.JohnPichtel theDeutscheNationalbibliothek BallStateUniversity TheDeutscheNationalbibliothekliststhis DepartmentofNaturalResourcesand publicationintheDeutscheNationalbibliografie; EnvironmentalManagement detailedbibliographicdataareavailableonthe WQ103 Internetathttp://dnb.d-nb.de. Muncie,IN47306 USA #2010WILEY-VCHVerlagGmbH&Co.KGaA, Weinheim Dr.AqilAhmad HigherCollegeofTechnology Allrightsreserved(includingthoseoftranslationinto DepartmentofAppliedSciences otherlanguages).Nopartofthisbookmaybe Al-Khuwair reproducedinanyform–byphotoprinting, SultanateofOman microfilm,oranyothermeans–nortransmittedor Oman translatedintoamachinelanguagewithoutwritten and permissionfromthepublishers.Registerednames, AligarhMuslimUniversity trademarks,etc.usedinthisbook,evenwhennot DepartmentofBotany specificallymarkedassuch,arenottobeconsidered Aligarh202002 unprotectedbylaw. India Cover Formgeber,Eppelheim Typesetting ThomsonDigital,Noida,India PrintingandBinding StraussGmbh,Mörlenbach PrintedintheFederalRepublicofGermany Printedonacid-freepaper ISBN:978-3-527-32519-1 V Contents Preface XI List of Contributors XIII 1 NitricOxide:Chemistry,Biosynthesis,andPhysiologicalRole 1 ShamsulHayat,SyedAimanHasan,MasakiMori,QaziFariduddin, andAqilAhmad 1.1 Introduction 1 1.2 NitricOxideChemistry 2 1.3 BiosynthesisofNitricOxide 3 1.4 PhysiologicalRoleofNitricOxide 5 1.4.1 EffectofNitricOxideonSeedDormancy 5 1.4.2 EffectofNitricOxideonGrowth 6 1.4.3 EffectofNitricOxideonSenescence 6 1.4.4 EffectofNitricOxideonNitrateReductaseActivity 7 1.4.5 EffectofNitricOxideonRespiration 7 1.4.6 EffectofNitricOxideonStomatalMovement 7 1.4.7 EffectofNitricOxideonChlorophyllContent 7 1.4.8 EffectofNitricOxideonPhotosynthesis 8 1.4.9 EffectofNitricOxideonAntioxidantSystem 8 1.4.10 EffectofNitricOxideonProgrammedCellDeath 9 1.5 NitricOxideandCrossTalkwithClassicalPlantHormones 10 1.5.1 AuxinsandNitricOxide 10 1.5.2 AbscisicAcidandNitricOxide 11 1.5.3 Cytokinins,Gibberellins,andNitricOxide 11 1.5.4 EthyleneandNitricOxide 12 References 12 2 ElectronParamagneticResonanceasaTooltoStudyNitric OxideGenerationinPlants 17 SusanaPuntarulo,SebastiánJasid,AlejandroD.Boveris, andMarcelaSimontacchi 2.1 Introduction 17 VI Contents 2.1.1 ChemistryofNitrogen-ActiveSpecies 17 2.1.2 BiologicalEffectsofNO 18 2.2 MethodsofNODetection 19 2.2.1 DeterminationofNObySpecificElectrodes 19 2.2.2 DeterminationofNObySpectrophotometric andFluorometricMethods 19 2.2.3 DeterminationofNObyElectronParamagneticResonance 20 2.2.3.1 SpecificExperimentalAdvances 20 2.3 UseofEPRMethodologyforAssayingEnzymeActivities 22 2.3.1 NOS-LikeDependentNOGeneration 24 2.3.2 NitrateReductase-DependentNOGeneration 24 2.4 ApplicationofEPRMethodstoAssessNOGenerationDuring PlantDevelopment 26 2.5 Conclusions 27 References 27 3 Calcium,NO,andcGMPSignalinginPlantCellPolarity 31 AnaMargaridaPrado,JoséA.Feijó,andDavidMarshallPorterfield 3.1 Introduction 31 3.2 CellPolarityandPlantGametophyteDevelopment 33 3.3 CalciumSignalinginPollenandFernSpores 34 3.4 NO/cGMPSignalinginPollenandFernSpores 35 3.5 NO/cGMPinPollen–PistilInteractions 38 3.6 OvuleTargetingandNO/cGMP 39 3.7 Ca2þ/NO/cGMPConnection? 42 3.8 ClosingPerspectives 46 References 48 4 NitricOxideandAbioticStressinHigherPlants 51 FranciscoJ.Corpas,JoséM.Palma,MarinaLeterrier,LuisA.delRío, andJuanB.Barroso 4.1 Introduction 51 4.2 NitricOxideandRelatedMolecules 52 4.2.1 ChemistryofNitricOxideinPlantCells 52 4.2.2 ReactiveNitrogenSpecies 52 4.3 CellularTargetsofNO 54 4.3.1 NitrosylatedMetals 54 4.3.2 ProteinS-Nitrosylation 55 4.3.3 ProteinTyrosineNitration 55 4.3.4 Nitrolipids 55 4.3.5 NucleicAcidNitration 56 4.3.6 NOandGeneRegulation 56 4.4 FunctionsofNOinPlantAbioticStress 57 4.4.1 Salinity 57 Contents VII 4.4.2 UltravioletRadiation 58 4.4.3 Ozone 58 4.4.4 MechanicalWounding 59 4.4.5 ToxicMetals(CadmiumandAluminum) 59 4.5 ConcludingRemarks 60 References 61 5 PolyaminesandCytokinin:IsNitricOxideBiosynthesisthe KeytoOverlappingFunctions? 65 RinukshiWimalasekeraandGüntherF.E.Scherer 5.1 Introduction 65 5.2 Cytokinin-andPolyamine-InducedNOBiosynthesis 66 5.3 TissueDistributionofZeatin-InducedandPA-Induced NOFormation 67 5.4 NitricOxide,Cytokinin,andPolyaminesinPlantGrowthand DevelopmentandinAbioticandBioticStresses 68 5.4.1 Embryogenesis 68 5.4.2 Flowering 69 5.4.3 Senescence 69 5.4.4 ProgrammedCellDeath 69 5.4.5 AbioticStresses 70 5.4.6 BioticStresses 71 References 73 6 RoleofNitricOxideinProgrammedCellDeath 77 MichelaZottini,AlexCosta,RobertoDeMichele,andFiorellaLoSchiavo 6.1 ProgrammedCellDeathinPlants 77 6.1.1 PCDHallmarksandRegulation 78 6.2 NOasaSignalingMolecule 79 6.2.1 NOIsAbletoInduceorInhibitPCD 79 6.2.2 NitricOxideandPCDinHypersensitiveResponse 80 6.2.3 SignalingComponentinSA-InducedNOProduction 80 6.3 RoleofMitochondriainNO-InducedPCD 84 6.4 Conclusions 85 References 85 7 NitrateReductase-DeficientPlants:AModeltoStudyNitric OxideProductionandSignalinginPlantDefenseResponse toPathogenAttack 89 IoneSalgado,HalleyCaixetadeOliveira,andMarciaReginaBraga 7.1 Introduction 89 7.2 PhysicochemicalBasisofNOSignaling 91 7.3 DefenseResponsesMediatedbyNO 92 7.3.1 AccumulationofDefensiveCompounds 92 VIII Contents 7.3.2 HypersensitiveResponse 93 7.3.3 SystemicResponses 94 7.3.4 StomatalClosure 94 7.4 SubstratesforNOProductionDuringPlant–Pathogen Interactions 95 7.4.1 ProductionofNOfromL-Arginine 95 7.4.2 ProductionofNOfromNitrite 95 7.5 TheRoleofNitrateReductaseinNOProductionDuring Plant–PathogenInteractions 97 References 98 8 EffectivePlantProtectionWeaponsAgainstPathogensRequire ‘‘NOBullets’’ 103 LuziaV.Modolo 8.1 Introduction 103 8.2 NitricOxideandReactiveOxygenSpeciesinthe HypersensitiveResponse 104 8.3 NitricOxideandPhytoalexinProduction 107 8.4 NitricOxideandtheSalicylicAcidSignalingPathway 108 8.5 NitricOxideandtheJasmonicAcidSignalingPathway 109 8.6 NitricOxideandGeneRegulation 109 8.7 NitricOxideandProteinRegulation 110 8.8 ConcludingRemarks 111 References 111 9 TheRoleofNitricOxideasaBioactiveSignalingMoleculein PlantsUnderAbioticStress 115 Gang-PingHaoandJian-HuaZhang 9.1 Introduction 116 9.2 BiosynthesisofNitricOxideUnderAbioticStress 116 9.2.1 NOGeneratedfromNOS-LikeActivityUnderAbioticStress 116 9.2.2 NOGeneratedfromNRUnderAbioticStress 120 9.3 NOSignalingFunctionsinAbioticStressResponses 121 9.3.1 FunctionofNOUnderDroughtStress 122 9.3.2 FunctionofNOUnderSaltStress 123 9.3.3 FunctionofNOUnderUltravioletRadiation 125 9.3.4 FunctionofNOUnderHeatandLowTemperature 126 9.3.5 FunctionofNOUnderHeavyMetalStress 126 9.3.6 FunctionofNOUnderOtherAbioticStresses 127 9.4 NOSignalTransductioninPlantsUnderAbioticStress 128 9.4.1 cGMP-DependentSignaling 128 9.4.2 DownstreamSignalingforNOAction 129 9.5 InteractionsofNOSignalingwithOtherSignalingMoleculesin PlantResponsetoAbioticStress 131 References 135