植物生理学 英文课件 Chapter 3 Photosynthesis学习资料

植物的营养问题(NutritionofPlant)

土壤营养：Water,MineralelementsFood?植物的营养问题(NutritionofPlant)

土壤营养：Water,Mineralelements空气营养：Photosynthesis

(光合作用)碳素同化作用（carbonassimilation）Chapter3Photosynthesis3.1

Thediscoveryofphotosynthesis3.2

Theimportanceofphotosynthesis3.3

Chloroplastandchloroplastpigments3.4

Mechanismsofphotosynthesis3.5

Factorsaffectingphotosynthesis3.6

Solarenergyutilizationofplants3.7

Translocationofphotosynthetic

products3.1ThediscoveryofphotosynthesisPhotosynthesis:

CO2+H2O（CH2O）+O2

LightGreenplant

1771,JosephPriestly:mint/candle,“oxygen”

1779,JanIngenhousz:Light

1782,J.Senebier：CO2andO2

1804,N.T.deSaussure:H2Oand[CO2]=[O2]

1864,J.Sachs:CH2O(Starchgrain/light)Backcarbohydrate1st3.2TheimportanceofphotosynthesisGreenfactorytoproduceorganiccompounds粮、棉、油/衣、食、住、行

EnergytransformationLight(solar)energyChemicalenergy food,oil,coal,gas,wood.H2SourceofoxygenBreath;Ozonelayer;RecycleofCO2

FoodO2Energy“大海航行靠舵手，万物生长靠太阳”

Rice/noodle/meatphotosyntheticscissorsmagicFutureshapeofhumanbeings?!GreenBack3.3Chloroplast叶绿体

andchloroplastpigments3.3.1Placesofphotosynthesis2ndconceptofphotosynthesis3rdconceptofphotosynthesis3.3.2Morphology,structureandchemicalcomponentsofchloroplastMorphology:flatandellipsoid(椭圆体)Chloroplastcanmovement,functions?Structure:2D被膜=外+内间质类囊体基粒类囊体间质类囊体Lipiddroplet脂滴Structure:3DChemicalcomponents:Mainchemicalcompound?Proteins:(onlamellae)proteincomplex,plastocyanin(质体蓝素PC)Water(75%)(instroma)enzymes(Rubisco,etc)Lipids:glycosylglyceride糖基甘油酯;fewP-,S-DNAandRNA:Minerals:Mg,etcCarbohydrates:starch,etcPigments:8%ofdryweightofchloroplastAbsorblightSemiautonomousorganelles蛋白质灰分贮藏物质脂类色素?Chlorophylls

叶绿素Carotenoids

类胡萝卜素Phycobilins

藻胆素3.3.3ChemicalcharacteristicsofphotosyntheticpigmentsChlorophylls:color,solubility,structure

Chla:C55H72O5N4Mg;blue-green

Chlb:C55H70O6N4Mg;yellow-greenCH3叶绿酸(双羧酸)的脂pyrrole吡咯环副环porphyrin卟啉环共轭电子系统C20H39ChlaHPolarity?Chlb﹤CHOCuCarotenoids:color,solubility,structure

carotene胡萝卜素:C40H56;orange-yellow；abg

xanthophyll/carotenol叶黄素:C40H56O2;yellowPolarity?b-caroteneXanthophyll/Carotenol紫罗兰酮环OHOHCarotenol

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CarotenePhycobilins/Phycobiliproteins:color,structure

phycoerythrin:red

phycocyanin:bluephycoerythrinphycocyanin研究叶绿体色素而获诺贝尔奖者威尔施泰特(德)1915：叶绿素是由叶绿醇和含镁的叶绿酸所形成的酯。研究叶绿体色素而获诺贝尔奖者费歇尔(德)1930：研究卟啉类化合物，其中包括叶绿素和铁血红素，经费歇尔修订的叶绿素分子结构一直沿用至今。制成人造血红素。研究叶绿体色素而获诺贝尔奖者卡勒因(瑞士)1937：研究类胡萝卜素等物质的化学结构。研究结果对研究类胡萝卜素这一天线色素的光合特性具有重要作用。研究叶绿体色素而获诺贝尔奖者伍德沃德(美)1965：于1960年合成叶绿素3.3.4OpticalcharacteristicsofphotosyntheticpigmentsIrradiationenergy:energyoflight Lighthascharacteristicsofbothaparticle(photon)andawave.Planck’slaw:E(quantum)=hv=hc/l

Absorptionspectrum吸收光谱:Solarspectrum:300-2600nm?Visiblespectrum:400-700nmAbsorptionspectrumLight连续光谱吸收光谱LightAbsorptionspectrumcurveChlaChlbPhycoerythrinb-caroteneBacteriochlorophyllaPeakandotherwavelengthsFluorescence荧光现象

and

Phosphorescence磷光现象Fluorescence:excitedchlorophyllcanre-emitaphotonwhenitreturnstoitsgroundstate.Invitro/solution: greenintransmission,redinreflection?Invivo/livingcell:?Phosphorescence:第一三线态

groundstateCH3吡咯环副环卟啉环共轭电子系统C20H393.3.5Biosynthesisandbreakdownofchlorophyllarecomplexpathways谷氨酸MgFeChlorophyllFerroheme亚铁血红素胆色素原x2原卟啉IXPlantAnimal原脱植基叶绿素aChlb脱植基叶绿素a叶绿醇（植醇）叶绿素的降解叶绿素b还原酶类囊体内的叶绿素b

叶绿素a

叶绿素酶脱镁整合酶

脱植基叶绿素a

脱镁叶绿素a氧化酶脱镁叶绿素a

水溶性无色产物3.3.6BiosynthesisofcarotenoidsChlorophylls/carotenoids=3Chla/Chl

b=3Xanthophyll/Carotene

=2??anthocyanin?Breakdown3.3.7LeafcolorLight:?EtiolationTemperature:?Mineralelements:N:?Mg:?Fe:?Mn:?Cu:?Zn:??Back3.3.8FactorsaffectingchlorophyllbiosynthesisChlorosis缺绿3.4Mechanismsofphotosynthesis“太阳公公为什么能把树晒大呢？”

“太阳公公为什么能把树晒大呢？”

Solarenergy

Electricenergy Absorption,transferandconversionofsolarenergy

ElectricenergyActivechemicalenergy(ATP/NADPH) Electrontransportandphotophosphorylation

ActivechemicalenergyStablechemicalenergy(carbohydrate) Carbonassimilation同化3.4Mechanismsofphotosynthesis3.4.1AbsorptionandtransferofsolarenergymostChla;allChlb,carotenoidsandphycobilinsLight-harvesting/Antennapigments:Reactioncenterpigments:fewspecialChlaResonance共振photon

3.4.2ConversionofsolarenergytoelectricenergyReactioncenter:Reactioncenterpigments:P700/P680(Chla)PrimaryelectrondonorandacceptorPhotochemicalreactionDonor.P.AcceptorLightDonor.P*.AcceptorDonor.P.Acceptor-Donor.P+.Acceptor-Donor+.P.Acceptor-LightenergyElectricenergyRedoxReactionReduction/Oxidation

Twophotosyntheticunit/Photosystem(PS)Twophotosystemsworkintandem接力Howweretwophotosystemsdiscovered?Reddrop:Emerson,bythelate1950s.680nmEnhancementeffect/EmersoneffectBytwophotosystemsworkingintandem接力Emerson,1957:700nm680nmTwophotosystemsworkintandem接力Structure:2D被膜=外+内间质类囊体基粒类囊体间质类囊体Lipiddroplet嗜饿滴/脂滴Organizationoftheproteincomplexesofthethylakoidmembrane3.4.3ElectronandprotontransportOriginaledonor:WaterEndeacceptor:NADP+ZschemePhotosyntheticchainCyclic-Noncyclic-autonomously?StromaLumenPSIIComplexes=CoreComplexPSIIlight-harvestingcomplex:LHCIIOxygen-evolvingcomplex:OECOECLocation:lumensideComponents:peptides(33,23,18kD)

Mncomplex,Ca+,Cl-Oxygenevolvingmechanism(Wateroxidizingclock)2H2OO2+4H++4e-CO2+2H2O*（CH2O）+O*2+H2OlightchloroplastNewconceptofphotosynthesis2H2OO2+4H++4e-CO2+2H2O*（CH2O）+O*2+H2OlightchloroplastNewconceptofphotosynthesis2H2OO2+4H++4e-CO2+2H2O*（CH2O）+O*2+H2OlightchloroplastNewconceptofphotosynthesisPSIComplexes德国科学家戴森霍弗、胡贝尔、米歇尔测定出了光合作用反应中心膜蛋白-色素的三维空间结构，揭示出电子和能量的传递作用，共同获得1988年度诺贝尔奖。3.4.4PhotosyntheticphosphorylationPmfH+H+H+H+H+H+H+Photophosphorylation光合磷酸化ATPSynthaseorCouplingfactorProductsofCyclic/NoncyclicElectronTransportATP,Oxygen,NADPH?OxygenNADPHATPAssimilatorypower同化力3.4.5Carbonassimilation同化Location:stromaofchloroplast3pathways: CalvinCycle/C3pathway C4pathway CAM3.4.5.1TheCalvincycle(M.Calvin)PhotosyntheticcycleReductivepentose戊糖

phosphatepathway(RPPP)C3pathway14CO2Paperchromatograph层析AutoradiogramsshowingtheproductsofphotosynthesisinthealgaChlorellapyrenoidosaaftervarioustimesofexposureto14CO2(bottom)2s,(middle)7s,(top)60s.Notetheincreasingimportanceof3-PGA(3-phosphoglycerate,3-磷酸甘油酸)andothersugarphosphatesastheexposuretimeisshortenedRuBP:Ribulose-1,5-bisphosphate核酮糖-1,5-二磷酸Carboxylationinorganic-organicx6RuBPCaseReductionx6x1x5x3激酶6x磷酸丙糖3CRegeneration4C5C6C3C

x53C6C7C7C5C5Cx1x1x1x1x13C5C5C5C5Cx3赤藓糖景天庚酮糖木酮糖核糖3x3x6xx66x5x3xGlyceraldehyde-3-phosphate1xFoodofplants“太阳公公为什么能把树晒大呢？”

RegulationoftheCalvincycleLight-dependentenzymeactivationregulatestheCalvincycleLight-dependentionmovementsregulatetheCalvincycleLightreaction??? 1stand2ndmainstepsDarkreaction??? 3rdmainstep3.4.5.2TheHatch-SlackPathwayDiscoveredbyHatchandSlackfromAustraliain1966,insometropicalandsub-tropicalplants,e.g.cereals,whichhavespecialleafanatomic解剖

featuresC4monocot,maizeC3monocot,riceC4-pathway:C4-dicarboxylicacidpathwayC4-plantsC3-plantschloroplastKranz=wreathchloroplastcontaining维管束鞘Manyplasmodesmata胞间连丝

betweenmesophyllandbundlesheathcells碳酸酐酶TheHatch-SlackPathway（C4pathway）草酰乙酸苹果酸PEPCase

丙酮酸4CPEPPEPC2xATP3.4.5.3CAM(Crassulacean

AcidMetabolism)NightDay景天酸代谢Open?Close?NightStomatamovementOpen?Close?DayCAMplantsProblem???4Chigh[organicacid]low[organicacid]C4Plants:?C3Plants:?CAMPlants:?MaizeRiceCrassulaceaeDifferentplanttypesclassifiedaccordingtophotosyntheticpathways3.4.5.4PhotorespirationC2OxidativePhotosyntheticCarbonCycleDarkrespirationPathway(inthreeorganelles):Concept:RuBP磷酸乙醇酸RuBP

Oxygenase

RubiscoRuBPC2C3CCO2CHLOROPLAST乙醛酸PEROXISOME[CO2]/[O2]CO2BiologicalFunctionofPhotorespirationUnderhighlightintensityandlowintercellularCO2concentration,dissipateexcessATPandreducingpowerfromthelightreactions.IsUnknownBack3.5FactorsaffectingphotosynthesisPhotosyntheticrate:mgCO2/dm2/h;µmolCO2/m2/sApparentphotosyntheticrate=NetphotosyntheticrateTruephotosyntheticrate=NPR+Respirationrate3.5.1Light

Lightintensity:µmol/m2/s光补偿点Lightsaturationpoint???合理密植?Xhighersaturationpointlowerlightcompensationpoint滨藜属，藜科细辛属，马兜铃科

Photoinhibition:Mechanismisunclear?LightofdifferentwavelengthCloudy:moreblueandgreenlightUndercanopy:moregreenlightUpperwater:greenalgaeDeepwater:moregreenandbluelight,redalgaePhycoerythrin3.5.2CO2CO2saturationpoint???>>

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CO2intheatmosphereCO2compensationpoint:C4<C3CO2intheatmosphere:360ppmatpresentGreenhouseeffectWaterplants?3.5.3TemperatureEnzymeactivitiesApparentphotosyntheticrate?Respiration3.5.4WaterMaterial:1%StomamovementTransportation3.5.5Essentialmineralelements

Iron---FeManganese---MnBoron---BCopper---CuZinc---ZnMolybdenum---MoChlorine---ClNickel---NiSodium---NaNitrogen---NPhosphorus---PPotassium---KCalcium---CaMagnesium---MgSulfur---SSilicon---SiWhyphotosyntheticrateofC4plantsismuchmorehigherthanthatofC3plants?

Specialleafanatomicfeatures:KranzPhysiologicalfeatures:Affinity亲和性

toCO2:PEPC>RuBPCCO2compensationpoint补偿点:C4<C3CO2pump:C4

Photorespiration:C4<C3Back3.6Solarenergyutilizationofplants3.6.1Efficiencyforsolarenergyutilization3.6.2Howtoincreasephotosyntheticrate?elongatephotosynthetictime:temporalandspatial

increasephotosyntheticarea:

合理密植、改变株型increaseCO2concentration:aeration,CO2fertilizerdecreasephotorespiration?BackPhotosynthesislocation:leavesConsumedplaces:alllivingcells

Storageorgans:seed,fruit,tuber,rhizoma,3.7TranslocationofphotosyntheticproductsSource源:cells,tissues,organs,produceandexportphotosyntheticproducts.leafSink库:?stemetc.Thetransitionoftheleaffromsinktosourcestatustranslocation3.7.1PathwaysoftranslocationWaterandmineralsaretransportedin???Photosyntheticproductsaretranslocatedinphloem!Howdoyouknow?

Girdling环割OnthestemsofbranchesOnthemainstem?Hibiscusrosa-sinensis朱槿（大红花）Onthemainstem?人要脸，树要皮？Howdoweknow?

Girdling环割ApplicationinfruitproductionHowdoweknow?

Girdling环割ApplicationinseedlingproductionCalliandra

haematocephala朱缨花

Isotopetracing同位素示踪Isotopetracing+surgicaltreatmentBidirectionaltransportationSievetubeelementsandcompanioncellsarelivingcells!3.7.2Typeofphotosyntheticproductstranslocated????????????Whattype?Howdoyouknow?Aphid蚜虫styletChemicalcompositionofphloemsap:Water:Drysubstances:Sucrose:90%ofdryweightRaffinose,StachyoseandVerbascose:few棉子糖水苏糖毛蕊花糖半乳糖Aminoacids:Hormones:Inorganicions:K,P,Cl,etc.ProteinsOrganicacids3.7.3.1Phloemloading3.7.3.2Translocationinsieveelements3.7.3.3Phloemunloading3.7.3Detailtranslocationsteps3.7.3.1Phloemloading

Inmesophyll/bundlesheathcells:ChloroplastCytosolMesophyll/bundlesheathcellsSieveelements

Frommesophyll/bundlesheathcellsPhloemparenchymacellsFromphloemparenchymacellsSieveelementsHowdoessucrosei