Table Of ContentPlant
Cell Death
Processes
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Plant
Cell
Death
Processes
Edited by
Larry D. Noodén
DepartmentofBiology
UniversityofMichigan
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03 04 05 06 07 08 9 8 7 6 5 4 3 2 1
CONTENTS
Contributors xvii
Foreword—AgingandDeath xxi
Preface xxv
1. Introduction 1
I. WhatthisBookCovers 1
II. TheProcesses—Senescence,Aging,ProgrammedCellDeath,
Apoptosis,etc.—EvolvingConcepts 2
III. ApoptosisinAnimals 6
IV. ApoptosisinPlants 8
V. TheSenescenceSyndrome 10
VI. HormonalControls 13
VII. Evolution 14
References 14
2. Plant Cell Death and Cell Differentiation 19
I. Introduction 19
II. TheScopeofPCDinPlants 20
III. PrereproductiveCellDeath 22
A. AleuroneandScutellum 22
B. Xylem 24
C. Aerenchyma 25
D. AbscissionZones 26
IV. ReproductiveCellDeath 27
A. FloralOrgans 27
B. Gametophytes 29
C. Pollination 29
D. Embryogenesis 30
V. Conclusions 31
References 31
v
vi Contents
3. Cell Death in Plant Disease: Mechanisms and
Molecular Markers 37
I. Introduction 37
II. RoleofCellDeathduringPlant–PathogenInteractions 38
III. StructuralandBiochemicalChangesAccompanyingCellDeath
duringPlantDisease 38
IV. DefinitionofStepsInvolvedintheSignalingProcessof
CellDeathInductionduringPlant–PathogenInteractions 39
V. MolecularComponentsforCellDeathControlduring
Plant–PathogenInteractions 41
VI. GlogalAnalysesofMarkersforCellDeathInductionby
PlantPathogens 43
A. IsolationandCharacterizationofHRMarkers 43
B. ComparativeAnalysesofCellDeathPathwaysInducedby
AvirulentPathogensandSenescence 44
C. FutureApproachestoGlobalAnalysisofCell
Death-relatedGenes:Genomic-scaleGeneExpression
MonitoringtoMappingofIntersectingSignalingPathways 45
References 47
4. Changes in Gene Expression during Senescence 51
I. Introduction 51
II. ChangesinPatternsofNucleicAcidsandProteinsduring
Senescence 52
III. SimilaritiesbetweenSenescingandRipeningTissues 52
IV. IdentificationandClassificationofSenescence-relatedGenes 52
V. Senescence-relatedGenes 53
A. RegulationofSRGeneExpressionbyStresses 60
B. RegulationofSRGeneExpressionbyCytokinins 60
C. RegulationofSRGeneExpressionbyEthylene 61
VI. FunctionofSRGenesinSenescence 62
A. GenesInvolvedinProteinDegradation 62
B. GenesInvolvedintheBreakdownofNucleicAcids 63
C. GenesInvolvedinMembraneDisassembly 64
D. GenesInvolvedinRemobilizationofNitrogen 64
E. GenesEncodingProtectiveorDefense-Response
Proteins 65
F. GenesInvolvedinEthyleneBiosynthesisandPerception 66
VII. Summary 66
References 67
5. Genes that Alter Senescence 73
I. Introduction 73
II. SenescenceasaGeneticallyProgrammedProcess 74
Contents vii
III. GenesInvolvedinExecutionofSenescence 76
A. Overview 76
B. ChlorophyllBreakdown 77
C. MembraneandLipidBreakdown 78
D. OtherGenesInvolvedinMacromoleculeDegradation 78
IV. GenesAffectingSenescencethroughActionontheHormonal
Controls 79
A. Overview 79
B. DelayofSenescencebyCytokinin 79
C. RegulationofSenescencebyEthylene 80
D. ChangeofSenescencebyBrassinosteroidBiosynthesis 80
V. GenesthatAlterSenescenceinResponsetoEnvironmental
Factors 80
A. GenesinLightSignaling 80
B. Stress-relatedGenes 81
VI. GenesControllingVegetativeGrowth(Regeneration)and
MonocarpicSenescence 81
VII. RegulatoryGenesandIntracellularSignaling 82
VIII. Conclusions 85
References 87
6. Senescence and Genetic Engineering 91
I. Introduction 91
II. TheRelationshipofCytokininsandSenescence 92
A. ManipulationofCytokininLevels 92
B. CytokininSignalingandMechanismofAction 95
III. TheRelationshipofEthyleneandSenescence 97
IV. ConcludingRemarks 100
References 101
7. Proteolysis 107
I. Introduction 107
II. SelectiveHydrolysisofPeptideBonds 108
A. ProteinMaturationandRemovalofPeptidesfromLarger
Precursors 108
B. DegradationofDamagedProteinsandofUnassembled
Subunits 108
C. ProteinTurnoverandAdaptationoftheMetabolismto
ChangingConditions 108
D. DegradationofMatureProteinsinRelationtoNitrogen
Remobilization 108
III. ProteolyticActivitiesinPlants 109
A. ClassificationofPeptideHydrolases 109
B. ComplexProteolyticSystems 110
C. ATP-DependencyofProteolysis 113
viii Contents
D. Compartmentation 113
IV. ProteolysisinRelationtoCellDeath 114
A. CellDeathduringDifferentiation 114
B. Senescence 115
C. HypersensitiveResponse 116
V. RegulationofProteinCatabolism 116
A. ChangesinthePatternofProteolyticEnzymes 116
B. AlteredSusceptibilityofSubstrateProteins 116
C. AspectsofCompartmentation 118
D. InfluenceofSolutes 118
VI. Conclusions 119
References 120
8. Ethylene Signaling in Plant Cell Death 125
I. Introduction 125
II. EthyleneBiosynthesisPathways 126
A. ACCPathway 126
B. Non-ACCPathway 126
III. TemporalandSpatialRegulationofEthyleneBiosynthesis 127
IV. EthyleneSignalTransductionPathway 127
V. EthyleneCrossTalkwithOtherPlantHormones 129
A. Ethylene,AbscisicAcidandSugar 131
B. EthyleneandAuxin 132
C. EthyleneandCytokinins 132
D. EthyleneandPolyamines 133
VI. ProteaseInvolvementandEthyleneBiosynthesisinPCD 134
VII. HormonalRegulationofPlantPCD 134
VIII. Perspective 136
References 137
9. Jasmonates – Biosynthesis and Role in Stress Responses and
Developmental Processes 143
I. Introduction 143
II. JasmonatesandRelatedCompounds 143
III. LOX-derivedCompoundsandtheBiosynthesisof
OctadecanoidsandJasmonates 144
A. TheLOXPathway 144
B. OctadecanoidandJasmonateBiosynthesis 145
C. SignalingPropertiesofJasmonatesand
Octadecanoids—theOxylipinSignature 146
IV. Jasmonate-inducedAlterationofGeneExpression 147
V. JasmonatesandPlantDefenseReactions 147
VI. JasmonatesinPlantDevelopmentandSenescence 150
References 152
Contents ix
10. Leaf Senescence and Nitrogen Metabolism 157
I. Introduction 157
II. LeafSenescenceandNitrogenLoss 157
A. FateofChloroplastsandtheirConstituents 158
III. ProteinMetabolismduringLeafSenescence 158
A. FatesofDNAandRNA 158
B. SynthesisandDegradationofRubisco 159
C. MolecularBasisforChangesinRubiscoSynthesisduring
Senescence 160
D. Light-HarvestingChlorophylla/bBindingProteinofPSII 161
E. D1protein 162
IV. Proteindegradation 163
V. RemobilizationofNitrogeninSenescingLeaves 163
A. AmideFormationinSenescingLeaves 163
VI. Conclusion 164
References 165
11. Photosynthesis and Chloroplast Breakdown 169
I. Introduction 169
II. DeclineofPhotosyntheticFunction 170
A. DecreaseintheNumberofChloroplastsorGradual
DegradationofPersistingChloroplasts? 170
B. PhotosyntheticPerformance 170
C. RiskofPhotoinhibition 171
III. ChloroplastBreakdown 172
A. SequenceofSenescence-associatedEvents 172
B. DegradationofPlastidConstituents 172
IV. ExpressionofNuclearGenesEncodingPhotosynthesis-related
Proteins 175
A. CalvinCycle-relatedGenes 175
B. GenesEncodingThylakoidMembraneProteins 175
C. RegulatoryMechanismsUnderlyingtheDown-regulation
ofPhotosynthesis-relatedNuclearGenes 176
V. PlastidGeneExpressionduringSenescence 176
A. TranscriptionalActivity 176
B. TranscriptLevels 178
C. ProteinSynthesis 178
D. ExpressionofPlastidGenesnotRelatedto
Photosynthesis 179
VI. RegulationofChloroplastSenescence 180
A. TheDilemmaoftheChloroplast:OngoingPhotosynthesis
duringSenescence-associatedBreakdown 180
B. CoordinationofSenescence-relatedDegradationof
Proteins 180
C. PhotosynthesisandSenescenceinStay-GreenMutantsand
TransgenicPlants 181