行为生态学崔旭红ppt课件

Chapter17BehavioralEcology\nBehavioriswhatananimaldoesandhowitdoesit.Everybehaviorresultsfromacombinationofgenetic(“nature”)andenvironmental(“nurture”)influencesInstinctivebehaviorLearningisalsoconsideredabehavioralprocess.Behaviorsareadaptive,theyaretheresultofalongevolutionaryprocess2\nReedwarblerCuckooAspeciesinteractionthatnicelyillustratesmanyaspectsofanimalbehavior...\nInstinctsallowreedwarblerstobuildnestsandtolayeggswithoutlearninghowtodosoEvenso,nestingbehaviorsonlyoccurwhentheappropriateenvironmentalcuesarepresent,andtheyimprovewithexperience(throughlearning)\nReedwarblerchicksinstinctivelybegforfood\nCuckoosremoveahosteggandlayoneoftheirowninanest\nCuckoosinstinctivelyparasitizereedwarblers’nests\nCuckoochicksinstinctivelyremovereedwarblereggsandyoung\n\nCuckoochicksinstinctivelyexploittheinnatefeedingbehaviorofthereedwarbleradult\nFemalecuckoochicksbecomeadultsthatinstinctivelyparasitizereedwarblernests\nTheaforementioned,specificbehaviorsareinstinctiveadaptationsthatarosethroughevolutionbynaturalselectionAssuch,eachrequiresanenvironmentalcontextandcanbemodifiedbyexperienceandlearning\n\nHybridlovebirdsprovideanotherexampleofthedualnatureofbehavior(combininggeneticandenvironmentalinputs)…\nHybridlovebirdsprovideanotherexampleofthedualnatureofbehavior(combininggeneticandenvironmentalinputs)…\nHybridlovebirdsprovideanotherexampleofthedualnatureofbehavior(combininggeneticandenvironmentalinputs)…\nHybridlovebirdsprovideanotherexampleofthedualnatureofbehavior(combininggeneticandenvironmentalinputs)…Hybridbehaviorhaselementsinheritedfrombothparents,andcanbemodifiedbyexperienceandlearning\nEverybehaviorhasbothproximateandultimatecausesProximatecausesor“how”,questionsaboutbehaviorfocusontheenvironmentalstimulithattriggerabehavior.Aswellas,focusonthegenetic,physiological,andanatomicalmechanismsunderlyingabehavioralact.EthologymainlyconcernsproximatecausesThreeethologistssharedtheNobelPrizein1973:KarlvonFrischKonradLorenzNikoTinbergen\nEverybehaviorhasbothproximateandultimatecausesUltimatecausesconcerntheevolutionarysignificanceofabehavior;i.e.,thebalancebetweenfitnesscostsandbenefitsthatselectivelyfavorsthebehaviorBehavioralecologyattemptstounderstandbothproximateandultimatecauses\nProximatecause:daylengthchangestriggerthereleaseofhormonesthatstimulateanest-buildingresponseUltimatecause:Thefitnessbenefitshaveoutweighedthecostsduringtheevolutionofnest-buildingbehavior\n17.3InstinctandlearningInstinctivebehavior(本能行为)Learnedbehavior(学习行为)\nInnatebehaviorInnate(instinctive)behaviorsdonothavetobelearned;theanimalperformsthemcorrectlywithnopriorexperienceAsimplekindofinnatebehaviorisafixedactionpattern(FAP)inresponsetoanexternalsensorystimulus(asignstimulus)\nLorenzandTinbergen(1938)examinedegg-rollingbehaviorinthegreylaggoose.Iftheeggslippedaway,shecontinuedthemotion.Oncestarted,thebehaviormustbecompletedinaspecificway.Stereotypicalbehavior\nFixedActionPatternsAfixedactionpattern(FAP)isasequenceofunlearned,innatebehaviorsthatisunchangeable.Onceinitiated,itisusuallycarriedtocompletion.\nFixedActionPatternsAFAPistriggeredbyanexternalsensorystimulusknownasasignstimulus.Theegg,forexample.\nFixedActionPatternsInmalesticklebacks,thesignstimulusforattackbehavior,istheredundersideofanintruder.(a)Amalethree-spinedsticklebackfishshowsitsredunderside.\nFixedActionPatternsWhenpresentedwithunrealisticmodels,aslongassomeredispresent,theattackbehavioroccurs.Noattackoccurswiththerealisticmodelthatlacksred.\nMalesticklebacksusesimplecuestorecognizeothermalesTinbergenshowedthatvariouslyshapedmodelscouldstimulatetheresponsejustaswellasrealmales,supportingthesign-stimulushypothesisMalesrespondedtored-belliedmodelsMalesdidnotrespondtorealisticmodelsthatlackedredbellies\nFixedActionPatterns\nHumanbabieshaveaninnatedivereflexWateronthefaceisthesignstimulusClosingthemouth,holdingbreath,andkickingconstitutethefixedactionpattern\nTinbergenalsoshowedthatgullchicksbeginresponsetoanappropriatelycoloreddotAnappropriatelycoloreddotisthesignstimulusBeggingisthefixedactionpattern\nLearningcanmodifyinnateresponsesparrot\nControlofBehaviorBiologistsstudythewaysbothgenesandtheenvironmentinfluencethedevelopmentofbehavioralphenotypes.Behaviorthatisdevelopmentallyfixediscalledinnatebehaviorandisunderstronggeneticinfluence.\nTheGeneticsofBehaviorHereditarytransmissionofbehaviorisoftencomplex.Occasionally,abehaviorwillfollowMendelianrules.\nTheGeneticsofBehavior“Hygienic”beesuncaphivecellsandremoverottinglarvae.Controlledbytwogenes.Homozygousrecessiveindividualsshowthetrait.\nLearnedbehaviorLearnedbehaviorsarethosethathavebeenmodifiedbyexperienceLearnedbehaviorsrangefromverysimpletoverycomplex.\nHabituation适应:ananimalignoresaninsignificantstimulustowhichitiscontinuallysubjected，examples:wildanimalslosetheirfearofhuman.Valuablelearning:helptonotwasteenergyandtimebyrespondingtounimportantstimuliWequicklyhabituateto“background”noisesLearnedbehavior\nHabituation(习惯化)\nAsanimalsbecomeaccustomedtoastimulus,habituationmayoccur.\nImprinting–isatypeofbehaviorthatincludesbothlearningandinnatecomponentsandisgenerallyirreversiblelearningthatislimitedtoasensitiveperiodinananimal’slifeLearnedbehavior\nImprintingisdistinguishedfromothertypesoflearningbyasensitiveperiod–alimitedphaseinananimal’sdevelopmentthatistheonlytimewhencertainbehaviorscanbelearned.\nImprintingAnexampleofimprintingisyounggeesefollowingtheirmother.\nImprintingKonradLorenzshowedthatwhenbabygeesespentthefirstfewhoursoftheirlifewithhim,theyimprintedonhimastheirparent.Lorenz’sclassicstudiessupportedhisimprintinghypothesis\nImprinting\nImprintingConservationbiologistshavetakenadvantageofimprintinginprogramstosavethewhoopingcranefromextinction.美洲鶴\nAcrane(鶴)imprintedonhumans.Thefemalewhoopingcranehasbeenhand-rearedandthereforehadimprintedonhumans.Sherejectedthemateprovidedforherbutcouldbeenticedtolayeggs(artificiallyfertilized)by"dancing"withhumans.\nImprintingYoungmalewhite-crownedsparrowslearntheirsongbylisteningtotheirfather.Abirdraisedinisolationwillhaveanabnormalsong.Ifhehearsarecordingofthesongduringacriticalperiod,hewilllearnit–eventhelocaldialect.Hecanonlylearnthesongofhisspecies.\nWehumanshaveasensitiveperiodforlearninglanguageLearnedbehavior\nAssociative(联想)–occurswhenananimalmakesaconnectionbetweenastimulusandanoutcomeAnanimalcanavoiddangeroranticipatebeneficialeventsbyassociationbehaviorLearnedbehaviorE.g.,IvanPavlov’sclassicexperiments\nClassicalconditioning(经典条件反射):occurswhenaninvoluntary,nature,reflexibleresponsetoanaturalstimulusistransferredfromthenaturalstimulustoanewstimulusConditionedresponse:theresponseproducedbythenewstimulus\nClassicalconditioningClassicalconditioningwasfirstdescribedbythefamedRussianbiologistIvanPavlov.Classicalconditioning–learningtoassociateanarbitrarystimuluswitharewardorpunishment\nUponpresentationofalight,meatpowderwouldbeblownintothedog'smouth,causingittosalivate.\nOperantconditioning(a.k.a.trial-and-errorlearning)–learningtoassociateoneofananimal’sownbehaviorswitharewardorpunishmentAssociativelearning–theabilitytoassociateonestimuluswithanother;includesseveralformsofconditioningLearnedbehavior\nOperantconditioning(操作性条件反射):theassociationofaparticularoutcomewithaspecificstimulus.TheanimalisrewardedorpunishedfollowingitsresponseDiffersfromclassicalconditioningRepeatactsthatbringgoodresultsArewardorpunishmentisreceivedaftertheanimalhasengagedinaparticularbehaviorTheresponseistypicallyamorecomplicatedbehaviorthanasimplereflex\nWequicklylearntoassociatetouchingflameswithapainful,burningsensationOperantconditioningLearnedbehavior\nOperantconditioningLearnedbehaviorToadslearntoavoidstinginginsectsthroughone-traillearning\n\nelectricshockPoisonousfood;\nOperantconditioning–theSkinnerboxLearnedbehaviorBothpositive&negativeoperantconditioningcanproducelearnedbehaviors\nB.F.Skinner,oneofthemostimportanttwentieth-centurypsychologists.\nAlthoughyoungbirdshavetheinnateabilitytofly,theycanimprovewithpractice.\nAyoungerbirdcrashesheadfirstintotheground.\nObservationallearning(imitation)(观察学习/模仿):aformofassociativelearningthatconsistsofacomplexsetofassociationsformedwhilewatchinganotheranimalbeingrewardedorpunishedafterperformingaparticularbehavior\nProblemsolvingcanbelearnedbyobservingthebehaviorofotheranimals.\nExploratorylearning:ananimalmovesabout,observesitssurroundings,andstoreinformationaboutitssurroundingsInformationstoredinmemorymaybevaluableimmediatelyorinthefutureExample:returntothefoodsourcerepeatedly\nInsightisdemonstratedwhenananimalevaluatesanewsituationandperformsthecorrect,non-instinctivebehaviorwithoutpriorexperience.aspecialkindoflearninginwhichpastexperiencesarereorganizedtosolvenewproblemsInsight(顿悟)Insightisproblem-solvingwithouttrail-and-errorlearning,soitreliesonpreviousexperienceincontextsotherthanthosethatcharacterizetheproblemathand\nChimpstacksupboxestoreachsuspendedfoodDogisunabletoworkouthowtoreachfoodInsight\nInsightSomeadultravenscansolvethisproblemthefirsttimetheyarepresentedwithit\nInstinctandlearninginthesameanimalDistributionoflearnedandinstinctivebehaviors\nAcomparisonofinstinctandlearningInstinctlearningBornwiththebehaviorNotbornwiththebehaviorGeneticallydeterminedNotgeneticallydetermined,butpartlyhereditaryNoexperienceisrequired;donecorrectlythefirsttimePerformanceimproveswithexperience;behaviorrequirespracticeCannotbechangedCanbechangedMemoryisnotimportantMemoryisimportantCanevolveasgenefrequencieschangeNottheresultofgeneticchangesTypicalofsimpleanimalswithshortlives,littlecontactwiththeirparentsTypicalofmorecomplexanimalswithlonglives,extensivecontactwithparentsOnlybepassedfromparentofoffspringbygeneticmeansAllowsacquiredbehaviorstobepassedfromparenttooffspringbyculturalmeans\n17.8SelectedtopicsinbehavioralecologyReproductivebehavior(繁殖行为)Territorialbehavior(领域行为)Dominancehierarchy(优势等级)Avoidingperiodsofscarcity(避免阶段性资源缺乏)Navigationandmigration(导航和迁移)Biologicalclocks(生物钟)Socialbehavior(社会行为)\nReproductivebehaviorFindingeachotherMostspeciesinvolvetheproductionofsignalsthatcanbeinterpretedbyothersofthesamespeciesAssuringfertilizationRaisingtheyoung:providingtheyoungwiththeresourcestheyneedtolivetoadulthoodThefemalemothreleasesapheromonetoattractmales\nGypsymoth舞毒蛾\n\nCourtship–behaviorpatternsthatprecedecopulation(orgametereleaseinspecieswithexternalfertilization)\nTerritorialbehaviorTerritory领地:thespaceusedforfood,mating,orotherpurposes,thatananimaldefendsagainstothersofthesamespeciesTerritorialbehavior领域行为:behaviorsofsecuringanddefendingtheterritoryAwaytodistributearesourcethatisinshortsupplyOftentiedtosuccessfulreproduction\n\nDominancehierarchy优势等级Arelativelystable,mutuallyunderstoodorderofprioritywithinthegroupismaintainedOftenestablishedinsocialanimalsDominanceindividual→second-ranking→…Thedominancehierarchyallowsindividualstogetpreferential优先的treatmentwhenresourcesarescarce\nAvoidingperiodsofscarcityResourceallocationbecomesmostcriticalduringperiodsofscarcity缺乏Becomeinactive(aestivate夏眠):frogs,toads,insectsHibernation冬眠:bats,marmotssquirrelsBuilt-inbehavior:squirrels,chickadees山雀\nNavigationandmigrationRequirement:MemoryofsurroundingsAwayofdeterminingdirection,e.g.magneticfield,landmarks,scenttrails,orreferencetotheSunandstarsInformationaboutdistance\nAmorecomplicatedmeansofrememberinginformationaboutlocationsinvolvescognitivemaps–internalrepresentationsofthespatialrelationshipsofobjectsinananimal’shomerange\nMovementthroughspaceMigration–relativelylong-distanceperiodicmovement(e.g.,annual)\nMovementthroughspaceMigration–relativelylong-distanceperiodicmovement(e.g.,annual)Animalsuseone,oracombination,ofthefollowingtofindtheirway:Piloting–movingfromonefamiliarlandmarktothenextOrientation–ananimaldetectscompassdirectionsandmovesinastraight-linepathtowardsitsdestinationNavigation–ananimalusesorientationcombinedwiththeabilitytodetermineitspresentlocationrelativetoitstargetlocation\nMovementthroughspaceMigration–relativelylong-distanceperiodicmovement(e.g.,annual)ExperimentswithEuropeanstarlingssuggestthatjuvenilesuseorientation,whereasadultsmakeinformedmovementsbycombiningorientationwithcognitivemaps(truenavigation)\nMovementthroughspaceTinbergenshowedthatwaspsusesimplelandmarkfeaturestofindtheirnestsInthisexample,femalewaspsmaysimplylearntolookforacertainpatternofobjects\nBiologicalclocksAnnualclocks:seasonalmigration,matingDailyclocks:jetlag(时差)Paloloworm矶沙蚕hunting,inpacificocean\nSocialbehaviorSociety:社会interactinggroupwithdivisionoflaborandgreaterspecialization,thewholeismorethanthesumofitspartsAfricanwilddogsociety\nHowdidvarioustypesofsocietiesdevelop?Whatselectiveadvantagedoesamemberofasocialgrouphave?Inwhatwaysaresocialgroupsbetteradapttotheirenvironmentthannonsocialorganisms?Howdosesocialorganizationaffectthewaypopulationgrowandchange?Difficultquestions\nSocialorganizationSolitarySomebenefits…DecreasedintraspecificcompetitionDecreasedriskofdiseaseDecreasedriskofdetectionbypredatorsSomecosts…DecreasedabilitytocooperateinfindingfoodDecreasedabilitytocooperateagainstpredatorsDecreasedlikelihoodoffindingamateSolitaryvs.Groupliving\nSolitaryvs.GrouplivingGrouplivingSomebenefits…IncreasedabilitytocooperateinfindingfoodIncreasedabilitytocooperateagainstpredatorsIncreasedlikelihoodoffindingamateSomecosts…IncreasedintraspecificcompetitionIncreasedriskofdiseaseIncreasedriskofdetectionbypredatorsSocialorganization\nSocialorganizationSolitaryGroupliving\nSolitaryGrouplivingSocialorganizationEusociality–aspecialtypeofgrouplivinginwhichonlyoneorafewmembersofthegroupeverreproduce\nSolitaryGrouplivingSocialorganizationEusociality–aspecialtypeofgrouplivinginwhichonlyoneorafewmembersofthegroupeverreproduceNon-reproductivemembersofeusocialsocietiessacrificetheirownindividualfitnesstoincreasethefitnessofothersinthegroup,i.e.,theyarealtruistic\nConceptmapsStimulusInstinctbehaviorLearningbehaviorAssociationThinkingImprintingInsightTerritorialbehaviorResponseinheritedlearningconnectionsMemory,conceptprotectionreorganizing\nSummaryInstinctbehaviorandlearningbehaviorKindsoflearning:habituation,association,exploratorylearning,imprinting,andinsightTopicsinbehavioralecology:reproductivebehavior,Territorialbehavior,dominancehierarchy,avoidingperiodsofscarcity,navigationandmigration,biologicalclocks,socialbehavior\nBehaviorservesmanyfunctions…\nForagingbehaviorForagingbehaviorcomprisesallofthemeansbywhichananimalsearchesfor,recognizes,andmanipulatesfooditems\nPlayTherearetwomainhypothesesfortheultimatereasonsforplay:\nPlayTherearetwomainhypothesesfortheultimatereasonsforplay:1.Thepracticehypothesispostulatesthatplayallowsanimalstoperfectbehaviorsneededlaterinlife2.Theexercisehypothesispostulatesthatplayhelpsmuscularandcardiovascularsystemsdevelopproperly\nMovementthroughspaceKinesis–changeinactivityorturningrateinresponsetoastimulusTaxis–orientedmovementtowardorawayfromastimulusSowbugsincreaseratesoftravelindryareas,whichhelpskeeptheminmoistareasSowbugsmoveawayfromlight,whichhelpskeepthemindarkplacesSeeFig.51.7\nMovementthroughspaceDispersal(ananimalbehaviorist’sdefinition)–one-timemovementawayfromthenatalhomerangeFig.51.35\nFig.51.14MovementthroughspaceTinbergenshowedthatwaspsusesimplelandmarkfeaturestofindtheirnestsInthisexample,femalewaspsmaysimplylearntolookforacertainpatternofobjects