第五節(jié)-植物的光形態(tài)建成-植物生理學課件

第五節(jié)植物的光形態(tài)建成光形態(tài)建成的概念(photomorphogenesis)向光色素(phototropin)隱花色素(cryptochrome)光敏素(phytochrome)避蔭反應（shadeavoidance）第五節(jié)植物的光形態(tài)建成光形態(tài)建成的概念(photomorp1

自然界中的光Wavelength(nanometers)700600500400

自然界中的光Wavelength(nanometers)2第五節(jié)-植物的光形態(tài)建成——植物生理學課件3第五節(jié)-植物的光形態(tài)建成——植物生理學課件4植物的光形態(tài)建成（photomorphogenesis）植物的光形態(tài)建成（photomorphogenesis）5Light-grownArabidopsisLight-grownArabidopsis6第五節(jié)-植物的光形態(tài)建成——植物生理學課件7EtiolatedcharacteristicsDe-etiolatedcharacteristicsDistinct"apicalhook"(dicot)orcoleoptile(monocot)NoleafgrowthNochlorophyllRapidstemelongation

LimitedradialexpansionofstemLimitedrootelongation

Limitedproductionoflateralroots

ApicalhookopensorcoleoptilesplitsopenLeafgrowthpromoted

ChlorophyllproducedStemelongationsuppressed

RadialexpansionofstemRootelongationpromoted

Lateralrootdevelopmentaccelerated

EtiolatedcharacteristicsDe-et8LightControlsPhotosynthesisRateofGrowthDirectionofGrowthPigmentationFloweringFruitingLeafFallOnsetofDormancyLightControlsPhotosynthesis9光敏素隱花色素向光蛋白光敏素隱花色素向光蛋白10第五節(jié)-植物的光形態(tài)建成——植物生理學課件11Phototropins

向光素Phototropins

向光素12第五節(jié)-植物的光形態(tài)建成——植物生理學課件13擬南芥的phot突變體擬南芥的phot突變體14第五節(jié)-植物的光形態(tài)建成——植物生理學課件15擬南芥的phot突變體擬南芥的phot突變體16PHOT1andPHOT2mediate

bluelight-dependentstomatalopeningKinoshitaetal.2001Nature:414,656PHOT1andPHOT2mediate

blue17PhototropinproteinscontaintwoLOV(Light,Ogen,Voltagesensitive)domainsandadownstreamserine/threoninekinase.ThechromophoreisaFlavinmononucleotide(FMN)whichisnon-covalentlyboundwithineachLOVdomain.Phototropinproteinscontain18第五節(jié)-植物的光形態(tài)建成——植物生理學課件19SchematicillustrationoftheLOVandkinasedomainsofphototropininlight-inducedphosphorylation.

(a)Inthedark,thephosphorylationactivityofthekinasedomainofphototropinisinhibitedbythebindingoftheLOV2domain.(b)Underlightconditions,thephosphorylationactivityofthekinaseisinducedbythereleaseoftheLOV2domain.

Volume9,Issue5,October2006,Pages503-508

Schematicillustrationofthe20第五節(jié)-植物的光形態(tài)建成——植物生理學課件21Cryptochromes隱花色素http://./Research/cryptochrome/Cryptochromes隱花色素http://.ks.ui22Cryptochromeandtheavian"pass"Cryptochromeandtheavian"pa23TheHY4genemaydefineabluelightphotoreceptor

CRYPTOCHROME1(CRY1)TheHY4genemaydefineablue24Phenotypesofcry2mutantcyr2islatefloweringunderlongdayconditionsPhenotypesofcry2mutantcyr225ThestructureofcryptochromesThestructureofcryptochromes26Cryptochromecontrolleddeetiolation.InArabidopsis,cry1andcry2regulatemostoftheblue-light-specificdevelopmentalprograms.Shownareonlyphenotypiceffectsduringdeetiolationcausedbymutationsincry1orcry2.Arabidopsiswildtype(WT)andphotoreceptormutantsgrownfor3daysindarkness(D),whitelight(WL)orcontinuousbluelightof30μmolm-2s-1(highblue,HB)or1μmolm2s1(lowblue,LB),respectivelygivenfromthetop.Inwildtypeseedlings,hypocotylgrowthisinhibitedbybluelightwhereasopeningofthehypocotylhookandcotyledonopeningandexpansionispromotedbybluelight.HBismoreefficientthanLB.Thelackofcry1(cry1)ismostevidentunderHBconditions,whereasthelackofcry2(cry2)beesmoreevidentunderLBconditions,whichisseeninparticularforthedoublemutant(cry1/cry2)(takenfromBatschaueretal.,2007).Cryptochromecontrolleddeetio27第五節(jié)-植物的光形態(tài)建成——植物生理學課件28http://./content/102/34/12270.full.pdf+htmlhttp://./content/102/329Phytochrome

光敏素Phytochrome

光敏素30不同波長的光對萵苣種子發(fā)芽的影響不同波長的光對萵苣31萵苣種子發(fā)芽的作用光譜萵苣種子發(fā)芽的作用光譜32LettuceseedgerminationisatypicalphotoreversibleresponsecontrolledbyphytochromeLettuceseedgerminationisa33影響‘GrandRapid’萵苣種子發(fā)芽的光可逆性照射程序發(fā)芽率(%)無(黑暗對照組)14#4*R7098R-Fr

62R-Fr-R7497R-Fr-R-Fr60R-Fr-R-Fr-R7695R-Fr-R-Fr-R-Fr71R-Fr-R-Fr-R-Fr-R8198R-Fr-R-Fr-R-Fr-R-Fr7.#：Tooleetal.,1953(發(fā)芽溫度27℃)

*：BewleyandBlack,1982。(發(fā)芽溫度24℃，640-680nm的紅光，0.9W/m2，每次照1.5min；>710nm的遠紅光，2.9W/m2，每次照4min)影響‘GrandRapid’萵苣種子發(fā)芽的光可逆性照射程序34Photoreversibility:

adistinctivefeatureofphytochromePhotoreversibility:

adistinc35chromophoreapoproteinholoproteinchromophoreapoproteinholoprote36SteroStructureofPhytochromeSteroStructure37第五節(jié)-植物的光形態(tài)建成——植物生理學課件38Chromophorestructure15105Chromophorestructure1510539PfrPrtransisomercisisomerStructureconversionofthe

chromophore(phytochromobilin)PfrPrtransisomercisisomerStr40光敏素蛋白的兩種構(gòu)象模型PfrPr光敏素蛋白的兩種構(gòu)象模型PfrPr41PhytochromeisencodedbyamultigenefamilyPhyA→

responsestoFRPhyB→

responsestoRorwhitelightPre-dominantformsHaveuniquerolesinregulatingresponsestoredandfar-redPhytochromeisencodedbyamu42PhytochromesindifferentspeciesArabidopsis:PHYA,PHYB,PHYC,PHYD,PHYETomato:PHYA,PHYB1,PHYB2,PHYE,PHYFCottonwood（棉白楊）:PHYA,PHYB1,PHYB2Norwayspruce（挪威云杉）:atleast5PHYgenesPhytochromesindifferentspec43光敏素類型及其基因類型編碼基因存在含量Pfr穩(wěn)定性作用PIPHYA黃化組織高低去黃化等PIIPHYB/C黃化組織綠色組織低高HIR等光敏素類型及其基因類型編碼基因存在含量Pfr作用PIPHYA44植物光敏素反應類型反應類型縮寫所需光強光受體光可逆性

實例低輻照度反應LFR1~1000μmolm-2光敏素R/FR可逆萵苣種子萌發(fā)擬南芥種子萌發(fā)轉(zhuǎn)板藻葉綠體轉(zhuǎn)動極低輻照反應VLFR10-4~10-2μmolm-2光敏素，隱花色素向光色素無燕麥胚芽鞘生長和中胚軸伸長抑制擬南芥種子萌發(fā)高輻照度反應HIR10~105μmolm-2光敏素，隱花色素向光色素無幼苗下胚軸伸長抑制彎鉤伸直花青素合成植物光敏素反應類型反應類型縮寫所需光強光受體光可逆性實45LFR&HIRactionspectrumfortheinhibitionofhypocotylelongationofdark-grownlettuceseedlingsLFR&HIRactionspectrumforth46Phytochromeisanautophosphorylating

serine/threoninekinasePhytochromeisanautophosphor47第五節(jié)-植物的光形態(tài)建成——植物生理學課件48第五節(jié)-植物的光形態(tài)建成——植物生理學課件49第五節(jié)-植物的光形態(tài)建成——植物生理學課件50Nuclearlocalizationofphy–GFPfusionproteinsinepidermalcellsofArabidopsishypocotylsPHYA:GFPPHYB:GFPContinuousfar-redlightWhitelightCanmoveinProrPfrformfasttransport(~15min)

maximaltransportundercontinuousFRlight(HIR)MovesonlyinPfrformSlowtransport(severalhours)

RequiresRfortransport;inhibitedbyFRUndercircadiancontrolNuclearlocalizationofphy–GF51PIF3

phytochrome-interactingfactor3helix-loop-helixproteintranscriptionfactorsPIF3helix-loop-helixproteint52PIFlocalizationPIFlocalization53CACGTG

GTGCAC光調(diào)節(jié)元件（LRE）CACGTG

GTGCAC光調(diào)節(jié)元件（LRE）54第五節(jié)-植物的光形態(tài)建成——植物生理學課件55第五節(jié)-植物的光形態(tài)建成——植物生理學課件56BothPHYTOCHROMEINTERACTINGFACTOR1(PIF1)andSPATULA(SPT)inhibitgerminationbyregulatingthelevelofactivegibberellicacid(GA).TheyactastranscriptionalrepressorsofGAbiosyntheticgenes(GA3ox1andGA3ox2),whicharerequiredtoconverttheGAprecursortoitsactiveform.Inaddition,PIF1activatestheexpressionofaGAcatabolicgene(GA2ox2).Underinductivelightconditions(inwhicharelativelyhighred:far-redratioconvertssufficientphytochromeBtotheactive,nuclear-localizedPfrform),phytochromebindingtriggersthedegradationofPIF1throughubiquitin-mediatedproteolysis,therebyblockingtherepressionofGAlevels.Similarly,coldtreatmentrepressesSPTactivitytopromotegermination.BothPHYTOCHROMEINTERACTINGF57第五節(jié)-植物的光形態(tài)建成——植物生理學課件58ConstitutivelyPhotomorphogenic(COP)mutantsPhotomorphogenicresponseisalwaysonincopmutants:ConstitutivephotomorphogenicphenotypeinthedarkHypersensitivephenotypeinthelightConstitutivelyPhotomorphogeni59COP1encodesanE3ubiquitinligaseCOP10encodesanE2ubiquitinligasevarianttheother8encodemembersofaplexknownastheCOP9signalosome(CSN).someCSNponentsresemblecomponentsofthe26Sproteasome10copmutantsidentifiedCOP1encodesanE3ubiquitinl60LightsignalingpathwayLightsignalingpathway61第五節(jié)-植物的光形態(tài)建成——植物生理學課件62植物的避蔭反應

shadeavoidance植物的避蔭反應

shadeavoidance63第五節(jié)-植物的光形態(tài)建成——植物生理學課件64第五節(jié)-植物的光形態(tài)建成——植物生理學課件65ShadeAvoidanceSyndrome(SAS)ShadeAvoidanceSyndrome(SAS)66ThereducedR:FRratiooflightactsasasignalthatotherplantsarenearbyThereducedR:FRratioofligh67第五節(jié)-植物的光形態(tài)建成——植物生理學課件68PlantsofDaturaferoxgrowinginfrontofselectivereflectingmirrorsplacedatca.6cmsouth(southernhemisphere)oftheplants.ThoseinfrontofthemirrorsselectivelyreflectingFRshowincreasedinternodeelongation.KnowTheNeighborthroughPhytochromePlantsofDaturaferoxgrowing69PhotomorphogenesisUnderContinuousLightPhotomorphogenesis70第五節(jié)-植物的光形態(tài)建成——植物生理學課件71degradationdegradation72第五節(jié)-植物的光形態(tài)建成——植物生理學課件73MutantsHelpsRevealTheMechanismofShadeAvoidanceMutantsHelpsReveal74第五節(jié)-植物的光形態(tài)建成——植物生理學課件75第五節(jié)-植物的光形態(tài)建成——植物生理學課件76第五節(jié)-植物的光形態(tài)建成——植物生理學課件77第五節(jié)-植物的光形態(tài)建成——植物生理學課件78Rapidsynthesisofauxinviaanewtryptophan-dependentpathwayisrequiredforshadeavoidanceinplantsYiTao,1,2

Jean-LucFerrer,3,4

KarinLjung,5

FlorencePojer,1,4

FangxinHong,1,2,6JeffA.Long,2

LinLi,2

JavierE.Moreno,7

MarianneE.Bowman,1,4

LaurenJ.Ivans,2,8

YoufaCheng,8

JasonLim,1,2

YundeZhao,8

CarlosL.Ballaré,7

G?ranSandberg,9

JosephP.Noel,1,4

andJoanneChory1,2*1HowardHughesMedicalInstitute2PlantBiologyLaboratory,TheSalkInstituteforBiologicalStudies,LaJolla,CA92037,USA3InstitutdeBiologieStructuraleCEA-CNRS-UJF,LaboratoiredeCristallographieetCristallogenèsedesProtéines,41RueJulesHorowitz,38027GrenobleCedex1,France4TheJackH.SkirballCenterforChemicalBiologyandProteomics,TheSalkInstituteforBiologicalStudies,LaJolla,CA92037USA5Ume?PlantScienceCentre,DepartmentofForestGeneticsandPlantPhysiology,SwedishUniversityofAgriculturalSciences,SE-90183Ume?,Sweden6DepartmentofBiostatisticsandputationalBiology,Dana-FarberCancerInstitute,HarvardSchoolofPublicHealth,44BinneyStreetBoston,MA02115,USA7InstitutodeInvestigacionesFisiológicasyEcológicasVinculadasalaAgricultura(IFEVA),ConsejoNacionaldeInvestigacionesCientíficasyTécnicas,andUniversidaddeBuenosAires,AvenidaSanMartín4453,C1417DSEBuenosAires,Argentina8DivisionofBiologicalSciences,SectionofCellandDevelopmentalBiology,UniversityofCa