Seed Development, Dormancy and Germination PDF
Preview Seed Development, Dormancy and Germination
P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 Seed Development, Dormancy and Germination i P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 ii P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 Seed Development, Dormancy and Germination Editedby KENTJ.BRADFORD DepartmentofPlantSciences SeedBiotechnologyCenter UniversityofCalifornia Davis,CA95616-8780 USA and HIROYUKINONOGAKI DepartmentofHorticulture OregonStateUniversity Corvallis,OR97331-7304 USA iii P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 (cid:2)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-10:1-4051-3983-8 ISBN-13:978-14051-3983-0 LibraryofCongressCataloging-in-PublicationData Seeddevelopment,dormancy,andgermination/editedbyKentBradford andHiroyukiNonogaki. p. cm.–(Annualplantreviews) Includesbibliographicalreferencesandindex. ISBN-13:978-1-4051-3983-0(hardback:alk.paper) ISBN-10:1-4051-3983-8(hardback:alk.paper) 1.Seeds–Development. 2.Seeds–Dormancy. 3.Germination. I.Bradford, K.J.(KentJ.) II.Nonogaki,Hiroyuki. QK661.S4152007 581.4(cid:3)67–dc22 2006026447 AcataloguerecordforthistitleisavailablefromtheBritishLibrary Setin10/12ptTimes byTechBooks,NewDelhi,India PrintedandboundinIndia byReplikaPressPvtLtd,Kundli Thepublisher’spolicyistousepermanentpaperfrommillsthatoperateasustainableforestry policy,andwhichhasbeenmanufacturedfrompulpprocessedusingacid-freeandelementary chlorine-freepractices.Furthermore,thepublisherensuresthatthetextpaperandcoverboard usedhavemetacceptableenvironmentalaccreditationstandards. ForfurtherinformationonBlackwellPublishing,visitourwebsite: www.blackwellpublishing.com iv P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 Contents ListofContributors xiii Preface xv 1 Geneticcontrolofseeddevelopmentandseedmass 1 MASA-AKIOHTO,SANDRAL.STONEANDJOHNJ.HARADA 1.1 Introduction 1 1.2 Overviewofseeddevelopmentinangiosperms 1 1.3 Geneticcontrolofembryodevelopment 3 1.3.1 Centralregulatorsofembryogenesis 3 1.3.2 Genesinvolvedinthemorphogenesisphaseofembryo development 4 1.3.3 Regulatorsofthematurationphaseofembryo development 5 1.4 Geneticcontrolofendospermdevelopment 6 1.4.1 Genesrequiredforcerealendospermdevelopment 7 1.4.2 Genesthatrepressautonomousendosperm development 7 1.5 Geneticaspectsoftestadevelopment 8 1.5.1 Geneticregulationofflavonoidbiosynthesisand accumulation 9 1.5.2 Regulatorsofmucilagebiosynthesisandaccumulation 9 1.6 Controlofseedmass 10 1.6.1 Geneticfactorsaffectingseedmass 10 1.6.2 Testadevelopmentandseedmass 11 1.6.3 Endospermdevelopmentandseedmass 11 1.6.4 Sugartransportandmetabolismduringseed development 13 1.6.5 Metaboliccontrolofseeddevelopmentandsize 15 1.7 Perspective 17 References 17 2 Seedcoatdevelopmentanddormancy 25 ISABELLEDEBEAUJON,LO¨ICLEPINIEC,LUCILLE POURCELANDJEAN-MARCROUTABOUL 2.1 Introduction 25 2.2 Developmentandanatomyoftheseedcoat 25 2.2.1 Theseedenvelopes 25 2.2.2 TheArabidopsistesta 27 v P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 vi CONTENTS 2.3 Roleoftheseedcoatinseeddormancyandgermination 29 2.3.1 Constraintsimposedbytheseedcoat 29 2.3.2 FlavonoidsinArabidopsisseeds 31 2.3.2.1 Mainflavonoidend-productspresentin seeds 31 2.3.2.2 Moleculargeneticsofflavonoidmetabolism 32 2.3.2.3 Effectsofflavonoidsonseeddormancyand germination 35 2.3.3 Flavonoidsinseeddormancyandgermination ofvariousspecies 38 2.3.3.1 Solanaceae 38 2.3.3.2 WaterpermeabilityoftestaeinLeguminosae andotherspecies 39 2.3.3.3 Flavonoidsandotherphenolicsasdirectand indirectgerminationinhibitors 39 2.3.3.4 Pre-harvestsprouting(PHS)incereals 40 2.3.3.5 Heteromorphismandphysiological heterogeneityamongseeds 40 2.3.3.6 Interactionswithendosperm 41 2.4 Linkbetweenseedcoat-imposeddormancyandlongevity 41 2.5 Concludingremarks 42 References 43 3 Definitionsandhypothesesofseeddormancy 50 HENKW.M.HILHORST 3.1 Introduction 50 3.2 Classificationsofdormancy 50 3.2.1 Endogenousdormancy 52 3.2.2 Exogenousdormancy 53 3.3 Definitionsofdormancy 54 3.4 Primarydormancy 57 3.4.1 Inductionofprimarydormancy 57 3.4.1.1 RoleofABAindormancyinduction 57 3.4.1.2 Developmentalprogramsanddormancy induction 58 3.4.2 Releaseofprimarydormancy 59 3.4.2.1 After-ripening 59 3.4.2.2 Regulationofdormancyinimbibed seeds 60 3.5 Secondarydormancy 63 3.6 Signalingindormancy 64 3.6.1 Stresssignaling 64 3.6.2 Signalingnetworks 65 3.6.3 Environmentalsignals 65 3.7 Challengesforthefuture 67 References 67 P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 CONTENTS vii 4 Modelingofseeddormancy 72 PHILS.ALLEN,ROBERTOL.BENECH-ARNOLD,DIEGO BATLLAANDKENTJ.BRADFORD 4.1 Introduction 72 4.2 Typesandphenologyofseeddormancy 73 4.3 Environmentalcontrolofdormancy 76 4.3.1 Factorsaffectingdormancylevelsofseedpopulations 76 4.3.1.1 Temperature 76 4.3.1.2 After-ripening 78 4.3.1.3 Stratification 79 4.3.2 Factorsthatstimulategermination 80 4.3.2.1 Fluctuatingtemperature 80 4.3.2.2 Light 81 4.3.2.3 Nitrate 82 4.3.3 Conceptualschemeofdormancyanditsrelationshipto modeling 82 4.4 Approachestomodelingseeddormancy 82 4.4.1 Temperatureresponsemodelsandthermaltime 83 4.4.2 Waterpotentialresponsesandhydrotimemodels 87 4.4.3 Interactionsoftemperatureandwaterpotential 89 4.4.4 Modelingresponsestootherfactorsaffecting dormancyandgermination 90 4.5 Examplesofseeddormancymodels 91 4.5.1 Solanumtuberosum 91 4.5.2 Bromustectorum 92 4.5.3 Polygonumaviculare 96 4.5.3.1 Modelingseedgerminationresponsesto temperature 97 4.5.3.2 Modelingseedresponsesto germination-stimulatingfactors 100 4.6 Population-basedthresholdmodelsofseeddormancy 100 4.7 Conclusionsandfuturedirections 105 References 106 5 Geneticaspectsofseeddormancy 113 LEO´NIEBENTSINK,WIMSOPPEAND MAARTENKOORNNEEF 5.1 Introduction 113 5.2 MutantapproachesinArabidopsis 114 5.3 Mutantapproachesinotherspecies 118 5.4 Geneticanalysesofnaturalvariation 119 5.4.1 GeneticanalysisofnaturalvariationinArabidopsis 120 5.4.2 Naturalvariationfordormancyingrasses 120 5.5 Whatdothegeneticsteachusaboutdormancyand germination? 125 References 127 P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 viii CONTENTS 6 Lipidmetabolisminseeddormancy 133 STEVENPENFIELD,HELENPINFIELD-WELLSAND IANA.GRAHAM 6.1 Introduction 133 6.2 MetabolicpathwaysforTAGbreakdownandconversion tosucrose 135 6.2.1 TAGhydrolysisandactivation 135 6.2.2 Importoffattyacidsintotheperoxisome 135 6.2.3 Activationoffattyacidstoacyl-CoAthioestersfor β-Oxidation 136 6.2.4 β-Oxidation 137 6.2.4.1 Acyl-CoAoxidases 137 6.2.4.2 Multifunctionalprotein 137 6.2.4.3 3-l-Ketoacyl-CoAthiolase 138 6.2.4.4 Peroxisomalcitratesynthase 138 6.2.5 Glyoxylatecycleandgluconeogenesis 138 6.2.5.1 Isocitratelyase 139 6.2.5.2 Malatesynthase 139 6.2.5.3 Phosphoenolpyruvatecarboxylase 139 6.3 Lipidmetabolismandseeddormancy 140 6.3.1 ImportanceoftheABCtransporterforthetransition fromdormancytogermination 140 6.3.2 Defectsinβ-oxidationenzymes,butnotinLACS, affectseeddormancy 142 6.3.3 Storagelipidmobilization(glyoxylatecycleand gluconeogenesis)isnotrequiredforseeddormancy release 144 6.4 Mechanismsfortheinvolvementofβ-oxidationindormancy release 145 6.4.1 β-Oxidationdoesnotfuelseedgermination 145 6.4.2 β-Oxidationandhormonalsignaling 145 6.4.3 Possiblebiosyntheticrolesforβ-oxidationin regulatinggermination 147 6.4.4 β-Oxidation,reactiveoxygenspecies,andredoxcontrol 148 6.5 Conclusions 149 References 149 7 Nitricoxideinseeddormancyandgermination 153 PAULC.BETHKE,IGORG.L.LIBOURELAND RUSSELLL.JONES 7.1 Nitricoxideinplantgrowthanddevelopment 153 7.2 ChallengesinNOchemistryandbiology 153 7.3 ToolsusedinNOresearch 154 7.4 RolesofNOandotherN-containingcompoundsinseed dormancyandgermination 157 P1:OTE/SPH P2:OTE BLUK053-Bradford September29,2006 21:43 CONTENTS ix 7.4.1 Nitrate,nitrite,andammonium 158 7.4.2 Cyanideandazide 159 7.4.3 NOdonorsandgermination 160 7.4.4 NOscavengersandgermination 162 7.5 BiochemicalandmolecularbasisofNOactioninseeds 162 7.5.1 SynthesisofNObyplants 162 7.5.2 NObindingtometal-containingproteins 164 7.5.3 NOasanantioxidant 166 7.6 InteractionsbetweenNOandphytochromeorABA 167 7.7 EcologicalsignificanceofNO 168 7.7.1 Nitrogenandvegetationgapsensing 168 7.7.2 SmokeandNO 169 7.8 Unresolvedquestionsandconcludingremarks 169 References 171 8 Amergingofpaths:abscisicacidandhormonalcross-talk inthecontrolofseeddormancymaintenanceand alleviation 176 J.ALLANFEURTADOANDALLISONR.KERMODE 8.1 Introduction 176 8.2 Abscisicacid 177 8.2.1 ABAinseedmaturationandtheinductionofprimary dormancy 177 8.2.2 Transcriptionfactorsandcombinatorialcontrolofseed developmentandmaturation 180 8.2.3 ABAindormancymaintenanceandtermination 182 8.2.3.1 ABAsynthesisandhomeostasisduring dormancymaintenanceandtermination 182 8.2.3.2 ABAsignalingfactorsandthecontrolof dormancymaintenanceandtermination 186 8.3 Gibberellin 190 8.3.1 GAisantagonistictoABAduringseeddevelopment 190 8.3.2 GApromotesthetransitiontogermination 191 8.4 Lightinteractions 195 8.4.1 GAsynthesisandsignalingarepromotedbylight throughtheactionofphytochrome 195 8.4.2 ABA-associatedsignalingprocessesareopposedby lightsignaling 196 8.5 Ethylene 197 8.5.1 EthylenecounteractsABAduringseeddevelopment 197 8.5.2 Ethylenepromotesthetransitionfromdormancyto germination 198 8.6 Auxinandcytokinin 201 8.6.1 Auxinandcytokininestablishtheembryobodyplan duringseeddevelopment 201
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