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PlantEnvironmentalStressPhysiologyFanHaiPlantEnvironmentalStressPhy1WhatisPlantStressPlantstressistheadversereactionofplantstoenvironmentalconditionsthatareunfavorabletogrowth,suchaslackofsufficientnutrients,inadequatewatering,flooding,highorlowtemp.,diseaseorinsectinfestation.WhatisPlantStressPlantstre2StressBioticstresspathogenmicrobesinsectsallelopathyAbioticstresswaterTemp.lightdroughtfloodingHightemp.Lowtemp.:chilling,freezingphotoinhibitionUV
damagechemicalsSalinityAlkalinityInsecticide,herbicideAirpollutionStressBioticstresspathogenmi3PlantstressesgreatlydiminishcropyieldsBioticandabioticstressescanreduceaverageproductivityby65%~87%Plantstressesgreatlydiminis4Table:Averageyieldsandrecordyieldsof8majorcropsCropRecordyieldAverageyieldAveragelossesAbioticlosses(%ofrecordyield)BioticAbioticCorn19300460019521270065.8Wheat1450018807261190082.1Soybean73901610666512069.3Sorghum20100283010511620080.6Oat106001720924796075.1Barley114002050765859075.4Potato9410028300177755090054.1Sugarbeet12100042600171006130050.7Table:Averageyieldsandreco5WhatisStrainStrainisthebiologicchangesofplantsundertheinfluenceofplantstress.Straincanbeelasticorplasticdependingonthedegreeandlastingtimeofstress.Elasticstrain:recoverable,temporaryPlasticstrain:non-recoverable,permanentWhatisStrainStrainisthebi6I.DamagesofstresstotheplantstressPrimarydamageSecondarydamagePrimarydirect-damagePrimaryindirectdamage(injurytotheplasmalemma)(dysfunctionofmetabolism)(e.g.waterstressinsaltstress)I.Damagesofstresstothepl7I.DamagesofstresstotheplantBio-membraneChangeinmembranephaseChangeinmembraneconstituentsChangeinmembranepermeabilityI.Damagesofstresstothepl8PhotosynthesisDecreaseinPnDecreaseinstomatalapertureDecreaseinFv/FmDecreaseinquantumyieldPhotosynthesis9RespirationChangeinrespirationpathE.g.TurnEMP-TCApathwaytoPPPunderdrough,pathogenandwoundingstress.Decreaseinrespirationrateunderfreezing,hotandfloodingstressIncreaseinrespirationrateunderwoundingandpathogenstressUnderchillinganddroughtstress,therespirationrateincreasefirstandthendecreaseRespiration10ChangesofphytohormonelevelDecreaseinIAA,CTK,GBSignificantincreaseinABAIncreaseofEthunderfloodingChangesofphytohormonelevel11Oxidativestressinducedbyreactiveoxygenspecies(ROS)Reactiveoxygenspeciesarehighlyreactivemoleculescontainingatomoxygen.Itmainlyinclude4species:O2.-,·OH,H2O2and1O2
ROScanattackchloroplast,mitochondriaandotherorganelles.ROScanalsoattackmembranebycausingmembraneperoxidation.Oxidativestressinducedbyre12II.PlantsRespondtoStressinSeveralDifferentWaysEnvironmentalStressStressresponseToleranceAvoidanceEscapeII.PlantsRespondtoStressi13StressEscape:Plantavoidtheinjuryofstressbyregulatingitslifecycletoavoidmeetingwithstress.E.g.someshort-lived,desertephemeralplants
germinate,growandflowerveryquicklyfollowingseasonalrains.Theythuscompletetheirlifecycleduringaperiodofadequatemoistureandformdormantseedsbeforetheonsetofdryseason.StressEscape:Plantavoidthe14Stressavoidance:plantsavoidtheinjuryofstressbybuildingupabarriertopreventstressfactorsenteringtheplant.E.g.alfafasurvivedryhabitatsbysendingdowndeeprootsystemsthatpenetratethewatertable.Salt-secretionhalophytessecretethesaltsoutfromtheleafthusreducesaltcontentintheleaf.Stressavoidance:plantsavoid15Stresstolerance:Plantsadapttothestressenvironmentbyregulatingtheirmetabolismandrepairthedamagecausedbystress.E.g.HighlysalttoleranthalophytessuchassuaedasalsasurvivesaltyhabitatbymanystrategiessuchashighROSscavengingability,highosmoticadjustmentability,highioncompartmentalizationabilityandsoon.Stresstolerance:Plantsadapt16Adaptationvs.AcclimationAdaptationandacclimationbotharemeansofachievingtolerancetoaparticularstress.Adaptationreferstoheritablemodificationsinstructureorfunctionthatincreasethefitnessoftheorganisminthestressfulenvironment.e.g.CAMplantstodesert.Acclimationreferstononheritablephysiologicalmodificationsthatoccuroverthelifeofanindividual.Thesemodificationsareinducedbygradualexposuretothestress.E.g.slowdryingincreasesthedroughtresistanceofplants.Theprocessofacclimationisknownashardening.Adaptationvs.AcclimationAdap17Howplantsgetadaptedtostress?OsmoticadjustmentStressproteinsScavengeROSSynthesisABACross-adaptationHowplantsgetadaptedtostre181.OsmoticadjustmentWaterdeficitcanbeinducedbymanyenvironmentalconditionsDroughtSalinityColdtemperatureOsmoticadjustmentisabiochemicalmechanismthathelpsplantstoacclimatetodrought,salinityetc.Osmoticadjustmentisanactiveprocessofincreasingthenumberofsoluteparticlesintheplantthatresultinginareducedosmoticandwaterpotentialandenablestheplanttoabsorbwaterindroughtandsalinehabitat.1.OsmoticadjustmentWaterdef19植物逆境生理課件20CompatiblesolutessharespecificbiochemicalattributesHighlysolubleLowmolecularweightDonotinterferewithcellularmetabolismNeutrallychargedatphysiologicalpHMainlydistributedinthecytosol,notvacuoles
Compatiblesolutessharespeci21MostcompatibleosmolytesareAA,QACSorpolyalcoholsMostcompatibleosmolytesare22SomecompatiblesolutesmayserveotherprotectivefunctionsinadditiontoosmoticadjustmentMinimizingtheperturbingimpactofionsProtectingenzymes(pro,glycinebetain)ScavengingROS(pro,mannitol,sorbitol)Somecompatiblesolutesmayse232.StressproteinsHSPs(Heatshockproteins)CRP(Coldregulatedproteins)anaerobicpeptidesLEAs(lateembryogenensisabundantproteins)OsmotinPathogenesis-relatedproteins2.StressproteinsHSPs(Heatsh243.ScavengingROSEnzymes:SOD(superocidedismutase)CAT(Catalase)POD(AscorbicacidPeroxidase)Oxidants:caroteneVitEDMSOVitCGSH3.ScavengingROSEnzymes:254.ABA-stresshormoneKeepmembranestabilityPromotesynthesisofosmolytesReducewaterlossIncreasewaterconductivity4.ABA-stresshormoneKeepmemb26Cross-adaptationCross-adaptationmainsexposureofanorganismtoagivenadverseenvironmentmodifiesitsresponsetootheradversefactors.E.g.exposureofplantstovariousstressessuchaschillingtemperature,heat,drought,andradiationconferredresistancetootherstresses.Cross-adaptationCross-adaptati27Causesofcross-adaptationABAisinducedundermanystresses.Stressinducesmanystressproteins,andstressproteinfunctionsinmanystresses.OsmolytesareinducedbymanystressesMembranechange.ROSscavengingsystemisactivated.Causesofcross-adaptationABA28III.FreezingStressFreezinginjuryiscausedbylowtemp.<0℃SupercoolingIII.FreezingStressFreezingi29IntercellularcrystallizationIcecrystalsformbetweencells.IntracellularcrystallizationIcecrystalsforminthecell.Intercellularcrystallization30FreezingInjury:Directinjury:injurybycrystalformationIndirectinjury:dehydration,suffocatingInjurymechanism:MembraneinjuryMechanicalinjury-SHtheoryFreezingInjury:31植物逆境生理課件32Strategiesofincreasingplantfreezingtolerance:LowerwatercontentReducephotosynthesisIncreaseABA/GBDormantIncreaseosmolytesAlltheabovecanbeattainedbycoldhardeningStrategiesofincreasingplant33IV.ChillingStressCausedbylowtemp.>0℃DamageMembranephaseRootwaterabsorptionabilityDysfunctionofrespiration,accumulationofethanol.DysfunctionofmetabolismIV.ChillingStressCausedbyl34Mechanismofchillinginjury:membranephasetransition:LCphase→→GelphaseMechanismofchillinginjury:35Liquidcrystallinephase.Thetypicalphaseinbiologicalmembranes.ThelipidshavebothlateralandkineticmotionandcontainmembraneproteinsGelphase.ThemembranelipidshavelesskineticenergyandlateralmotionthanintheliquidcrystallinephaseresultinginaregularspacingbetweentheacyltailsLiquidcrystallinephase.The36Strategiesofimprovingplantchillingtolerance:IncreaseIUFA(indexofunsaturatedfattyacid),whichleadstothedecreaseofphasetransitiontemp.Synthesisofchilling-tolerantisoenzymes.Strategiesofimprovingplant37V.DroughtStressDroughtstressincludes:Soildrought,norainforlongtimeandno-availablewaterinthesoil.
Airdrought,RH<20%inatmosphere,transpiration>>waterabsorption.Iflonger,soildroughtoccurs.V.DroughtStressDroughtstres38Symptomsinplantfacingtodrought:stun,redcolorinbase,smallcellandleafarea,leafyellowishandabscission.Youngleavesor/andreproductiveorganswilttodeath.Symptomsinplantfacingtodr39Mechanismsofdroughtstress:MembranedamageMetabolicdisorder:WaterredistributionintheorganPhotosynthesisreductionRespirationrisesthendropsDecreaseinnuclearacidsandproteins
AccumulationofprolineABAaccumulationMechanicaldamageFormationofS-SMechanismsofdroughtstress:40StrategiesofincreasingdroughtresistanceStressavoidance:absorptionofwater,declinationoftranspirationandincreaseinwatertransportation.DevelopedrootsystemandhigherratioofroottoshootThickleaf,smallerleafareaandthickcuticleAccumulationofABAandstomatalcontrolStresstoleranceOsmoticadjustmentChangeofphotosyntheticpathwayStrategiesofincreasingdroug41VI.FloodstressInjuryofflood:(Secondaryinjury)Anaerobicstress:anaerobicrespiration→ethanolEthyleneevolvingVI.FloodstressInjuryoffloo42(1)InjuryinmorphologyandanatomybyO2deficiency:growth↓,leafyellowish(nutritiondeficiency),epinasty(Eth),stemhollow(tissuedegradationcausedbyEth).(2)InjuryinmetabolismbyO2deficiency,photosynthesis↓——stomatalblock,inhibitionofCO2entrance.Anaerobicrespiration↑,toxicants:alcohol,acetaldehyde,NH3,lactate,H2S(1)Injuryinmorphologyanda43(3)Nutritiondisorder:absorption↓,soilN、P、K、CalossbutH2S、Fe、Mn↑,microele
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