第五節(jié)-植物的光形態(tài)建成-植物生理學(xué)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(yīng)（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(xué)課件3第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)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(xué)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(xué)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(xué)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(xué)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(xué)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(xué)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(xué)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(yuǎn)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(xué)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(yīng)類型反應(yīng)類型縮寫所需光強(qiáng)光受體光可逆性

實(shí)例低輻照度反應(yīng)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(dòng)極低輻照反應(yīng)VLFR10-4~10-2μmolm-2光敏素，隱花色素向光色素?zé)o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(yīng)HIR10~105μmolm-2光敏素，隱花色素向光色素?zé)o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(yīng)類型反應(yīng)類型縮寫所需光強(qiáng)光受體光可逆性實(shí)45LFR&HIRactionspectrumfortheinhibitionofhypocotylelongationofdark-grownlettuceseedlingsLFR&HIRactionspectrumforth46Phytochromeisanautophosphorylating

serine/threoninekinasePhytochromeisanautophosphor47第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件48第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件49第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)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(xué)課件55第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)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(xué)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(xué)課件62植物的避蔭反應(yīng)

shadeavoidance植物的避蔭反應(yīng)

shadeavoidance63第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件64第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)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(xué)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(xué)課件71degradationdegradation72第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件73MutantsHelpsRevealTheMechanismofShadeAvoidanceMutantsHelpsReveal74第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件75第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件76第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)課件77第五節(jié)-植物的光形態(tài)建成——植物生理學(xué)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