FM BLBK197-Ostergaard September12,2009 10:29 CharCount= ANNUAL PLANT REVIEWS VOLUME 38 Fruit Development and Seed Dispersal Edited by Lars Østergaard JohnInnesCentre,NorwichResearchPark,Norwich,UK A John Wiley & Sons, Ltd., Publication Fruit Development and Seed Dispersal Edited by Lars Østergaard © 2010 Blackwell Publishing Ltd. ISBN: 978-1-405-18946-0 i FM BLBK197-Ostergaard September12,2009 10:29 CharCount= Thiseditionfirstpublished2010 (cid:1)C 2010BlackwellPublishingLtd BlackwellPublishingwasacquiredbyJohnWiley&SonsinFebruary2007.Blackwell’s publishingprogrammehasbeenmergedwithWiley’sglobalScientific,Technical,andMedical businesstoformWiley-Blackwell. 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Annualplantreviews(Print)ISSN1460-1494 Annualplantreviews(Online)ISSN1756-9710 (cid:1) Setin10/12ptPalatinobyAptaraR Inc.,NewDelhi,India PrintedinSingapore 1 2010 ii FM BLBK197-Ostergaard September12,2009 10:29 CharCount= AnnualPlantReviews Aseriesforresearchersandpostgraduatesintheplantsciences.Eachvolume inthisseriesfocusesonathemeoftopicalimportanceandemphasisisplaced onrapidpublication. EditorialBoard: Prof.JeremyA.Roberts(Editor-in-Chief),PlantScienceDivision,Schoolof Biosciences,UniversityofNottingham,SuttonBoningtonCampus, Loughborough,Leicestershire,LE125RD,UK; DrDavidEvans,SchoolofBiologicalandMolecularSciences,Oxford BrookesUniversity,Headington,Oxford,OX30BP; Prof.HidemasaImaseki,Obata-Minami2419,Moriyama-ku,Nagoya463, Japan; DrMichaelT.McManus,InstituteofMolecularBioSciences,Massey University,PalmerstonNorth,NewZealand; DrJocelynK.C.Rose,DepartmentofPlantBiology,CornellUniversity, Ithaca,NewYork14853,USA. Titlesintheseries: 1. Arabidopsis EditedbyM.AndersonandJ.A.Roberts 2. BiochemistryofPlantSecondaryMetabolism EditedbyM.Wink 3. FunctionsofPlantSecondaryMetabolitesandtheirExploitationin Biotechnology 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.Laing,andA.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.Moller 14. PlantPigmentsandtheirManipulation EditedbyK.M.Davies 15. MembraneTransportinPlants EditedbyM.R.Blatt iii FM BLBK197-Ostergaard September12,2009 10:29 CharCount= 16. IntercellularCommunicationinPlants EditedbyA.J.Fleming 17. PlantArchitectureandItsManipulation EditedbyCGNTurnbull 18. Plasmodeomata 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 24. PlantHormoneSignaling EditedbyP.HaddenandS.G.Thomas 25. PlantCellSeparationandAdhesion EditedbyJ.R.RobertsandZ.Gonzalez-Carranza 26. SenescenceProcessesinPlants EditedbyS.Gan 27. SeedDevelopment,DormancyandGermination EditedbyK.J.BradfordandH.Nonogaki 28. PlantProteomics EditedbyC.Finnie 29. RegulationofTranscriptioninPlants EditedbyK.Grasser 30. LightandPlantDevelopment EditedbyG.Whitelam 31. PlantMitochondria EditedbyDavidC.Logan 32. CellCycleControlandPlantDevelopment EditedbyD.Inze´ 33. IntracellularSignalinginPlants EditedbyZYang 34. MolecularAspectsofPlantDiseaseResistance EditedbyJaneParker 35. PlantSystemsBiology EditedbyGloriaM.CoruzziandRodrigoA.Gutie´rrez 36. TheMossPhyscomitrellaPatens EditedbyCeliaKnight 37. RootDevelopment EditedbyTomBeeckman 38. FruitDevelopmentandSeedDispersal EditedbyLarsØstergaard iv FM BLBK197-Ostergaard September12,2009 10:29 CharCount= CONTENTS Contributors ix Preface xii 1 CarpelEvolution 1 Aure´lieC.M.Vialette-GuiraudandCharlieP.Scutt 1.1 Theimportanceofhavingcarpels 1 1.2 Hypothesesofcarpelorigin 3 1.3 Aphylogeneticframeworkforstudiesofcarpelevolution 7 1.4 Amorphologicalportraitoftheancestralcarpel 10 1.5 Thegeneticcontrolofcarpeldevelopmentinthefirst floweringplants 12 1.6 AmajorrolefortheE-functionintheoriginofthecarpel? 16 1.7 Carpelspecificationinmonocots 18 1.8 Geneduplicationandcarpelevolutioninthecoreeudicots 20 1.9 TheA-functionfindsaroleinfruitdevelopment 22 1.10 Themultipleoriginsandmechanismsofsyncarpyin theangiosperms 23 1.11 Afruitbyanyothername:evolutionaryconvergence betweenangiospermsandgymnosperms 26 References 27 2 GynoeciumPatterninginArabidopsis:ABasicPlanBehinda ComplexStructure 35 EvaSundbergandCristinaFerra´ndiz 2.1 Introduction 35 2.2 Thebasicplaninlateralorgans 36 2.3 TheArabidopsisgynoecium 41 2.4 Geneticandhormonalfactorscontrollinggynoecium development 43 2.5 Conclusion 57 Acknowledgements 57 References 57 3 TheInsandOutsofOvuleDevelopment 70 RaffaellaBattaglia,MonicaColomboandMartinM.Kater 3.1 Introduction 70 3.2 Originoftheovule 71 v FM BLBK197-Ostergaard September12,2009 10:29 CharCount= vi (cid:1) Contents 3.3 OvuledevelopmentinArabidopsis 72 3.4 Sporophytictissues 73 3.5 Gametophytictissue 81 3.6 Interactionbetweenthefemalegametophyteandthe maternalsporophyte 91 3.7 Ovuleidentitydetermination 93 References 97 4 FertilisationandFruitInitiation 107 SaraFuentesandAdamVivian-Smith 4.1 Introduction 107 4.2 Pollination 110 4.3 Femalereceptivityandthecessationofgynoecialgrowth 113 4.4 Additionalrestraintsonflowerdevelopmentand fruitinitiation 115 4.5 Fertilisation 117 4.6 Hormonalcuesduringfruitinitiation 123 4.7 RNAsilencingduringfruitinitiation 141 4.8 Signaltransductionfromovuletocarpelandvascular canalisation 145 4.9 Currentmodelsoffruitinitiation 147 4.10 Concludingremarks 150 Acknowledgements 151 References 151 5 ArabidopsisFruitDevelopment 172 AntonioMart´ınez-LabordaandAntonioVera 5.1 Introduction 172 5.2 MorphologyoftheArabidopsissilique 174 5.3 Determiningtheboundarybetweenvalveandreplum: valvemargingenes 177 5.4 Themakingofvalvesandreplumrequiresrepression ofvalvemargingenes 179 5.5 Suppressorsoftherplphenotype:settingupterritories 182 5.6 Amodelforpatterningthemediolateralaxisofthe Arabidopsissilique 185 5.7 Auxin:asignalingmoleculeforthemediolateralaxis? 192 5.8 Abiotechnologicalview 195 Acknowledgements 196 References 196 6 Long-DistanceSeedDispersal 204 FrankM.Schurr,OrrSpiegel,OferSteinitz,AnaTrakhtenbrot, AsafTsoarandRanNathan 6.1 Introduction 205 6.2 SixgeneralizationsonLDDmechanisms 214 FM BLBK197-Ostergaard September12,2009 10:29 CharCount= Contents (cid:1) vii 6.3 Avector-basedperspectiveontheevolutionand predictabilityoflong-distanceseeddispersal 223 6.4 Futuredirections 230 Acknowledgements 231 References 231 7 SeedDispersalandCropDomestication:Shattering, GerminationandSeasonalityinEvolutionUnder Cultivation 238 DorianQ.FullerandRobinAllaby 7.1 Introduction 239 7.2 Lossofnaturalseeddispersalinwheatandbarley: archaeobotanicalevidence 240 7.3 Non-shatteringinothercereals:rice,pearl milletandmaize 246 7.4 Thegeneticsofnon-shatteringcereals 249 7.5 Reductioninseeddispersalaids 252 7.6 Non-cerealalternative:appendagehypermorphy infibrecrops 254 7.7 Lossofnaturalseeddispersalinpulses andothercrops 255 7.8 Germinationtraitsindomestication:theimportanceof lossofdormancy 257 7.9 Thegeneticbasisfordormancyandgermination 260 7.10 Germinationandseedlingcompetition:changes inseedsize 261 7.11 Thegeneticsofseedsize 271 7.12 Seasonalitycontrols:photoperiodicityand vernalization 273 7.13 Discussion:evolutionanddevelopmentof domesticatedseedtraits 278 References 280 8 FactorsInfluencingtheRipeningandQualityofFleshyFruits 296 CorneliusS.Barry 8.1 Introduction 296 8.2 Controloffruitripening 297 8.3 Transcriptionfactorsserveasmasterregulatorsof fruitripening 298 8.4 Hormonalcontroloffruitripening 302 8.5 Theinfluenceoflightonfruitquality 306 8.6 Thediscoveryofaromaandflavourgenesinfruit 307 8.7 Cellwallchangesinfluencefruitquality 309 8.8 Thecuticleinfluencesfruitqualityand postharvestlongevity 310 FM BLBK197-Ostergaard September12,2009 10:29 CharCount= viii (cid:1) Contents 8.9 GenomicsResources 311 8.10 Conclusionsandfutureperspectives 314 Acknowledgements 314 References 314 9 ParthenocarpyinCropPlants 326 TizianaPandolfini,BarbaraMolesiniandAngeloSpena 9.1 Introduction 326 9.2 Parthenocarpy 328 9.3 Auxin-synthesisparthenocarpy 329 9.4 Parthenocarpyviaauxinsignaltransduction 330 9.5 Parthenocarpyviagibberellinsignaltransduction 334 9.6 Aucsia-silencingparthenocarpy 334 9.7 Auxinsensitivityandparthenocarpy 335 9.8 Apetalousparthenocarpyandtheroleofother floralorgans 336 9.9 Stenospermocarpy 336 9.10 Parthenocarpyinperennialcropplants 337 9.11 Parthenocarpyandfruitcropbreeding 337 9.12 Fromgreenplantstofruitcropplants 340 References 341 Index 346 Colorplatesectionappearsatthestartofthebook,beforeChapterOne FM BLBK197-Ostergaard September12,2009 10:29 CharCount= CONTRIBUTORS RobinAllaby WarwickHRI,UniversityofWarwick,Wellesbourne,Warwick,United Kingdom CorneliusS.Barry A32PlantandSoilSciences,MichiganStateUniversity,EastLansing,MI, USA RaffaellaBattaglia DipartimentodiBiologia,Universita` degliStudidiMilano,ViaCeloria26, Milano,Italy MonicaColombo DipartimentodiBiologia,Universita` degliStudidiMilano,ViaCeloria26, Milano,Italy CristinaFerra´ndiz InstitutodeBiolog´ıaMolecularyCelulardePlantas(UPV-CSIC),Campus delaUniversidadPolite´cnicadeValencia,AvdadelosNaranjoss/n, Valencia,Spain SaraFuentes DepartmentofCropGenetics,JohnInnesCentre,Norwich,UnitedKingdom DorianQ.Fuller InstituteofArchaeology,UniversityCollegeLondon,London,United Kingdom MartinM.Kater DipartimentodiScienzeBiomolecolarieBiotecnologie,Universita` di Milano,viaCeloria26,Milano,Italy AntonioMart´ınez-Laborda Divisio´ndeGene´tica,UniversidadMiguelHerna´ndez,CampusdeSan Juan,Ctra.deValencias/n,03550-SanJuandeAlicante,Spain ix