Table Of ContentSenescence Processes in Plants
Editedby
SUSHENGGAN
DepartmentofHorticulture
CornellUniversity
Ithaca
NY,USA
Senescence Processes in Plants
Senescence Processes in Plants
Editedby
SUSHENGGAN
DepartmentofHorticulture
CornellUniversity
Ithaca
NY,USA
(cid:2)C 2007byBlackwellPublishingLtd
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Firstpublished2007byBlackwellPublishingLtd
ISBN:978-1-4051-3984-7
LibraryofCongressCataloguing-in-PublicationData
Senescenceprocessesinplants/editedbySushengGan.
p. cm.—(Annualplantreviews;v.26)
Includesbibliographicalreferencesandindex.
ISBN:978-1-4051-3984-7(hardback:alk.paper)
1.Plants—Aging.I.Gan,Susheng.
QK762.5.S472007
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Contents
Contributors xv
Preface xvii
1 Mitoticsenescenceinplants 1
SUSHENGGAN
1.1 Introduction 1
1.2 Terminologyandtypesofsenescence 1
1.3 Plantsexhibitmitoticsenescence,postmitoticsenescenceand
cellquiescence 3
1.4 Mitoticsenescence:arrestofSAM 4
1.4.1 InitiationofSAM 4
1.4.2 MaintenanceofSAM 4
1.4.3 ArrestofSAM:amitoticsenescenceinnature 5
1.4.3.1 Physiologicalregulation 5
1.4.3.2 Geneticregulation 7
1.5 Roleoftelomereandtelomeraseinmitoticsenescence 7
1.5.1 Telomere 7
1.5.2 Telomerase 7
1.5.3 Telomereshorteningandreplicativesenescencein
animals 8
1.5.4 Telomerebiologyinplants 8
1.6 Closingremarks 9
Acknowledgment 10
References 10
2 Chlorophyllcatabolismandleafcoloration 12
STEFANHO¨RTENSTEINERANDDAVIDW.LEE
2.1 Introduction 12
2.2 Chlorophyllcatabolites 12
2.2.1 Greencatabolites 12
2.2.1.1 Chlorins 12
2.2.1.2 Phytol 15
2.2.2 Cataboliteswithatetrapyrrolicstructure 15
2.2.2.1 Redchlorophyllcatabolites 15
2.2.2.2 Fluorescentchlorophyllcatabolites 16
vi CONTENTS
2.2.2.3 Nonfluorescentchlorophyllcatabolites 16
2.2.2.4 AreNCCsdegradedfurther? 17
2.3 Thechlorophylldegradationpathway 18
2.3.1 Chlorophyllcycle 18
2.3.2 Reactionsongreenpigments 18
2.3.2.1 Chlorophyllase 18
2.3.2.2 Mgdechelation 19
2.3.3 Lossofgreencolor 20
2.3.3.1 Pheophorbideaoxygenase 20
2.3.3.2 Redchlorophyllcatabolitereductase 21
2.3.4 ReactionsonpFCC 21
2.3.4.1 Hydroxylation 21
2.3.4.2 Glucosylation 22
2.3.4.3 Malonylation 22
2.3.4.4 Demethylation 22
2.3.4.5 Tautomerization 22
2.4 Chlorophyllcatabolicmutants 23
2.5 Significanceofchlorophyllbreakdown 23
2.5.1 Topologyofchlorophyllbreakdown 23
2.5.2 Chlbreakdownandcelldeath 24
2.5.3 Chlbreakdownandnitrogeneconomy 25
2.6 Thepigmentsofsenescingleaves 26
2.7 Thefunctionofanthocyaninsinleafsenescence 28
2.7.1 Physiologicalexplanations 28
2.7.2 Ecologicalexplanations 29
2.7.3 Reconcilingtheseexplanations 30
2.8 Conclusionsandperspectives 30
References 31
3 Membranedynamicsandregulationofsubcellularchanges
duringsenescence 39
MARIANNEHOPKINS,LINDAMcNAMARA,CATHERINE
TAYLOR,TZANN-WEIWANGANDJOHNTHOMPSON
3.1 Introduction 39
3.2 Lossofmembranestructuralintegrityduringsenescence 40
3.2.1 Senescence-associatedchangesinthemolecular
organizationofmembranelipidbilayers 40
3.2.2 Roleoflipases 42
3.2.2.1 Initialfateofde-esterifiedfattyacidsin
senescingmembranes 43
3.2.2.2 Autocatalyticnatureofmembranefattyacid
de-esterification 44
3.2.3 Roleofgalactolipases 45
3.3 Roleofproteolysisinmembranesenescence 48
CONTENTS vii
3.4 Dismantlingofmembranesinsenescingtissue 51
3.4.1 Plastoglobuli 51
3.4.2 Cytosoliclipid-proteinparticles 54
3.4.2.1 Sitesofcytosoliclipid-proteinparticle
ontogeny 54
3.5 Roleofautophagy 55
3.6 Metabolismofmembranefattyacidsinsenescingtissues 57
3.6.1 Galactolipidfattyacids 58
3.6.2 Fateofthylakoidfattyacidsduringstress-induced
senescence 59
3.7 Translationalregulationofsenescence 61
References 62
4 Oxidativestressandleafsenescence 69
ULRIKEZENTGRAF
4.1 Introduction 69
4.2 Antioxidativecapacity,oxidativestressandlifespan 71
4.3 Antioxidants 72
4.4 ROSsignaling 74
4.5 Roleofdifferentcellcompartments 77
4.5.1 Peroxisomes 77
4.5.2 Chloroplasts 78
4.5.3 Mitochondria 79
4.5.4 Nucleus 80
4.6 Concludingremarks 81
References 81
5 Nutrientremobilizationduringleafsenescence 87
ANDREASM.FISCHER
5.1 Overview 87
5.2 Macro-andmicronutrientremobilization 88
5.2.1 Carbon 89
5.2.2 Sulfur 90
5.2.3 Phosphorus 90
5.2.4 Potassium 91
5.2.5 Magnesium,calciumandmicronutrients 91
5.3 Nitrogenremobilization 92
5.3.1 Proteindegradationinsenescingleaves 93
5.3.1.1 Classificationofpeptidases 93
5.3.1.2 Compartmentationofpeptidases 94
5.3.1.3 Regulationofpeptidasesduringleaf
senescence 96
5.3.2 Aminoacidmetabolisminsenescingleaves 98
5.3.3 Nitrogentransporttodevelopingsinks 99
viii CONTENTS
5.4 Outlook 101
Acknowledgments 102
References 102
6 Environmentalregulationofleafsenescence 108
AMNONLERS
6.1 Introduction 108
6.2 Lightirradiance 111
6.2.1 Lightintensity 111
6.2.1.1 Lowlight 111
6.2.1.2 Darkness 112
6.2.1.3 Highlight 113
6.2.2 Photoperiod 114
6.2.3 Wavelength 114
6.2.3.1 Red/Farred 114
6.2.3.2 Bluelight 116
6.2.3.3 Ultraviolet 116
6.3 Ozone 118
6.4 Temperature 119
6.5 Droughtstress 120
6.6 Flooding 121
6.7 Salinity 122
6.8 Environmentalpollution–toxicmaterials 123
6.9 Oxidativestressinvolvementinenvironmentalregulationof
senescence 124
6.10 Nutrient/mineralshortage 125
6.11 AtmosphericCO 126
2
6.12 Bioticstress 127
6.13 Concludingremarks 130
References 133
7 Developmentalandhormonalcontrolofleafsenescence 145
JOSH.M.SCHIPPERS,HAI-CHUNJING,JACQUESHILLE
ANDPAULP.DIJKWEL
7.1 Introduction 145
7.2 Developmentalsenescence:aplantgenomeisoptimisedfor
earlysurvivalandreproduction 145
7.3 Developmentalprocessesthatregulateleafsenescence 147
7.3.1 Reactiveoxygenspecies 147
7.3.2 Metabolicflux 148
7.3.3 Proteindegradation 148
7.4 Hormonalcontrolofleafsenescence 149
7.4.1 Hormonesthatdelayleafsenescence 150
7.4.1.1 Gibberellicacid 150
7.4.1.2 Auxin 150
7.4.1.3 Cytokinins 151