Annual Plant Reviews Volume 44: The Plant Hormone Ethylene

420 Pages·2012·15.49 MB·English

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ANNUAL PLANT REVIEWS
VOLUME 44
i
BLBK404-FM BLBK404-McManus January12,2012 19:2 Trim:234mm×156mm Series:APR CharCount=
ANNUAL PLANT REVIEWS
VOLUME 44
The Plant Hormone Ethylene
Edited by
Michael T. McManus
InstituteofMolecularBiosciences
MasseyUniversity
NewZealand
A John Wiley & Sons, Ltd., Publication
iii
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p.cm.–(Annualplantreviews;v.44)
Includesbibliographicalreferencesandindex.
ISBN978-1-4443-3003-8(hardcover:alk.paper)
1.Ethylene. 2.Planthormones. I.McManus,MichaelT. II.Series:Annualplant
reviews; v.44.
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1 2012
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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.MichaelT.McManus,InstituteofMolecularBiosciences,Massey
University,PalmerstonNorth,NewZealand
DrJocelynK.C.Rose,DepartmentofPlantBiology,CornellUniversity,
Ithaca,NY14853,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.LaingandA.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
15. MembraneTransportinPlants
EditedbyM.R.Blatt
16. IntercellularCommunicationinPlants
EditedbyA.J.Fleming
17. PlantArchitectureandItsManipulation
EditedbyC.G.N.Turnbull
18. Plasmodesmata
EditedbyK.J.Oparka
BLBK404-FM BLBK404-McManus January12,2012 19:2 Trim:234mm×156mm Series:APR CharCount=
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
EditedbyD.C.Logan
32. CellCycleControlandPlantDevelopment
EditedbyD.Inze´
33. IntracellularSignalinginPlants
EditedbyZ.Yang
34. MolecularAspectsofPlantDiseaseResistance
EditedbyJ.Parker
35. PlantSystemsBiology
EditedbyG.M.CoruzziandR.A.Gutie´rrez
36. TheMossPhyscomitrellapatens
EditedbyC.D.Knight,P-F.PerroudandD.J.Cove
37. RootDevelopment
EditedbyT.Beeckman
38. FruitDevelopmentandSeedDispersal
EditedbyL.Østergaard
39. FunctionandBiotechnologyofPlantSecondaryMetabolites
EditedbyM.Wink
40. BiochemistryofPlantSecondaryMetabolism
EditedbyM.Wink
41. PlantPolysaccharides
EditedbyP.Ulvskov
42. NitrogenMetabolisminPlantsinthePost-GenomicEra
EditedbyC.FoyerandH.Zhang
43. BiologyofPlantMetabolomics
EditedbyR.D.Hall
44. ThePlantHormoneEthylene
EditedbyM.T.McManus
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CONTENTS
ListofContributors xv
Preface xxiii
1 100YearsofEthylene–APersonalView 1
DonGrierson
1.1 Introduction 1
1.2 Ethylenebiosynthesis 2
1.3 Ethyleneperceptionandsignalling 7
1.4 Differentialresponsestoethylene 9
1.5 Ethyleneanddevelopment 10
1.6 Lookingahead 13
Acknowledgements 14
References 14
2 EarlyEventsintheEthyleneBiosyntheticPathway–Regulation
ofthePoolsofMethionineandS-Adenosylmethionine 19
KatharinaBu¨rstenbinderandMargretSauter
2.1 Introduction 20
2.2 ThemetabolismofMetandSAM 22
2.3 RegulationofdenovoMetsynthesis 25
2.4 RegulationoftheSAMpool 27
2.4.1 RegulationofSAMSgenesbyethyleneandof
SAMSenzymeactivitybyprotein-S-nitrosylation 29
2.5 Theactivatedmethylcycle 30
2.6 TheS-methylmethioninecycle 32
2.7 ThemethionineorYangcycle 35
2.7.1 TheYangcycleinrelationtopolyamineand
nicotianaminebiosynthesis 39
2.7.2 RegulationoftheYangcycleinrelationto
ethylenesynthesis 40
2.8 Conclusions 42
Acknowledgement 43
References 44
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viii (cid:2) Contents
3 TheFormationofACCandCompetitionBetweenPolyamines
andEthyleneforSAM 53
SmadarHarpaz-Saad,GyeongMeeYoon,AutarK.Mattoo,and
JosephJ.Kieber
3.1 Introduction 53
3.2 IdentificationandcharacterizationofACCsynthase
activityinplants 54
3.2.1 Historicaloverview 54
3.2.2 PurificationandpropertiesoftheACCsynthase
protein 56
3.3 AnalysisofACCsynthaseatthetranscriptionallevel 58
3.3.1 MolecularcloningofACCsynthasegenes 58
3.3.2 TranscriptionalregulationoftheACCsynthase
genefamily 59
3.4 Post-transcriptionalregulationofACS 62
3.4.1 Identificationandcharacterizationofinteractions
withETO1 62
3.4.2 RegulationofACSdegradation 64
3.5 DoesACCactasasignal? 65
3.6 Biosynthesisandphysiologyofpolyamines 67
3.6.1 SAMisasubstrateforpolyamines 67
3.6.2 Physiologyofpolyamineeffectsinvitroandinvivo 67
3.6.3 Concurrentbiosynthesisofethyleneand
polyamines 70
3.6.4 Doplantcellsinvokeahomeostaticregulationof
SAMlevels? 72
Acknowledgements 72
References 72
4 TheFateofACCinHigherPlants 83
SarahJ.DorlingandMichaelT.McManus
4.1 Introduction 83
4.2 HistoryofthediscoveryofACCoxidaseasthe
ethylene-formingenzyme 84
4.2.1 EarlycharacterizationofACCoxidase 84
4.2.2 Cloningoftheethylene-formingenzymeasan
indicatorofenzymeactivity 85
4.2.3 Initialbiochemicaldemonstrationof
ethylene-formingenzymeactivityinvitro 86
4.3 MechanismoftheACCoxidase-catalyzedreaction 86
4.3.1 InvestigationoftheACOreactionmechanism 87
4.3.2 MetabolismofHCN 89
4.3.3 EvidenceoftheconjugationofACC 91
4.4 TranscriptionalregulationofACCoxidase 92
4.4.1 ACOmulti-genefamilies 92
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Contents (cid:2) ix
4.4.2 DifferentialexpressionofmembersofACO
multi-genefamiliesinresponsetodevelopmental
andenvironmentalstimuli 94
4.4.3 TranscriptionalregulationofACOgeneexpression 96
4.4.4 Crosstalkbetweenethylenesignallingelements
andACOgeneexpression 97
4.5 TranslationalregulationofACCoxidase 97
4.6 EvidencethatACCoxidaseactsasacontrolpointin
ethylenebiosynthesis 100
4.6.1 Cell-specificexpressionofACCoxidase 102
4.6.2 DifferentialexpressionofACSandACOgenes 103
4.7 EvolutionaryaspectsofACCoxidase 104
Acknowledgements 105
References 105
5 PerceptionofEthylenebyPlants–EthyleneReceptors 117
BradM.Binder,CarenChangandG.EricSchaller
5.1 Historicaloverview 118
5.2 Subfamiliesofethylenereceptorsandtheirevolutionary
history 120
5.3 Ethylenebinding 123
5.3.1 Requirementsforametalcofactor 123
5.3.2 Characterizationoftheethylene-bindingpocket
andsignaltransduction 124
5.4 Signaloutputfromthereceptors 126
5.5 Overlappingandnon-overlappingrolesforthereceptor
isoformsincontrollingvariousphenotypes 128
5.6 Post-translationalregulationofthereceptors 131
5.6.1 Clusteringofreceptors 131
5.6.2 Ethylene-mediateddegradationofreceptors 132
5.6.3 RegulatoryroleofREVERSION-TO-ETHYLENE
SENSITIVITY1(RTE1)/GREEN-RIPE(GR) 133
5.6.4 Otherproteinsthatinteractwiththeethylene
receptors 134
5.7 Conclusionsandmodel 135
Acknowledgements 137
References 138
6 EthyleneSignalling:theCTR1ProteinKinase 147
SilinZhongandCarenChang
6.1 Introduction 148
6.2 DiscoveryofCTR1,anegativeregulatorofethylenesignal
transduction 148
6.2.1 IsolationoftheArabidopsisCTR1mutant 148
6.2.2 CTR1mutantphenotypesinArabidopsis 149
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x (cid:2) Contents
6.2.3 PlacementofCTR1intheethylene-response
pathway 150
6.3 CTR1Encodesaserine/threonineproteinkinase 151
6.3.1 Molecularcloningandsequenceanalysisofthe
ArabidopsisCTR1gene 151
6.3.2 CTR1biochemicalactivity 152
6.4 TheCTR1genefamily 153
6.4.1 TheCTRmulti-genefamilyintomato 153
6.4.2 FunctionalrolesoftomatoCTRgenes 153
6.4.3 TranscriptionalregulationofCTR-likegenes 155
6.5 RegulationofCTR1activity 156
6.5.1 PhysicalassociationofCTR1withethylene
receptors 158
6.5.2 MembranelocalizationofCTR1 159
6.5.3 AninhibitoryrolefortheCTR1N-terminus? 159
6.5.4 Otherfactorsthatpotentiallyinteractwithand
regulateCTR1activity 160
6.6 ElusivetargetsofCTR1signalling 161
6.7 CTR1crosstalkandinteractionswithothersignals 162
6.8 Conclusions 163
Acknowledgements 164
References 164
7 EIN2andEIN3inEthyleneSignalling 169
Young-HeeCho,SanghoLeeandSang-DongYoo
7.1 Introduction 169
7.2 OverviewofethylenesignallingandEIN2andEIN3 172
7.3 GeneticidentificationandbiochemicalregulationofEIN2 173
7.4 EIN3regulationinethylenesignalling 174
7.4.1 Geneticidentificationandbiochemicalregulation
ofEIN3 174
7.4.2 Structuralandfunctionalanalysisofein3function 178
7.4.3 FunctionofEIN3astranscriptionactivator 180
7.5 FunctionsofERF1andotherERFsinethylenesignalling 181
7.6 Futuredirections 183
Acknowledgements 184
References 184
8 EthyleneinSeedDevelopment,DormancyandGermination 189
RenataBogatekandAgnieszkaGniazdowska
8.1 Introduction 189
8.2 Ethyleneinseedembryogenesis 192
8.2.1 Ethylenebiosynthesisduringzygotic
embryogenesis 192
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Contents (cid:2) xi
8.2.2 Ethyleneinvolvementintheregulationofseed
morphology 194
8.3 Ethyleneinseeddormancyandgermination 194
8.3.1 Ethylenebiosynthesisduringdormancyrelease
andgermination 194
8.3.2 Theroleofethyleneinseedheterogeneity 199
8.4 Ethyleneinteractionswithotherplanthormonesinthe
regulationofseeddormancyandgermination 199
8.5 EthyleneinteractionswithROSintheregulationofseed
dormancyandgermination 202
8.6 Ethyleneinteractionswithothersmallgaseoussignalling
molecules(NO,HCN)intheregulationofseeddormancy
andgermination 204
8.7 Concludingremarks 207
Acknowledgements 209
References 209
9 TheRoleofEthyleneinPlantGrowthandDevelopment 219
FilipVandenbusscheandDominiqueVanDerStraeten
9.1 Introduction 219
9.2 Designofrootarchitecture 220
9.3 Regulationofhypocotylgrowth 225
9.4 Shootarchitectureandorientation:post-seedlinggrowth 229
9.4.1 Inhibitionofgrowthbyethylene 229
9.4.2 Stimulationofgrowthbyethylene 229
9.4.3 Shootgravitropism 231
9.4.4 Controlofstomataldensityandaperture 231
9.4.5 Activityoftheshootapicalmeristem 231
9.5 Floraltransition 232
9.6 Determinationofsexualformsofflowers 232
9.7 Ethyleneeffectsongrowthcontrollingmechanisms 233
9.8 Conclusions 234
Acknowledgements 234
References 234
10 EthyleneandCellSeparationProcesses 243
ZinniaH.Gonzalez-CarranzaandJeremyA.Roberts
10.1 Introduction 243
10.2 Overviewofthecellseparationprocess 244
10.2.1 Abscission 245
10.2.2 Dehiscence 249
10.2.3 Aerenchymaformation 251
10.2.4 Stomatadevelopmentandhydathodeformation 252
10.2.5 Rootcapcellsloughingandlateralrootemergence 254
10.2.6 Xylemdifferentiation 257