Intercellular Communication in Plants (Annual Plant Reviews, Volume 16) PDF
Preview Intercellular Communication in Plants (Annual Plant Reviews, Volume 16)
P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 Intercellular Communication in Plants i P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 Annual Plant Reviews Aseriesforresearchersandpostgraduatesintheplantsciences.Eachvolumeinthisseries focusesonathemeoftopicalimportance,andemphasisisplacedonrapidpublication. EditorialBoard: Professor Jeremy A. Roberts (Editor-in-Chief), Plant Science Division, School of Bio- sciences,UniversityofNottingham,SuttonBoningtonCampus,Loughborough,Leicester- shireLE125RD,UK;ProfessorHidemasaImaseki,Obata-Minami2419,Moriyama-ku, Nagoya463,Japan;DrMichaelT.McManus,InstituteofMolecularBioSciences,Massey University,PalmerstonNorth,NewZealand;DrJocelynK.C.Rose,DepartmentofPlant Biology,CornellUniversity,Ithaca,NewYork14853,USA. Titlesintheseries: 1.Arabidopsis EditedbyM.AndersonandJ.A.Roberts 2.BiochemistryofPlantSecondaryMetabolism EditedbyM.Wink 3.FunctionsofPlantSecondaryMetabolitesandtheirExploitationinBiotechnology EditedbyM.Wink 4.MolecularPlantPathology EditedbyM.DickinsonandJ.Beynon 5.VacuolarCompartments EditedbyD.G.RobinsonandJ.C.Rogers 6.PlantReproduction EditedbyS.D.O’NeillandJ.A.Roberts 7.Protein–ProteinInteractionsinPlantBiology EditedbyM.T.McManus,W.A.LaingandA.C.Allan 8.ThePlantCellWall EditedbyJ.K.C.Rose 9.TheGolgiApparatusandthePlantSecretoryPathway EditedbyD.G.Robinson 10.ThePlantCytoskeletoninCellDifferentiationandDevelopment EditedbyP.J.Hussey 11.Plant–PathogenInteractions EditedbyN.J.Talbot 12.PolarityinPlants EditedbyK.Lindsey 13.Plastids EditedbyS.G.Møller 14.PlantPigmentsandtheirManipulation EditedbyK.M.Davies 15.MembraneTransportinPlants EditedbyM.R.Blatt 16.IntercellularCommunicationinPlants EditedbyA.J.Fleming ii P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 Intercellular Communication in Plants Editedby ANDREWJ.FLEMING ProfessorofPlantSciences UniversityofSheffield,UK iii P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 (cid:1)c 2005byBlackwellPublishingLtd Editorialoffices: BlackwellPublishingLtd,9600GarsingtonRoad,OxfordOX42DQ,UK Tel:+44(0)1865776868 BlackwellPublishingAsiaPtyLtd,550SwanstonStreet,Carlton,Victoria3053,Australia Tel:+61(0)383591011 ISBN1-4051-2068-1 ISSN1460-1494 PublishedintheUSAandCanada(only)byCRCPressLLC,2000CorporateBlvd.,N.W.,Boca Raton,FL33431,USA OrdersfromtheUSAandCanada(only)toCRCPressLLC USAandCanadaonly: ISBN0-8493-2363-0 ISSN1097-7570 TherightoftheAuthortobeidentifiedastheAuthorofthisWorkhasbeenassertedin accordancewiththeCopyright,DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,or transmitted,inanyformorbyanymeans,electronic,mechanical,photocopying,recordingor otherwise,exceptaspermittedbytheUKCopyright,DesignsandPatentsAct1988,withoutthe priorpermissionofthepublisher. Thisbookcontainsinformationobtainedfromauthenticandhighlyregardedsources.Reprinted materialisquotedwithpermission,andsourcesareindicated.Reasonableeffortshavebeenmade topublishreliabledataandinformation,buttheauthorandthepublishercannotassume responsibilityforthevalidityofallmaterialsorfortheconsequencesoftheiruse. Trademarknotice:Productorcorporatenamesmaybetrademarksorregisteredtrademarks,and areusedonlyforidentificationandexplanation,withoutintenttoinfringe. Firstpublished2005 LibraryofCongressCataloging-in-PublicationData: AcatalogrecordforthistitleisavailablefromtheLibraryofCongress BritishLibraryCataloguing-in-PublicationData: AcataloguerecordforthistitleisavailablefromtheBritishLibrary Setin10/12ptTimes byTechbooks PrintedandboundinGreatBritain byMPGBooksLtd,Bodmin,Cornwall Thepublisher’spolicyistousepermanentpaperfrommillsthatoperateasustainableforestry policy,andwhichhasbeenmanufacturedfrompulpprocessedusingacid-freeandelementary chlorine-freepractices.Furthermore,thepublisherensuresthatthetextpaperandcoverboard usedhavemetacceptableenvironmentalaccreditationstandards. ForfurtherinformationonBlackwellPublishing,visitourwebsite: www.blackwellpublishing.com iv P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 Contents Contributors xi Preface xiii 1 Auxinasanintercellularsignal 1 JIRˇ´IFRIMLANDJUSTYNAWIS´NIEWSKA 1.1 Introduction 1 1.2 Auxintransport–knownpathways 2 1.2.1 Polarauxintransportpathway 2 1.2.2 Chemiosmoticmodel 4 1.2.3 Multicomponentauxineffluxcarriersystem 5 1.3 Molecularcomponents 6 1.3.1 Auxininflux–AUX1proteins 7 1.3.2 Auxinefflux–PINproteins 7 1.3.3 ABCtransporters 9 1.4 Subcellulardynamicsofauxincarriers 9 1.4.1 ConstitutiverecyclingofPINproteins 9 1.4.2 AEIsandvesicletrafficking 11 1.4.3 GNOMandPINdynamics 12 1.5 Theroleofauxingradientsinplantdevelopment 12 1.5.1 Monitoringofauxindistributioninplanta 13 1.5.2 Embryonicaxisformation 13 1.5.3 Postembryonicorganformation 15 1.5.4 Rootmeristemmaintenance 16 1.5.5 Tropisms 17 1.5.6 Downstreamofauxingradients 18 1.5.7 Auxinasmorphogen 21 1.6 Conclusions 21 Acknowledgements 22 References 22 2 Peptidesassignals 27 YIJIXIA 2.1 Introduction 27 2.2 Peptidesignalsinplantsandtheirbiologicalfunctions 28 2.2.1 Systeminsmediatesystemicandlocalwoundresponses 28 2.2.2 RALFregulatesplantgrowthanddevelopment 31 2.2.3 ENOD40regulatesnodulationandcellproliferation 31 v P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 vi CONTENTS 2.2.4 PSK(phytosulfokine)isamitogenicfactor 34 2.2.5 CLAVATA3(CLV3)regulatesstemcellhomeostasis 35 2.2.6 S-locuscysteine-richproteinsdeterminespecificityof self-incompatibilityintheBrassicaceae 38 2.3 Proteolyticprocessingofprohormones 40 2.4 Technologiesfordiscoveringnewpeptidesignals 41 2.5 Concludingremarks 43 References 44 3 RNAasasignallingmolecule 49 PATRICEDUNOYERANDOLIVIERVOINNET 3.1 IntercellularmovementofplantmRNAs 49 3.1.1 Cell-to-cellmovementofplantmRNAs 49 3.1.1.1 Plantplasmodesmata 49 3.1.1.2 Cell-to-cellmovementofatranscriptionfactorwithitsmRNA 50 3.1.2 Long-distancetransportofplantmRNAs 51 3.2 Intercellularmovementofviroids 53 3.2.1 Whatareviroids? 53 3.2.2 Intercellularmovementofviroids 54 3.2.2.1 Cell-to-cellmovementofviroids 54 3.2.2.2 Long-distancemovementofviroids 54 3.2.3 Cellularfactorsinvolvedinviroidmovement 55 3.2.3.1 PhloemLectin2 55 3.2.3.2 VirP1 55 3.3 IntercellularmovementofRNAsilencing 56 3.3.1 MechanismofRNAsilencing 56 3.3.1.1 Co-suppressioninpetunia 57 3.3.1.2 Double-strandedRNA:triggermoleculeofRNAsilencing 57 3.3.1.3 Shortinterfering(si)RNAsarethespecificitydeterminants ofRNAsilencing 58 3.3.1.4 RNA-inducedsilencingcomplexRISC 59 3.3.1.5 TransitiveRNAsilencing 60 3.3.1.6 BiologicalfunctionsofRNAsilencinginplants 61 3.3.2 ThediscoveryofsystemicRNAsilencing 61 3.3.3 InitiationofsystemicRNAsilencing 63 3.3.3.1 SpontaneousactivationofsystemicRNAsilencing 63 3.3.3.2 Exogenouslyinducedsystemicsilencing 64 3.3.4 PropagationofsystemicRNAsilencing 65 3.3.4.1 Long-distancemovementofRNAsilencing 65 3.3.4.2 Cell-to-cellmovementofRNAsilencing 67 3.3.5 Maintenanceofsystemicsilencing 71 3.3.6 Whatisthenucleicacidcomponentofthesilencingsignal? 72 3.3.6.1 Cell-to-cellmovementandphloemtransportofsilencing involveseparatemechanismsand,mostlikely, separatesignals 73 3.3.6.2 PossiblenatureoftheRNAspeciesinvolvedincell-to-cell movementofsilencing 73 P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 CONTENTS vii 3.3.6.3 NospecificRNAspecieshasbeencorrelatedwith long-distancetransportofsilencinginplants 74 3.3.7 PlantfactorsrequiredformovementofRNAsilencing 75 3.3.8 Biologicalfunctionsofnon-cellautonomousRNAsilencinginplants 75 3.3.8.1 Antiviraldefence 75 3.3.8.2 Aroleinnon-cellautonomousregulationofgeneexpression? 76 References 77 4 Theplantextracellularmatrixandsignalling 85 ANDREWJ.FLEMING 4.1 Introduction 85 4.2 Thecellwallandsignalling 86 4.3 Thecellwallasapotentialsourceofchemicalsignals 87 4.3.1 Polysaccharidesignals 88 4.3.2 Arabinogalactanproteinsassignals 90 4.3.3 Cutinandsignalling 94 4.3.4 Uncharacterisedcellwalldeterminantsinvolvedinsignalling 96 4.4 Thecellwallandbiophysicalsignalling 98 4.4.1 Connectionsbetweenthecellwallandthecytosolasaconduit forintercellularsignalling 101 4.5 Conclusions 102 Acknowledgements 103 References 103 5 Plasmodesmata–gatewaysforintercellularcommunicationinplants 109 TRUDIGILLESPIEANDKARLJ.OPARKA 5.1 Introduction 109 5.1.1 Plasmodesmata–keycomponentsofthesymplast 109 5.1.2 Plasmodesmata:simpledescription,complexfunction 110 5.1.3 Discoveryofplasmodesmata 110 5.2 Structure 110 5.2.1 Thegeneralultrastructureofplasmodesmata 110 5.2.2 Primaryandsecondary;simpleorbranched 113 5.2.3 Plasmodesmalfrequencyanddistribution:gainandloss 116 5.2.4 Plasmodesmalcomponents 117 5.2.5 Passagethroughthecytoplasmicsleeve 120 5.3 Macromoleculartrafficking 120 5.3.1 PassivetransportandthebasalSEL 122 5.3.2 Selectivetransportandgating:modulationoftheSEL 124 5.3.3 PhysiologicalmodulationofSEL 124 5.3.4 FineregulationofplasmodesmalSEL–roleofthecytoskeleton 129 5.3.5 ‘Coarse’regulationbycallose 130 5.3.6 Phosphorylation,proteinunfoldingandchaperones 131 5.3.7 Theemergingpictureofplasmodesmata 134 Acknowledgements 134 References 135 P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 viii CONTENTS 6 Lessonsfromthevegetativeshootapex 147 JOHNF.GOLZ 6.1 Introduction 147 6.2 Structureoftheangiospermshootapicalmeristem 147 6.2.1 Zonesofthemeristem 148 6.2.2 Layersofthemeristem 149 6.2.3 Symplasticfieldswithinthemeristem 149 6.3 Periclinalchimaerasrevealaroleforsignallinginplantdevelopment 149 6.4 Signallinginvolvedinmeristemmaintenance 151 6.4.1 TheCLAVATAmutants 151 6.4.2 TheCLAVATAsignallingpathway 154 6.4.3 Thewuschelmutant 156 6.4.4 TheCLAVATA–WUSCHELregulatoryloop 156 6.5 Maintainingindeterminatecellsinthemeristemrequireshomeoboxgenes 158 6.6 InteractionsbetweenKNOXgenesandhormonesregulatemeristemactivity 161 6.7 Signalsinvolvedinorganformation 162 6.7.1 Modelsofphyllotaxis 162 6.7.2 Theroleofauxininphyllotaxis 163 6.7.3 Organoutgrowthinvolvesphysicalforces 166 6.8 Signallingbetweenorganprimordiaandthemeristem 166 6.9 Conclusion 169 Acknowledgements 170 References 170 7 Intercellularcommunicationduringfloralinitiation anddevelopment 178 GEORGECOUPLAND 7.1 Introduction 178 7.2 Long-distancesignalingduringtheinductiontoflowering 179 7.2.1 Discoveryofaroleforlong-distancesignalingintheinduction offlowering 179 7.2.2 Mutationsthatimpairlong-distancesignalinginpeaandmaize 180 7.2.3 Moleculargeneticanalysisofflowering-timecontrolin Arabidopsisplacesthelong-distancesignalwithinaregulatory hierarchy 181 7.2.3.1 AnetworkofpathwayscontrolsfloweringofArabidopsis 181 7.2.3.2 Spatialregulationofflowering-timecontrol 184 7.2.3.3 Identifyingthefloralstimulus:aperspectivefrom Arabidopsismoleculargenetics 187 7.3 Intercellularcommunicationduringfloraldevelopment 188 7.3.1 Someofthetranscriptionfactorsthatcontrolfloralmeristemor organidentityactnon-cellautonomouslyinthedevelopingflower 189 7.3.2 Movementoftranscriptionfactorsbetweencellsdefinesone mechanismforshort-distancesignalinginthedevelopingflower 190 7.4 Perspectives 193 References 194 P1:FCG/FFX P2:FCG/FFX QC:FCG/FFX T1:FCG BY034-FM BY034-Fleming BY034-Fleming-v3.cls October30,2004 10:39 CONTENTS ix 8 Lessonsfromtherootapex 199 MARTINBONKE,SARITA¨HTIHARJUANDYKA¨ HELARIUTTA 8.1 Introduction 199 8.2 Organizationoftheroot 199 8.2.1 Anatomyoftherootmeristemandprocambiumintheapex ofagrowingroot 199 8.2.2 Cellularorganizationoftherootisestablishedduring embryonicdevelopment 200 8.2.3 Developmentofsecondaryroots 203 8.3 Cellfatestudiesofthegrowingroot 203 8.4 Moleculargeneticsofrootdevelopment 205 8.4.1 Distalpatterning 205 8.4.2 Geneticcontrolofinitiationofsecondaryroots 210 8.4.3 Moleculargeneticsofepidermalpatterning 211 8.4.4 Patterningofgroundtissue 215 8.4.5 Vascularpatterning 218 8.5 Futureprospects 220 Acknowledgements 220 References 220 9 Lessonsfromleafepidermalpatterninginplants 225 BHYLAHALLIPURUSHOTTAMSRINIVASAND MARTINHU¨LSKAMP 9.1 Overview 225 9.2 Introduction 225 9.3 Mechanismsoftrichomepatterning 225 9.3.1 Trichomedifferentiation 226 9.3.2 Whyisamechanismpostulatedtoexplainthetrichomespacing patternandwhatkindofunderlyingprinciplesareoperating? 226 9.3.3 Analysisoftrichomeinitiationmutants 227 9.3.3.1 Positiveregulatorsoftrichomeinitiation 228 9.3.3.2 Negativeregulatorsoftrichomeinitiation 229 9.3.4 Interactionsbetweenthetrichomeinitiationgenes 229 9.3.5 Localcell–cellinteractionsleadingtocellfatedecisions:amodel 231 9.3.6 Long-rangecontroloftrichomeinitiationbyhormones 233 9.4 Stomataldevelopmentandpatterning 233 9.4.1 Celldivisionpatternduringstomatapatterning 235 9.4.2 Cellsignallingandthecontrolofasymmetriccelldivisions duringstomatadevelopment 236 9.5 Perspective 237 References 237 10 Lessonsonsignallinginplantself-incompatibilitysystems 240 ANDREWG.MCCUBBIN 10.1 Introduction 240
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