Light and Plant Development Editedby GARRYC.WHITELAM DepartmentofBotany SchoolofBiologicalSciences UniversityofLeicester Leicester UK and KARENJ.HALLIDAY SchoolofBiologicalSciences TheUniversityofEdinburgh Edinburgh UK Light and Plant Development Light and Plant Development Editedby GARRYC.WHITELAM DepartmentofBotany SchoolofBiologicalSciences UniversityofLeicester Leicester UK and KARENJ.HALLIDAY SchoolofBiologicalSciences TheUniversityofEdinburgh Edinburgh UK (cid:1)C 2007byBlackwellPublishingLtd EditorialOffices: BlackwellPublishingLtd,9600GarsingtonRoad,OxfordOX42DQ,UK Tel:+44(0)1865776868 BlackwellPublishingProfessional,2121StateAvenue,Ames,Iowa50014-8300,USA Tel:+15152920140 BlackwellPublishingAsiaPtyLtd,550SwanstonStreet,Carlton,Victoria3053,Australia Tel:+61(0)383591011 The right of the Author to be identified as the Author of this Work has been asserted in accordancewiththeCopyright,DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem, ortransmitted,inanyformorbyanymeans,electronic,mechanical,photocopying,recording orotherwise,exceptaspermittedbytheUKCopyright,DesignsandPatentsAct1988,without thepriorpermissionofthepublisher. Firstpublished2007byBlackwellPublishingLtd ISBN:978-1-4051-4538-1 LibraryofCongressCataloging-in-PublicationData Lightandplantdevelopment/editedbyGarryC.WhitelamandKarenJ.Halliday. p. cm. Includesbibliographicalreferencesandindex. ISBN:978-1-4051-4538-1(hardback:alk.paper) 1.Phytochrome. 2.Plants—Photomorphogenesis. 3.Plants—Development. I.Whitelam,GarryC. II.Halliday,KarenJ. 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ForfurtherinformationonBlackwellPublishing,visitourwebsite: www.blackwellpublishing.com Contents Contributors xiii Preface xv PartI Photoreceptors 1 1 Phytochromes 3 ANDREASHILTBRUNNER,FERENCNAGYAND EBERHARDSCHA¨FER 1.1 Introduction 3 1.2 Historicalaspects 3 1.3 PropertiesofphyAinvivo 4 1.4 Propertiesinyeastcells 4 1.5 Invivopropertiesofphytochromes 5 1.5.1 Invivospectroscopy 5 1.6 Intracellularlocalisationofphytochromes 7 1.6.1 Classicalmethods 7 1.6.2 Spectroscopicmethods 7 1.6.3 Cellbiologicalmethods 8 1.6.4 Immunocytochemicalmethods 8 1.6.5 Novelmethods 8 1.7 IntracellularlocalisationofphyBindarkandlight 9 1.8 IntracellularlocalisationofphyAindarkandlight 11 1.9 IntracellularlocalisationofphyC,phyDandphyEindark andlight 12 1.10 Phytochrome/PIF3co-localisationandnuclearspeckles 12 1.11 Regulationofintracellularlocalisationofphytochromes 13 Acknowledgements 14 References 15 2 Cryptochromes 17 ALFREDBATSCHAUER,ROOPABANERJEEAND RICHARDPOKORNY 2.1 Introduction 17 2.2 Cryptochromegenesandtheirevolution 17 2.3 Cryptochromedomains,chromophoresandstructure 20 2.3.1 Domainstructureofthecryptochromes 20 2.3.2 Cryptochromechromophores 21 vi CONTENTS 2.3.3 Photolyaseandcryptochromestructure 22 2.3.3.1 Photolyasestructureandreactionmechanism 22 2.3.3.2 Cryptochromestructure 23 2.4 Cryptochromebiochemistryandspectroscopy 25 2.4.1 Phosphorylation 25 2.4.2 Nucleotide-bindingandkinaseactivity 25 2.4.3 DNA-bindingactivity 26 2.4.4 Electrontransfer 27 2.5 Expressionandbiologicalactivityofcryptochromes 28 2.5.1 Expressionandlightregulationofcryptochromesin planta 28 2.5.2 Cellularlocalization 31 2.5.3 Growthresponsescontrolledbycryptochromes 31 2.5.4 Regulationofgeneexpressionthroughcryptochromes 33 2.6 Cryptochromesignalling 35 2.6.1 Dimerizationandoutputdomains 36 2.6.2 Cryptochromepartners 38 2.6.2.1 InteractionwithCOP1 39 2.6.2.2 Interactionwithzeitlupe/ADAGIO1 39 2.6.2.3 Interactionwithphytochromes 40 2.6.3 Furtherdownstreamcomponents 41 2.7 Summary 43 Acknowledgements 43 References 43 3 PhototropinsandotherLOV-containingproteins 49 JOHNM.CHRISTIE 3.1 Introduction 49 3.2 Phototropinsandtheirbiologicalfunctions 50 3.2.1 Physiologicalrolesinhigherplants 50 3.2.2 Physiologicalrolesinlowerplants 52 3.3 Phototropinstructure,localizationandactivity 53 3.3.1 Phototropinstructureandlocalization 53 3.3.2 Phototropinautophosphorylation 55 3.4 LightsensingbytheLOVdomains 57 3.4.1 LOV-domainphotochemistry 57 3.4.2 LOV-domainstructure 59 3.4.3 FunctionalrolesofLOV1andLOV2 60 3.4.4 Light-inducedproteinmovements 62 3.5 Phototropinsignaling 63 3.5.1 Phototropin-interactingproteins 63 3.5.2 Downstreamsignalingtargets 65 3.6 OtherLOV-containingproteins 67 3.6.1 LOV-containingproteinsinArabidopsis 67 3.6.2 LOV-containingproteinsinfungi 70 3.6.3 LOV-containingproteinsinbacteria 72 CONTENTS vii 3.7 Conclusionsandfutureperspectives 73 Acknowledgements 73 References 74 PartII Photoreceptorsignaltransduction 79 4 Phytochrome-interactingfactors 81 PETERH.QUAIL 4.1 Introduction 81 4.2 Methodology 81 4.2.1 InitialidentificationofPIFs 81 4.2.2 Subsequentassayandcharacterizationofthe interaction 82 4.2.3 Reversegeneticassessmentoffunctionalrelevanceto physignalling 83 4.3 phy`-interactors 84 4.3.1 PIF3 84 4.3.2 OtherbHLHtranscriptionfactorsandtheactivephyB bindingdomain 90 4.3.3 Nucleosidediphosphatekinase2 93 4.3.4 Phytochromekinasesubstrate1 94 4.3.5 Type5proteinphosphatase 94 4.3.6 Proteinphosphatase2A 95 4.3.7 Earlyflowering3 95 4.3.8 Far-redelongatedhypocotyl1 96 4.4 Pre-selectedinteractiontargets 96 4.4.1 Arabidopsisresponseregulator4 96 4.4.2 Zeitlupe 97 4.4.3 Cryptochrome1and2 97 4.4.4 AUX/IAAproteins 98 4.4.5 COP1 98 4.5 Perspective 99 Acknowledgements 100 References 101 5 Phosphorylation/dephosphorylationinphotoreceptorsignalling 106 CATHERINELILLO,TRUDIEALLENANDSIMONGEIR MØLLER 5.1 Introduction 106 5.1.1 Thephotoreceptorsautophosphorylate,butthe classicalactivationloopisnotinvolved 107 5.1.2 Phosphatasesinphotoreceptorsignalling 108 5.2 Phytochromes 109 5.2.1 Phosphorylationofphytochrome 109 5.2.2 Phytochromekinasesubstrate1 110 5.2.3 Nucleosidediphosphatekinase2 111 viii CONTENTS 5.2.4 FYPP 112 5.2.5 PAPP5 113 5.3 Cryptochromes 114 5.3.1 Cryptochromephosphorylation 114 5.3.2 PhosphorylationoftheC-terminalendisnecessaryfor signaltransduction 115 5.4 Phototropins 116 5.5 Isphosphorylation/dephosphorylationimportantfor downstreamevents? 118 5.5.1 HY5 118 5.5.2 Longhypocotylinfar-redlight 119 5.5.3 Circadianclock-associatedandlateelongated hypocotyl 119 5.5.4 EID1 120 5.5.5 Aux/IAA 120 5.5.6 PP7 121 5.5.7 Downstreamofphototropin 121 5.6 Conclusions 122 References 123 6 Theroleofubiquitin/proteasome-mediatedproteolysis inphotoreceptoraction 128 SUHUAFENGANDXINGWANGDENG 6.1 Introduction 128 6.2 Overviewoftheubiquitin/proteasomesystem 129 6.2.1 Ubiquitinconjugationanddeconjugationpathways 129 6.2.2 DiversityofE3Ubligases 130 6.2.3 26Sproteasome 131 6.3 RoleofCOP/DET/FUSproteinsinphotoreceptor-mediated signaltransductionandubiquitin/proteasome-mediated proteolysis 132 6.3.1 COP/DET/FUSproteinsintegratedivergent photoreceptorsignalingpathwaysanddownstream geneexpression 132 6.3.2 COP1 134 6.3.2.1 LightregulationofCOP1localization 135 6.3.2.2 COP1actsasanE3Ubligase 136 6.3.2.3 InteractionbetweenphotoreceptorsandCOP1 137 6.3.3 CDDcomplex 137 6.3.3.1 COP10isanE2Ub-conjugatingenzyme variant 137 6.3.3.2 COP10formsacomplexwithDET1and DDB1 138 6.3.4 COP9signalosome 139 6.3.4.1 CompositionandstructureoftheCOP9 signalosome 139
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