Seeds, Second Edition: Ecology, Biogeography, and, Evolution of Dormancy and Germination

1601 Pages·2014·61.674 MB·English

by  Carol C. Baskin, Jerry M. Baskin

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
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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