植物间的相互作用.ppt

Plant-plantinteractions,PhotocourtesyofRonaldPierik,Plantssenseandrespondtootherplants,Often,butnotalways,plantscompetewitheachotherforlimitingresources,suchaslightandnutrientsHowdotheyperceivecompetitors?Arealloftheirinteractionscompetitive?Howdointeractionsbetweenplantsaffecthigherorganizationallevels(e.g.,communities)?,Photocredit:TomDonald,Outline,KeydefinitionsandconceptsCompetitionCompetitionforlightCompetitionbelowgroundDoplantsperceive“self”or“kin”?Cooperation/FacilitationEnvironmentalmodulationEnhancednutrientavailabilityStresscuesPuttingknowledgetowork,Photocredit:MaryWilliams,Keydefinitionsandconcepts,ReprintedbypermissionfromWileyfromDrew,M.C.(1975).Comparisonoftheeffectsofalocalisedsupplyofphosphate,nitrateandammoniumandpotassiumonthegrowthoftheseminalrootsystem,andtheshoot,inbarley.NewPhytol.75:479-490.ReprintedfromVandenbussche,F.,etal.(2005).Reachingoutoftheshade.Curr.Opin.PlantBiol.8:462-468withpermissionfromElsevier,ReprintedfromSultan,S.E.(2000).Phenotypicplasticityforplantdevelopment,functionandlifehistory.TrendsPlantSci.5:537-542withpermissionfromElsevier.SeealsoBradshaw,A.D.(1965).Evolutionarysignificanceofphenotypicplasticityinplants.Adv.Genet.13:115155.,Thecapacityofanindividual(oragenotype)toexhibitarangeofphenotypesinresponsetovariationintheenvironment,Phenotypicplasticity,Phenotypicplasticityinanimals,ReprintedbypermissionfromMacmillanPublishersLtd.fromAgrawal,A.A.,Laforsch,C.andTollrian,R.(1999).Transgenerationalinductionofdefencesinanimalsandplants.Nature.401:60-63;ReprintedbypermissionfromMacmillanPublishersLtd.FromWarner,D.A.andShine,R.(2008).Theadaptivesignificanceoftemperature-dependentsexdeterminationinareptile.Nature.451:566-568;SeealsoCoker,R.E.(1939).TheproblemofcyclomorphosisinDaphnia.Quart.Rev.Biol.14:137-148,andCrews,D.andBull,J.J.(2008).Sexdetermination:Somelikeithot(andsomedont).Nature.451:527-528.,Temperature-dependentsexdeterminationinreptilesandfish,Predator-inducedhelmetformationinDaphniacucullata,Phenotypicplasticityinplants,ReprintedbypermissionfromWileyfromDrew,M.C.(1975).Comparisonoftheeffectsofalocalisedsupplyofphosphate,nitrateandammoniumandpotassiumonthegrowthoftheseminalrootsystem,andtheshoot,inbarley.NewPhytol.75:479-490.PhotocreditMichaelClayton.,Rootplasticityinresponsetolocalizednutrientavailability,Shootplasticityinresponsetolight,AdaptedfromSanchez,P.,Nehlin,L.andGreb,T.(2012).Fromthintothick:majortransitionsduringstemdevelopment.TrendsPlantSci.17:113-121.ReprintedbypermissionfromMacmillanPublishersLtd.fromTsiantis,M.andHay,A.(2003).Comparativeplantdevelopment:thetimeoftheleaf?NatRevGenet.4:169-180.,Plantmorphologyishighlymodular,Eachmodulebehavesquasi-independently,althoughintegrationbetweenthemallowselaboratecoordinatedresponsesoftheentireplanttochanginggrowthconditions,Intheshoot,eachmoduleisaphytomerthatincludesanode,internode,axillarybudandleaf,Phenotypicplasticityinvolvesalternativeoutcomes,AdaptedfromNovoplansky,A.(2002)Developmentalplasticityinplants:implicationsofnoncognitivebehavior.EvolutionaryEcology16:177188.,Noteverydevelopmentalchangeisconsideredphenotypicplasticity.Plasticdevelopmentisallaboutalternatives,Phenotypicplasticity:CanbeobservedAdaptivephenotypicplasticity:Initially,onlyahypothesis.Whetherobservedplasticityisadaptivecannotbeassumedandhastobedemonstrated,Developmentaldifferentiation,PLASTICITY,Geneticdifferentiation,Metamorphosis,Notallplantsdisplaythesamerangeofphenotypicplasticities,Differentspeciesorvariants(differentgenotypes),Thetype(morphology,architecture,physiologyetc.)andhierarchyofphenotypicplasticityarevariable,Sometraitsaremoreplasticthanothers,AdaptedfromClausenetal.,1940,1948,ascitedinBradshaw,A.D.(1965).Evolutionarysignificanceofphenotypicplasticityinplants.Adv.Genetics13:115155.ImagesfromFTDandChinesebonsaigarden.,Allometry,ReprintedbypermissionofOxfordUniversityPressfromGayon,J.(2000).Historyoftheconceptofallometry.Amer.Zool.40:748-758,imagefromChampy,C.1924.Sexualitethormones.Doin,Paris.Enquist,ImageJournalofHeredity(1921)Volume12,pg421.PlantallometryimageadaptedfromZens,M.S.andWebb,C.O.(2002).Sizinguptheshapeoflife.Science.295:1475-1476,seealsoB.J.andNiklas,K.J.(2002).Globalallocationrulesforpatternsofbiomasspartitioninginseedplants.Science.295:1517-1520;,Someproportionschangeduringdevelopment,MaleDynastesbeetles:hornsgrowfasterthanbodies,Humans:Proportionalsizeofheaddecreaseswithdevelopmentalage,Allometrycanbeplastic:Environmentalconditionsorgeneticchangecancauseadeviationfromstandardallometry,Thestudyoftherelativesize,scaleorgrowthrateofparts,andtheconsequencesofallometricscaling,Signalsandcues,Signalsaregenerallyconsideredtobeanintendedinformationbroadcastandcaninclude:hormones,bacterialmetabolites,electricalsignals,lightquality,stresssignalsCuesareconsideredtobewithoutintentandcaninclude:Nutrientresources,water,light,Signalsandcuesconveyinformation,Semaphoresaresignals,Seeforexample,Aphalo,P.J.andBallare,C.L.(1995).Ontheimportanceofinformation-acquiringsystemsinplant-plantinteractions.Funct.Ecol.9:5-14.Aphalo,P.J.,Ballar,C.L.andScopel,A.L.(1999).Plant-plantsignalling,theshade-avoidanceresponseandcompetition.J.Exp.Bot.50:1629-1634.ImagefromNesnad.,Signalsandcuesaffectingplants,INFORMATION,InternalSignaling,PrimarymetabolitesHormonesElectricalpulses,ExternalCueingSignaling,Bioticonly,Abiotic:Physicalandchemicalenvironmentatmospheric,edaphic.Biotic:Secretions,exudates,volatilesetc,Externalcueing:CanbeobservedExternalsignaling:Initially,onlyahypothesis.Whetheritisadaptivefortheemitteraswellasthereceivermustbedetermined,Somehormones,suchasethyleneandstrigolactones,serveascommunicationvectorsbothinternallyandexternally,Cuesinformdecisionsaboutwhenandhowtoallocatefiniteresources,Likeapokerplayer,plantshavelimitedresources.Gamblingonabadhand,orexpendingresourcesatthewrongplaceortimecanbeabigmistake,Signalsandcuesindicatethe“bestbet”,Cuesthatindicatefutureconditionsandcircumstancesareparticularlyimportant.Plantsgrowslowly,andcantrunaway,sotheyhavetolivewiththeconsequencesoftheirbehaviors,SeeforexampleShemesh,H,BFZaitchik,TAcuna,andANovoplansky(2012)ArchitecturalplasticityinaMediterraneanwinterannual.PlantSignal.Behav.7:492501andShemesh,H.andNovoplansky,A.(2012)Branchingtherisks:architecturalplasticityandbet-hedginginMediterraneanannuals.PlantBiol.,Inpress.PhotocreditTomDonald.,Plantbehavior,Whataplantdoesinthecourseofitslifetime,inresponsetoaneventorchangeintheenvironment,Example:Phototropiccurvaturetowardsalightsource,Images:Wisconsinfastplants,Tangopaso,Plantbehavioraffectsmorphologicalandbiochemicalphenotypes,Franklin,K.A.andWhitelam,G.C.(2005).Phytochromesandshade-avoidanceresponsesinplants.Ann.Bot.96:169-175.Ryan,C.A.(1990).Proteaseinhibitorsinplants:Genesforimprovingdefensesagainstinsectsandpathogens.Annu.Rev.Phytopathol.28:425-449.,Plantscantrunawaytheyhavedifferentformsofbehavior,Plantbehaviorisaffectedbymanyenvironmentalparameters,Plantbehaviorismediatedthroughphenotypicplasticity,Casestudy:Plasticityofleafmorphologyinaquaticplants,Lin,B.-L.andYang,W.-J.(1999).BluelightandabscisicacidindependentlyinduceheterophyllousswitchinMarsileaquadrifolia.PlantPhysiol.119:429-434.,Submergedleafphenotype,Manyspeciespronetoperiodicsubmergenceshowphenotypicplasticityoftheirleafforms.Submergedleavesareoftenthinnerandwithoutstomataorcuticle.,ThehormoneABAisonesignalthatinitiatestheswitchtotheaerialform,andinthisplant,Marsileaquadrifolia,bluelightisanother.Theplantwasirradiatedwithbluelightatthepositionindictedbythearrowhead,Summary:Behavioristhevariableresponsetotheenvironment,Themechanismsbywhichplantsperceivetheirenvironmentandintegrateinformationintoabehavioralresponsearenotwellunderstood,butunderintenseinvestigation,Plantinteractionsinvolveperceptionthroughcuesandsignals,andplasticbehavioralresponses,Photocredit:TomDonald,VictoriaNuzzo,NaturalAreaConsultants;EasyStockPhotos;Hassanali,A.,Herren,H.,Khan,Z.R.,Pickett,J.A.andWoodcock,C.M.(2008).Integratedpestmanagement:thepushpullapproachforcontrollinginsectpestsandweedsofcereals,anditspotentialforotheragriculturalsystemsincludinganimalhusbandry.Phil.Trans.R.Soc.B363:611-621copyright2008TheRoyalSociety;Karban,R.,Baldwin,I.T.,Baxter,K.J.,Laue,G.andFelton,G.W.(2000).Communicationbetweenplants:Inducedresistanceinwildtobaccoplantsfollowingclippingofneighboringsagebrush.Oecologia.125:66-71.MaxPlanckInstituteforChemicalEcology,Jena,Germany/RaykoHalitschke.,Plantsareaffectedbyeachotherpositivelyandnegatively,Negativeeffect:Competitionforlight,Negativeallelopathiceffect:Here,theinvasivespeciesAlliariapetiolata(garlicmustard)suppressesallothers,Positiveeffect:Nutrientsharingandsuppressionofparasitism,Positiveeffect:Stresscues,“Wecandimlyseewhythecompetitionshouldbemostseverebetweenalliedforms,whichfillnearlythesameplaceintheeconomyofnature.”CharlesDarwin,1859,OntheOriginofSpecies,Ch3StruggleforExistence,Withsimilarneeds,competitionbetweenplantscanbeintense,Plantscompetewiththemselvesandothers,Treesgrowinginforestscangrowmorethan100metershigh,shadingplantsbelowthem,includingtheirownoffspring,Lightcanbealimitingresourceinmanyenvironments,PhotoscourtesyArielNovoplansky,Responsestoshading:confrontation,avoidance,tolerance,ReprintedfromVandenbussche,F.,Pierik,R.,Millenaar,F.F.,Voesenek,L.A.C.J.andVanDerStraeten,D.(2005).Reachingoutoftheshade.Curr.Opin.PlantBiol.8:462-468withpermissionfromElsevier,SeealsoNovoplansky,A.(2009)Pickingbattleswisely:plantbehaviourundercompetition.PlantCellEnviron.32:726-741.Photocredits:LennartSuselbeek;ArielNovoplansky,Light,Plantsperceivelightlevelsandcolor(or-lightquantityandquality),Detectionoflightlevelsandquality,Plantsperceivelightthroughphotoreceptorsthataresensitivetolightofvariouswavelengths,Phytochromedetectstheboundaryofphotosyntheticallyactiveradiance,Cryptochromedetectsblue(450nm)light,Actionspectrumforphotosynthesis,ReprintedbypermissionfromMacmillanPublishersLtdfromSmith,H.(2000).Phytochromesandlightsignalperceptionbyplants-anemergingsynthesis.Nature.407:585-591;AdaptedfromJaillais,Y.andChory,J.(2010).Unravelingtheparadoxesofplanthormonesignalingintegration.Nat.Struct.Mol.Biol.17:642-645.,Alowratioofredtofar-redlightisindicativeofvegetativeshading,Fullsun,R,FR,R,FR,Greenandredlightarenotabsorbedbythephotosyntheticpigmentstheyreflectandpassthroughleaves,Phytochromesconformationandabsorptionspectra“switch”,Phytochromechangesconformationwhenitabsorbslight:RedlightconvertsittothePfrform,whichmainlyabsorbsfar-redlight;Far-redlightconvertsittothePrform,whichmainlyabsorbsredlight,ReprintedbypermissionfromMacmillanPublishersLtdfromSmith,H.(2000).Phytochromesandlightsignalperceptionbyplants-anemergingsynthesis.Nature.407:585-591.,Transductionoflightinformationdownstreamofphotoreceptors,ReprintedfromGommers,C.M.M.,Visser,E.J.W.,Onge,K.R.S.,Voesenek,L.A.C.J.andPierik,R.(2013).Shadetolerance:whengrowingtallisnotanoption.TrendsPlantSci.18:65-71withpermissionfromElsevier.,Manyofthemoleculareventsthatcontributetoshadeavoidancehavebeenelucidated,andincludeeffectsontranscriptionfactoractivityandhormonelevelsandresponses,Shadeavoidanceisacollectionofresponsestovegetativeshading,ReprintedfromCasal,J.J.(2012)ShadeAvoidance.TheArabidopsisBook10:e0157.doi:10.1199/tab.0157,FromBallar,C.L.,Scopel,A.L.andSnchez,R.A.(1990).Far-redradiationreflectedfromadjacentleaves:Anearlysignalofcompetitioninplantcanopies.Science.247:329-332.reprintedwithpermissionfromAAAS,deWit,M.,Kegge,W.,Evers,J.B.,Vergeer-vanEijk,M.H.,Gankema,P.,Voesenek,L.A.C.J.andPierik,R.(2012).Plantneighbordetectionthroughtouchingleaftipsprecedesphytochromesignals.Proc.Natl.Acad.Sci.USA(2012)109:14705-14710.,Plantselongatelesswhenwearingacollarthatfiltersoutfar-redlightreflectedfromadjacentleaves,Touchisanothercuethatsignalsfuturecompetitionandstimulateselongation,Casestudy:Portulacaoleracea,lightresponsesinrecumbentplant,ReprintedwithpermissionfromNovoplansky,A.,Cohen,D.,andSachs,T.(1990)HowPortulacaseedlingsavoidtheirneighbours.Oecologia82:490-493.,LowR:FRlight,Growthdirection;probability,Futureshadecanbemoreimportantthanpresentshade,AdaptedfromNovoplansky,A.(1991).Developmentalresponsesofportulacaseedlingstoconflictingspectralsignals.Oecologia88:138-140.,FiltersweresetuponoppositesidesofPortulacaseedlings,Oneside(grey)transmittedverylittlephotosyntheticlight,equallylowinRandFR,Theothersidetransmittedandreflectedmorephotosyntheticlightbutmuchmorefar-redthanredlight,Dotheplantsgrowtowardsthesidewithmorelightnow(green)ormorelightlater(grey)?ThelowR/FRratioonthe“green”sideimpliesthepresenceofpotentialcompetitor,Littlered,Littlefar-red,Portulacagrowsawayfromfar-redlight,evenwhenitmeansthattheyaregrowingtowardslesslight,LowR:FRcuesindicatethepresenceofacompetitor,sogrowingawayfromthedirectionofsuchcuesmightimprovetheplantsfitness,Someplantshaveevolvedtotolerateshadelifeinthedimlane,Shade-tolerantspeciesareadaptedtolowlightenvironments,PhotoscourtesyTomDonaldandArielNovoplansky,Shadetolerancetakesmanyforms,ReprintedwithpermissionfromValladares,F.andPearcy,R.W.(1998).ThefunctionalecologyofshootarchitectureinsunandshadeplantsofHeteromelesarbutifoliaM.Roem.,aCalifornianchaparralshrub.Oecologia.114:1-10;Coley,P.D.(1988).Effectsofplantgrowthrateandleaflifetimeontheamountandtypeofanti-herbivoredefense.Oecologia.74:531-536.SeealsoValladares,F.andNiinemets,.(2008).Shadetolerance,akeyplantfeatureofcomplexnatureandconsequences.Annu.Rev.Ecol.Evol.Syst.39:237-257andGommers,C.M.M.,etal.(2013).Shadetolerance:whengrowingtallisnotanoption.TrendsPlantSci.18:65-71.PhotocourtesyofRonaldPierik,Long-livedshade-tolerantplantsinvestrelativelymoreresourcesintodefensesagainstherbivoresandpathogens,Shade,Sun,Reducedstem-elongationresponse,Energeticsofshadetolerance:max.assimilation,min.expenditure,Lightinformation:Angle,gradients,quality,andtimeofday,SeeforexampleSellaro,R.,Pacn,M.andCasal,J.J.(2012).DiurnaldependenceofgrowthresponsestoshadeinArabidopsis:Roleofhormone,clock,andlightsignaling.Mol.Plant.5:619-628.,LightinformationisMOREthanjustquantityandspectrum.Itislikelythatplantsrespondtoarichergamutoflightcues,Vertical,mid-dayshade(possiblylowR:FR)mightindicateaverytallneighbordontbothertryingtocatchup!,Horizontalandlate-daylowR:FRcuesmightindicatesimilarly-sizedneighborsgoforit!,Germinationplasticity:lightandothercuesforseedgermination,Photocredits:ForestHowardF.Schwartz,ColoradoStateUniversity,B,Oneofthemostimportantdecisionsaplantmakesiswhentogerminate,Manyseedsgerminateinresponsetowhiteorredlight,butfar-redlightisinhibitory,R,FR,Fire(heatandsmoke)canpromoteseedgermination,Imagesources:pfern,Hesperian,KurtStueber,2003,Firestimulatesseedreleaseorgerminationinsomeplants,Someconesandseedpodsarefire-serotinous,openinginresponsetofire,Banskiaspp,beforeandafterfire,Karrikinsaregermination-promotingcompoundsfoundinsmoke,ReprintedfromChiwocha,S.D.S.,Dixon,K.W.,Flematti,G.R.,Ghisalberti,E.L.,Merritt,D.J.,Nelson,D.C.,Riseborough,J.-A.M.,Smith,S.M.andStevens,J.C.(2009).Karrikins:Anewfamilyofplantgrowthregulatorsinsmoke.PlantScience.177:252-256withpermissionfromElsevier,andseealsoFlematti,G.R.,etal.,(2004).Acompoundfromsmokethatpromotesseedgermination.Science305:977.,Fire-inducedgerminationletsseedlingsbecomeestablishedwithlesscompetitionfromtallerplants.Karrikinsarecuesfromsmokethatpromotegermination.However,followingafire,therecanbeincreasedcompetitionbetweensimilarly-sizedseedlings.,PlantsproducemanyredundantorgansthatcompetewitheachotherSomaticcompetitioncanincreaseplantperformancebyputtingresourcesintomoresuccessfulorgans,SeeSachs,T.andA.Novoplansky(1995)Treeform:architecturemodelsdonotsuffice.IsraelJ.PlantSci.43:203-212;PhotoscourtesyArielNovoplansky.,Somaticcompetition:Whenplantscompetewiththemselves,Arethesebranchessheddingasadirectresultofthemgrowinginlowlight?Orisitaresultofcompetitionwithother,moresuccessfulbranchesonthesametree?,Branchautonomymayvaryaccordingtocircumstances,AdaptedfromKawamura,K.(2010).Aconceptualframeworkforthestudyofmodularresponsestolocalenvironmentalheterogeneitywithintheplantcrownandareviewofrelatedconcepts.EcologicalResearch.25:733-744.,Thereisexperimentalevidencetosupportallthreetypesofresponsesbetweenshoots:independent,competitiveandcooperative,Casestudy:Two-shootpeasandcorrelativeinhibition,SeeNovoplansky,A.,Cohen,D.,andSachs,T.(1989)Ecologicalimplicationsofcorrelativeinhibitionbetweenplantshoots.Physiol.Plant.77:136-140;Snow,R.(1931).Experimentsongrowthandinhibition.PartII.Newphenomonaoninhibition.Proc.Roy.Soc.B.108:305-316.Sachs,T.andA.Novoplansky(1997)Whatdoesaclonalorganizationsuggestconcerningclonalplants?indeKroon,H.andJ.vanGroenendael(eds.)TheEcologyandEvolutionofClonalGrowthinPlants,pp.55-78,SPBAcademicPublishing,Leiden,TheNetherlands.,TwoshootpeasRemovingtheshootfromapeaseedlingcausestwoshootstoregenerateagoodsystemtostudysomaticcompetition!,5-dayoldpea,Shootremoval,5days,“Two-shootpea”,Twoequalshootscanco-exist,butveryoftenonebecomesdominantandtheotherdiesWhydoesthesmallershootdie?,Casestudy:Two-shootpeasandcorrelativeinhibition,RedrawnwithpermissionfromNovoplansky,A.,Cohen,D.,andSachs,T.(1989)Ecologicalimplicationsofcorrelativeinhibitionbetweenplanshoots.Physiol.Plant.77:136-140;Snow,R.(1931).Experimentsongrowthandinhibition.PartII.Newphenomonaoninhibition.Proc.Roy.Soc.B.108:305-316.,Ashootinthedarkcansurvive10days(usingnutrientreserves),butincompetitionwithamoresuccessfulshootinthelight,thedarkenedshootdies.Theplantselectivelyallocatesreservestothestrongershoot.,Model:Resourcesareallocatedtothebestoption.InA,thebest(only)optionistheshootinthedark.InB,resourcesareallocatedtotheshootinthelight,attheexpenseoftheothershoot.,A.Darkonly,B.Onedarkandonelightshoot,Withinandbetweentrees,branchesinthebestconditionsprevail,Lesssuccessfulbranches,e.g.shadedbyneighborsorselfarediscriminatedagainstandshed,Photocredit:TomDonald,Summary:Perceptionofandresponsetovegetativeshading,LightisaresourceandalsoasourceofinformationthataffectsplantbehaviorAdaptiveresponsestocompetitionforlightcanbearchitectural(shootposition,sizeandnumber),morphological(stemelongati