Seeds, Second Edition: Ecology, Biogeography, and, Evolution of Dormancy and Germination PDF
Preview Seeds, Second Edition: Ecology, Biogeography, and, Evolution of Dormancy and Germination
Seeds Ecology, Biogeography, and Evolution of Dormancy and Germination To Drs. Lela Viola Barton and Marianna Georgievna Nikolaeva, two ladies who distinguished themselves by their contributions to our knowledge of seed germination biology Seeds Ecology, Biogeography, and Evolution of Dormancy and Germination Second Edition Carol C. Baskin Jerry M. Baskin Department of Biology University of Kentucky Lexington, Kentucky, USA AMSTERDAM(cid:129)BOSTON(cid:129)HEIDELBERG(cid:129)LONDON(cid:129)NEWYORK(cid:129)OXFORD(cid:129)PARIS SANDIEGO(cid:129)SANFRANCISCO(cid:129)SINGAPORE(cid:129)SYDNEY(cid:129)TOKYO AcademicPressisanimprintofElsevier AcademicPressisanimprintofElsevier 525BStreet,Suite1900,SanDiego,CA92101-4495,USA 32JamestownRoad,LondonNW17BY,UK 225WymanStreet,Waltham,MA02451,USA Copyrightr2014ElsevierInc.Allrightsreserved. Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmittedinanyform orbyanymeanselectronic,mechanical,photocopying,recordingorotherwisewithoutthepriorwritten permissionofthepublisher. PermissionsmaybesoughtdirectlyfromElsevier’s Science&TechnologyRightsDepartment inOxford,UK:phone(144)(0)1865843830;fax(144)(0)1865853333;email:[email protected]. Alternatively,visittheScienceandTechnologyBookswebsiteatwww.elsevierdirect.com/rightsfor furtherinformation. Notice Noresponsibilityisassumedbythepublisherforanyinjuryand/ordamagetopersonsorproperty asamatterofproductsliability,negligenceorotherwise,orfromanyuseoroperationofanymethods, products,instructionsorideascontainedinthematerialherein.Becauseofrapidadvancesinthe medicalsciences,inparticular,independentverificationofdiagnosesanddrugdosagesshould bemade. BritishLibraryCataloguing-in-PublicationData AcataloguerecordforthisbookisavailablefromtheBritishLibrary LibraryofCongressCataloging-in-PublicationData AcatalogrecordforthisbookisavailablefromtheLibraryofCongress ISBN:978-0-12-416677-6 ForinformationonallAcademicPresspublications visitourwebsiteatelsevierdirect.com TypesetbyMPSLimited,Chennai,India www.adi-mps.com PrintedandboundintheUnitedStatesofAmerica 14 15 16 17 18 10 9 8 7 6 5 4 3 2 1 Contents PrefacetotheSecondEdition xi PercentagesandRatesofGermination 28 PrefacetoFirstEdition xiii VeryFastGerminatingSpecies 30 LengthofGerminationTestPeriod 31 TestViabilityofNongerminatedSeeds 31 GerminatingSeedsYouKnowNothingAbout 33 1. Introduction 3. Types of Seeds and Kinds of Seed Purpose 1 Dormancy SeedGerminationEcology 1 ABookonSeedDormancyandGermination Purpose 37 fromaDifferentPerspective 2 TypesofSeeds 37 Need 2 ClassificationofSeedDormancy 39 Content 3 HierarchicalClassificationSystem 39 Nikolaeva’sFormulaSystem 41 DichotomousKeyforSeedDormancy 41 2. Ecologically Meaningful Germination OverviewofClassesofSeedDormancy 41 Studies PhysiologicalDormancy 41 Purpose 5 MorphologicalDormancy 63 DefinitionofSeedDormancy 5 MorphophysiologicalDormancy 68 GuidelinesforLaboratoryStudieson PhysicalDormancy 72 GerminationEcology 7 PhysicalPlusPhysiologicalDormancy 77 CollectSeedsatMaturity 7 CheckSeedsforPresenceofEmbryo 9 4. Germination Ecology of Seeds TestandUseSeedsImmediately with Nondeep Physiological afterHarvesting 10 Dormancy CheckforImbibitionofWater 12 UseIntactNaturalDispersalUnits 12 Purpose 79 Replications 13 DiscoveryofDormancyCycles 79 StatisticalAnalyses 14 ChangesintheStateofDormancy 80 PetriDishes 14 TheDormancyContinuum 86 Substrate 14 EnvironmentalFactorsCausingChangesin PlacementofSeedson/inSubstrate 16 DormancyStates 87 Water 17 Temperature 88 pH 17 Darkness 99 DisinfectantsandFungicides 19 Light 102 Constantvs.AlternatingTemperatureRegimes 19 Gases 104 LightandDarkness 20 Water 106 StoreSeedsunderNaturalorSimulated HydrothermalTimeModelandGermination 109 HabitatConditions 26 InorganicChemicals 110 TestSeedsFrequently 27 OrganicChemicals 112 v vi Contents 5. Germination Ecology of Seeds with LightandDark 164 Morphophysiological Dormancy MoistureStress 165 Flooding 166 Purpose 119 pH 166 LevelsofMorphophysiologicalDormancy 119 EnvironmentalControlofBreakingPhysical NondeepSimpleMorphophysiological Dormancy 166 Dormancy 119 HighandHighFluctuatingTemperatures 166 IntermediateSimpleMorphophysiological TwoStepsinBreakingPhysicalDormancy Dormancy 124 withHighTemperatures 169 DeepSimpleMorphophysiological DryingatHighTemperatures 171 Dormancy 124 Fire 172 DeepSimpleEpicotylMorphophysiological LowWinterTemperatures 175 Dormancy 127 TwoStepsinBreakingPhysicalDormancy NondeepSimpleEpicotyl withLowTemperatures 176 MorphophysiologicalDormancy 132 IncubationonMoistSubstrate 176 DeepSimpleorSimpleEpicotyl BurialinSoilintheField 177 MorphophysiologicalDormancy? 133 MicrobialAction 177 DeepSimpleDoubleMorphophysiological EffectsofAnimalsonSeedswithPhysical Dormancy 134 Dormancy 178 ASpecialKindofSimpleDouble Long-TermStorageofPermeableSeeds 180 MorphophysiologicalDormancy 137 SeedswithPhysicalandPhysiological NondeepComplexMorphophysiological Dormancy 181 Dormancy 137 IntermediateComplexMorphophysiological 7. Germination Ecology of Seeds in Dormancy 139 the Persistent Seed Bank DeepComplexMorphophysiological Dormancy 140 Purpose 187 AerialSeedBanks 187 6. Germination Ecology of Seeds DevelopmentofSerotiny 187 HowLongisSeedDispersalDelayed? 188 with Physical Dormancy Seed-holdingStructures 191 Purpose 145 SizeofSeedBank 191 GerminationPhenology 145 OpeningofSeed-holdingStructures 192 InternalMoistureConditionsofSeeds 145 LethalTemperatures 193 ArtificialSofteningofWater-Impermeable SeedDormancyandGermination 193 Seed(orFruit)Coat 150 SoilSeedBanks 195 MechanicalScarification 150 Formation 195 AcidScarification 150 SpatialDistribution 199 Enzymes 153 MethodologyofStudies 201 OrganicSolvents 153 TypesofPersistentSeedBanks 213 Percussion 154 HabitatsandSeedDensity 220 HighAtmosphericPressures 154 SeedSizeandShape 225 WetHeat 155 LongevityofSeedsintheSoil 228 DryHeat 157 InferredAge 228 DryStorage 159 ActualAge 230 Radiation 160 SurvivorshipCurves 231 Ultrasound 161 ImpactofAnimalsonWater-PermeableSeeds LowTemperatures 161 EnteringtheSeedBank 233 GerminationRequirementsof WhatCanHappentoSeedsintheSoil? 235 PermeableSeeds 162 ModelsofSeedDynamicsintheSoil 235 Imbibition 162 StudiesonSeedBankDynamics 236 Temperature 164 ChangesinDormancyStates 238 Contents vii InsituGermination 239 GerminationRequirements 299 PredictingSeedlingEmergence 243 GerminationSpeed 302 SeedDeathDuetoAging 245 Storability 303 SeedDeathDuetoActivitiesofPredators SecondaryDormancy 303 andPathogens 249 VariationinSeedGerminationResponses 303 SeedDeath:EffectsofFire 256 TypesofVariation 303 SeedDeathDuetoSolarizationand CommonGardenStudies 310 Composting 257 LocalAdaptation 312 SeedDeathDuetoFlooding 258 FactorsthatCausePreconditioningEffects 314 EffectsofSeedBanksonPopulations 258 CarbonDioxideLevels 314 SurvivalinRiskyEnvironments 259 Competition/MaternalCropEnvironment 314 PopulationStability 260 DayLength 315 PopulationVariability 261 Herbicides 316 EffectofSeedBanksonSelectionofOther Hormones 316 Traits 263 LengthofGrowingSeason 317 Density-escapingCharacteristics 264 LightQuality 317 CoexistenceofSpecies 264 MineralNutrition 318 PersistenceofCommunities 265 PhysiologicalAgeofPlants 319 RestorationofPlantCommunities 265 PositionofFruit/SeedonMonomorphic GeneBanksandRestoration 265 MotherPlants 320 RestorationPotentialofSeedBanks 266 PrematureDefoliation/Herbivory 322 SeedRainandTransientSeedBanks 266 SoilMoisture 323 PresenceofNativeandAlienWeedsin SolarIrradiance 323 SeedBank 271 Temperature 324 PresenceofRareSpeciesinSeedBank 272 Time:Year,SeasonandWithinSeason 326 Seed-RelatedStrategiestoPromote UltravioletLight 327 Restoration 272 VirusesandFungi 327 OtherFactors 327 8. Variation in Seed Dormancy and ChangesinSeedsasaResultofthe Germination within and between PreconditioningEnvironment 327 ChemicalCompounds 328 Individuals and Populations SeedSize,ColorandShape 329 of a Species StructuralandChemicalChanges 338 Purpose 277 HeritableChanges 340 InheritanceofSeedDormancy 277 VariationinSeedswithinaSpecies 340 SpecieswithHeritableSeedDormancy 277 MassSeedCollectionsandUnimodal TypesofInheritedSeedDormancy 277 (Monomorphic)SeedVariation 340 SelectionofPureLines 277 HeritabilityandUnimodalVariation 343 CrosseswithinaSpecies 278 Seed/FruitHeteromorphism 344 GenesandGeneMutations 280 GeneticPolymorphism 352 Maternal,PaternalandEmbryonicGenotypes 281 ConsequencesofSeedVariationon Maternal(Non-Mendelian)Influence 283 GerminationandSeedlingSize Chromosomes 284 andSurvival 353 InbreedingDepression 285 Cleistogamy,Amphicarpy,Geocarpy Heterostyly 293 andSeedGerminationEcology 362 Pollen(Microgametophyte)Competition CleistogamousFlowers 362 (PollenLoad) 293 Amphicarpy 363 NumberofPollenDonors(MultiplePaternity) 294 Geocarpy 365 InheritanceofDormancy-Breakingand CausesofCleistogamy 366 GerminationCharacteristics 295 SeedSize 368 DegreeofDormancy 295 SeedGermination 369 DormancyBreak 298 ConsequencesonSeedlingRecruitment 371 viii Contents 9. A Geographical Perspective on SpecialFactor:BiologicalCrust 588 Germination Ecology: Tropical SpecialFactor:NursePlants 589 and Subtropical Zones Purpose 375 10. A Geographical Perspective on EvergreenRainforests 375 Germination Ecology: Temperate ClimaxTrees 375 and Arctic Zones PioneerTrees 433 DormancyandGerminationofShrubs 439 Purpose 591 DormancyandGerminationofLianas 442 SclerophyllousWoodlandswithWinterRain 591 GerminationofHerbaceousSpecies 444 SeedDormancyandGerminationofTrees 592 SpecialBioticFactor:SeedDispersal 450 SeedDormancyandGerminationofShrubs 598 TropicalMountains 452 SeedDormancyandGerminationofVines 626 MontaneSpecies 452 SeedDormancyandGerminationof SeedDormancyandGerminationof HerbaceousSpecies 626 SubalpineandAlpineSpecies 468 SpecialFactor:Fire 645 SpecialFactor:InvasiveSpecies 468 SpecialFactor:Allelopathy 676 SemievergreenTropicalForests 470 SpecialFactor:Myrmecochory 678 GerminationPhenology 470 MoistWarmTemperatureWoodlands 683 DormancyandGerminationofTrees 471 SeedDormancyandGerminationofTrees 683 DormancyandGerminationofBamboos SeedDormancyandGerminationofShrubs andShrubs 499 andBamboos 699 DormancyandGerminationofLianas 501 SeedDormancyandGerminationofLianas DormancyandGerminationof andVines 706 HerbaceousSpecies 503 SeedDormancyandGerminationof ToleranceofSeedDamage 507 HerbaceousSpecies 708 TropicalDeciduousForests 508 SpecialFeature:Myrmecochory 713 DormancyandGerminationofTrees 508 Deciduous(Nemoral)Forests 713 DormancyandGerminationofShrubs 516 SeedDormancyandGerminationofTrees 714 DormancyandGerminationofVines 517 SeedDormancyandGerminationofShrubs DormancyandGerminationof andBamboos 722 HerbaceousSpecies 517 SeedDormancyandGerminationofVines 728 ChlorophyllousEmbryos 522 SeedDormancyandGerminationof SpecialBioticFactor:SeedPredation 523 HerbaceousWoodlandSpecies 728 TropicalDryWoodlands,NaturalSavannas SeedDormancyandGerminationof andGrasslands 524 HerbaceousNonwoodlandSpecies 735 DormancyandGerminationofTrees 525 SpecialFactors 758 DormancyandGerminationofShrubs 536 SteppesorGrasslandsoftheTemperate DormancyandGerminationofLianas 543 Zones 766 DormancyandGerminationofHerbaceous SeedDormancyandGerminationofTrees 766 Species 543 SeedDormancyandGerminationofShrubs 766 SpecialFactor:Fire 554 SeedDormancyandGerminationofVines 769 HotSemidesertsandDeserts 557 SeedDormancyandGerminationof DormancyandGerminationofTrees 558 HerbaceousSpecies 769 DormancyandGerminationofShrubs 561 SeedDormancyandGerminationofWeeds 788 DormancyandGerminationofVines 568 SpecialFactors 788 DormancyandGerminationofPerennial SemidesertsandDesertswith Succulents 569 ColdWinters 794 DormancyandGerminationofHerbaceous SeedDormancyandGerminationofShrubs 794 Perennials 574 SeedDormancyandGerminationof AnnualSpecies 577 HerbaceousSpecies 798 SpecialFactor:WaterasitRelatestoDispersal 587 SpecialFactors 803 Contents ix BorealConiferousandTemperate Halophytes 957 SubalpineZones 806 SaltMarshesandSaltDeserts 957 SeedDormancyandGerminationofSpecies Beaches,CliffsandForedunes 975 inBorealandNorth-temperate MarineAngiosperms 980 SubalpineZones 806 Mangroves 986 SeedDormancyandGermination PsammophytesofNonsalineSoils 993 ofSpeciesinSouth-temperate SeedDormancy 993 SubalpineZones 834 Dormancy-breakingRequirements 997 SpecialFactors 835 GerminationRequirements 998 ArcticandTemperate-ZoneAlpineTundra 837 SeedDormancyandGermination 12. Biogeographical and Evolutionary ofShrubs 838 Aspects of Seed Dormancy SeedDormancyandGerminationof HerbaceousSpecies 838 Purpose 1005 SpecialFactors 850 WorldBiogeographyoftheKindsof Mountains 852 SeedDormancy 1005 Montane 852 Dormancyvs.Nondormancy 1005 Woodland(Colline) 862 GeographicalDistributionofSeed Dormancy 1006 EnvironmentalGradients,SeedDormancy 11. Germination Ecology of Plants andLifeForms 1007 with Specialized Life Cycles DormancyProfiles:Future Considerations 1009 and/or Habitats SpecializedHabitatsorLifeCycles 1010 Purpose 869 RareSpecies 1011 ParasiticPlants 869 InvasiveSpecies 1015 Holoparasites 869 TheoreticalConsiderationsonthe AHoloparasitewithChlorophyll:Cuscuta 881 EvolutionofSeedDormancy 1017 AHemiparasiticMemberoftheLauraceae: PersistenceinRiskyEnvironments 1017 Cassytha 884 EscapefromCompetition 1024 RootedHemiparasitesWhoseSeedsRequire SurvivalinUnfavorableSeasons 1027 aHostStimulusforGermination: OptimizingtheTimefor StrigaandAlectra 885 Germination 1028 RootedHemiparasitesWhoseSeeds SeedDormancyandtheEvolution donotRequireaHostStimulus ofOtherTraits 1033 toGerminate 896 PaleoclimateasaBackgroundforthe NonrootedHemiparasites:Mistletoes 902 EvolutionofSeedDormancy 1033 Myco-HeterotrophicPlantsOtherThan RainfallPatterns 1034 Orchids 908 Temperature 1035 Orchids 910 FossilHistoryofSeedsasitRelatesto SeedDormancy 911 Dormancy 1038 Dormancy-breakingRequirements 912 FossilSeeds 1038 GerminationRequirements 913 FossilEmbryos 1042 CarnivorousPlants 925 SpeculationsonDormancy 1044 SeedDormancy 925 SeedDormancyandPhylogeneticPosition LongevityofStoredSeeds 927 ofFamily 1045 Dormancy-breakingRequirements 927 Gymnosperms 1045 GerminationRequirements 927 Angiosperms 1046 Aquatic(Nonsaline)Plants 929 SeedDormancyandPresumedageof SeedDormancy 929 PlantFamilies 1046 Dormancy-breakingRequirements 939 Gymnosperms 1046 GerminationRequirements 943 Angiosperms 1048
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