Table Of ContentFlowering and its Manipulation
Editedby
CHARLESAINSWORTH
DepartmentofAgriculturalSciences
ImperialCollege
WyeCampus
Ashford
Kent
UK
Flowering and its Manipulation
Annual Plant Reviews
Aseriesforresearchersandpostgraduatesintheplantsciences.Eachvolumeinthisseries
focusesonathemeoftopicalimportanceandemphasisisplacedonrapidpublication.
EditorialBoard:
Professor Jeremy A. Roberts (Editor-in-Chief), Plant Science Division, School of
Biosciences, University of Nottingham, Sutton Bonington Campus, Loughborough,
Leicestershire,LE125RD,UK;DrDavidEvans,SchoolofBiologicalandMolecularSci-
ences, Oxford Brookes University, Headington, Oxford, OX3 0BP; Professor Hidemasa
Imaseki,Obata-Minami2419,Moriyama-ku,Nagoya463,Japan;DrMichaelT.McManus,
InstituteofMolecularBioSciences,MasseyUniversity,PalmerstonNorth,NewZealand;Dr
Jocelyn K.C. Rose, Department of Plant Biology, Cornell University, Ithaca, New York
14853,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 Editedby
P.J.Hussey
11. Plant–PathogenInteractions EditedbyN.J.Talbot
12. PolarityinPlants EditedbyK.Lindsey
13. Plastids EditedbyS.G.Moller
14. PlantPigmentsandtheirManipulation EditedbyK.M.Davies
15. MembraneTransportinPlants EditedbyM.R.Blatt
16. IntercellularCommunicationinPlants EditedbyA.J.Fleming
17. PlantArchitectureanditsManipulation EditedbyC.Turnbull
18. Plasmodesmata EditedbyK.J.Oparka
19. PlantEpigenetics EditedbyP.Meyer
20. FloweringanditsManipulation EditedbyC.Ainsworth
21. EndogenousPlantRhythms EditedbyA.HallandH.McWatters
22. ControlofPrimaryMetabolisminPlants EditedbyW.C.PlaxtonandM.T.McManus
23. BiologyofthePlantCuticle EditedbyM.Riederer
Flowering and its Manipulation
Editedby
CHARLESAINSWORTH
DepartmentofAgriculturalSciences
ImperialCollege
WyeCampus
Ashford
Kent
UK
©2006byBlackwellPublishingLtd
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Floweringanditsmanipulation/editedbyCharlesAnisworth.
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1.Plants,Floweringof. I.Ainsworth,C.C.(CharlesColin),1954-
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Contents
Contributors xiii
Preface xv
PartI. CoreDevelopmentandGenetics
1 Adevelopmentalgeneticmodelfortheoriginoftheflower 3
DAVIDA.BAUMandLENAC.HILEMAN
1.1 Introduction 3
1.2 Whatisaflower? 4
1.3 Phylogeneticandpaleontologicalcontext 5
1.4 Evolutionarynoveltiesoftheflower 6
1.4.1 Bisexuality 6
1.4.2 Determinate/compressedaxes 7
1.4.3 Perianth 7
1.5 Orderingthekeystepsinfloralevolution 8
1.6 Developmentalgeneticbackground 9
1.6.1 Positionandidentityofthereproductiveorgans 9
1.6.2 Developmentalregulationoftheperianth 13
1.7 Modelsfortheoriginofbisexuality 14
1.8 Apicalmegasporophyllproductiononamicrosporangiateaxis? 16
1.9 Thecompressionofthefloralaxis 18
1.10 Theevolutionoftheperianth 19
1.11 Theoriginofadimorphicperianth 20
1.12 Conclusion 21
References 22
2 Floralinduction 28
REYNALDTREMBLAYandJOSEPHCOLASANTI
2.1 Introduction 28
2.2 Floraltransitionismarkedbydevelopmentalphasechanges 28
2.3 Floralinductionismediatedthroughmultiplepathways 30
2.4 Photoperiodicfloralinductionprovidesacuetoseasonalchanges 31
2.4.1 Photoreceptorstransducelightsignals 31
2.4.2 Thecircadianclockisself-reinforcing 32
2.4.3 Keygenesintegratephotoperiodicinduction 32
vi CONTENTS
2.4.4 COandFTgenefunctionisconservedinotherplant
species 34
2.4.5 PhotoperiodinductionthroughCO-independent
pathways 35
2.5 Autonomouspathway 36
2.5.1 FLOWERINGLOCUSCintegratesdifferentfloral
inductivepathways 36
2.6 Vernalization 38
2.6.1 MediationofvernalizationinArabidopsisbyFLC
repression 38
2.6.2 Vernalizationincereals 40
2.7 Hormonesandotherfactors 41
2.7.1 Nutrientdiversiontheoryoffloralinduction 41
2.7.2 Gibberellin 42
2.7.3 Long-distancefloralinductivesignals 44
2.7.4 Integrationandcommencementofthefloraltransition 45
2.7.4.1 LEAFYandAPETALA1 45
2.7.4.2 TERMINALFLOWER1 45
2.7.4.3 ConservationofLFYfunctioninhigherplants 45
2.8 Perspective 46
References 46
3 Floralpatterningandcontroloffloralorganformation 49
ELENAM.KRAMER
3.1 Introduction 49
3.2 TheABCmodeloffloralorganidentity 49
3.2.1 ThemajorgeneticplayersintheABCmodel 50
3.2.2 MembersoftheMADS-boxtranscriptionfactorfamily 51
3.2.2.1 Redundantandcomplexfunctionsamong
thefloralMIKCMADS-boxgenes 53
3.2.3 MembersoftheAP2/EREBPtranscriptionfactorfamily 55
3.3 Regulatingtheexpressionofthefloralorganidentitygenes 56
3.4 ConservationandmodificationoftheABCprogram 58
3.4.1 FloralorganidentitygenefunctioninPetunia 58
3.4.2 FloralorganidentitygenefunctioninOryza 60
3.5 Sexdeterminationasamodificationoffloralorganidentity 62
3.6 Futureperspectives 63
References 64
4 Thegeneticcontrolofflowersizeandshape 71
LYNETTEFULTON,MARTINEBATOUX,RAMKISHORYADAV
andKAYSCHNEITZ
4.1 Introduction 71
4.2 Flowerprimordiumoutgrowth 73
CONTENTS vii
4.3 Regulatingflowermeristemsize 74
4.3.1 Cell–cellcommunication,patternformationinthe
meristemandmeristemsize 75
4.3.2 Cellularfactorsregulatingfloralmeristemdevelopment 77
4.4 Earlycontroloforganogenesisintheflower 79
4.5 Generatingorganboundaries 82
4.6 Floralorgansize 84
4.7 Flowershapeandsymmetry 86
4.7.1 Floraldeterminacy 86
4.8 Dorsoventralsymmetry 88
4.8.1 Petalasymmetryandcontortaestivation 90
4.9 Outlook:toboldlygowherenoonehasgonebefore... 91
References 92
5 Inflorescencearchitecture–Movingbeyonddescription
todevelopment,genesandevolution 98
SUSANR.SINGER
5.1 Overview 98
5.2 Inflorescencetypologies 98
5.3 Inflorescencedevelopment 104
5.3.1 Inflorescencemeristem 104
5.3.2 Regulatingdeterminacy 105
5.3.3 Regulatingmeristemidentity 108
5.4 Evolutionofinflorescencearchitecture 108
5.4.1 Teosinte 108
5.4.2 CauliflowerandBroccoli 109
5.4.3 Phenotypicplasticity 110
5.5 Futuredirections:modelinginflorescencearchitecture 111
References 111
PartII. SpecialisedComponentsofDevelopment
6 Close,yetseparate:patternsofmaleandfemalefloral
developmentinmonoeciousspecies 117
RAFAELPERL-TREVESandPREMANANDRAJAGOPALAN
6.1 Introduction 117
6.2 Ecologicalandevolutionaryaspectsof
monoeciousplantdevelopment 118
6.2.1 Evolutionofdiversereproductivestrategiesinland
plants 118
6.2.2 Selectiveforcesthatfavortheevolutionofunisexual
flowers 119
6.2.3 Flexibilityinsexratiosinmonoeciousplants 119
6.2.4 Relationshipbetweenmonoecyandpollination 121
viii CONTENTS
6.2.5 Theandromonoeciousoption:morecompromises? 121
6.3 Howdounisexualflowersdevelop? 122
6.3.1 Structuraldifferencesbetweenmaleandfemaleflowers 122
6.3.2 Unisexualflowerdevelopmentinmaize 123
6.3.2.1 Developmentofthemaleandfemaleflorets 123
6.3.2.2 Mutationsthataffectstamendevelopment:
roleofgibberellin 124
6.3.2.3 Mutationsthataffectpistilabortion 126
6.3.2.4 Cytokinincounteractspistilabortion 127
6.3.2.5 Conservationofthemaizesex
determinationpathwayinotherspecies 128
6.3.3 Unisexualflowerdevelopmentincucumber 129
6.3.3.1 Cucumberasanexperimentalmodelforsex
expression 129
6.3.3.2 Sexexpressioninthecucumberbudand
alongtheshoot 129
6.3.3.3 Inheritanceofsexincucumber 130
6.3.3.4 Environmentalandhormonalregulationof
sexexpression 131
6.3.3.5 Amodelforcucumbersexexpression 132
6.3.3.6 Molecularstudiesonethylene-synthesisand
perceptiongenesinCucumis 134
6.3.3.7 Searchfor‘effectorgenes’thatcarryoutthe
developmentalprogramfollowingsex
determination 138
6.4 Concludingremarks 141
References 142
7 Cytoplasmicmalesterility 147
FRANÇOISEBUDAR,PASCALTOUZETandGEORGES
PELLETIER
7.1 Introduction 147
7.2 Pioneeringworkinplantsexualityandhybridisation 147
7.3 EarlystudiesofCMS 148
7.4 DefinitionofCMSasasystemwithtwogeneticdeterminants 149
7.5 UseofCMSinhybridseedproduction 150
7.6 ThesearchforCMS:intraspecificcrosses,alloplasmic
situationsandcybrids 153
7.7 SpontaneousCMSsresultfromgenomicconflictbetween
nuclearandcytoplasmicgenes 156
7.8 MolecularstudiesonCMSs:asteptowards
understandingorconfusion? 159
7.8.1 TheTexasstory 159
7.8.2 IdentificationofCMSgenes 161
Description:The flowering plants now dominate the terrestrial ecosystems of the planet, and there are good reasons for supposing that the flower itself has been a major contributing factor to the spread of the Angiosperms. The flowers of higher plants not only contain the organs of plant reproduction but are of
