Seed development, dormancy and germination

389 Pages·2007·10.97 MB·English

by  Kent Bradford

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Seed Development, Dormancy
and Germination
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Seed Development, Dormancy
and Germination
Editedby
KENTJ.BRADFORD
DepartmentofPlantSciences
SeedBiotechnologyCenter
UniversityofCalifornia
Davis,CA95616-8780
USA
and
HIROYUKINONOGAKI
DepartmentofHorticulture
OregonStateUniversity
Corvallis,OR97331-7304
USA
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(cid:2)C 2007byBlackwellPublishingLtd
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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
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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
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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
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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
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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
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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