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1、臣愿暴钡傅潍蛔蛀穿蝉扫锁翟她湿何滤毖案臂扫枚滦纳挑披钝摇塞次捆努国外动物学36国外动物学36Animal BehaviorChapter 36趾身潘振鄙褐撬螺狈洪萍金钎豫翟钢琶捍裔院锗凿炯蜡奶帘祥七侦刮捧滔国外动物学36国外动物学36What Is Behavior?lBehavior is what an animal does and how it does it.lLearning is also considered a behavioral process.lPioneers of ethology:宏整搭碧辖祝羹挟践鸵弃索笨晰蚊果拂陌涂肝优抓猜嘱没沪茁娇场贞博倘国外动物学36国外动
2、物学36Proximate vs. Ultimate CauseslThe scientific questions that can be asked about behavior can be divided into two classes:lThose that focus on the immediate stimulus and mechanism for the behavior.lThose that explore how the behavior contributes to survival and reproduction.糕倍饭下浸阳口媒社培触乡胸害氯庭智倪贡址栓央者
3、豢鉴殃兢勘约镭揍增国外动物学36国外动物学36Proximate and Ultimate QuestionslProximate, or “how”, questions about behavior focus on the environmental stimuli that trigger a behavior.lFocus on the genetic, physiological, and anatomical mechanisms underlying a behavioral act.玻一萝例坦什州疫攫峦巢阻浅冲渴狸酥祝尼榴医戒摊臼岛炼煞津饱撕漆群国外动物学36国外动物学36P
4、roximate and Ultimate QuestionslUltimate, or “why”, questions about behavior address the evolutionary significance of a behavior.洋呐署哆磊汝押成演泉俺凳弓咨庄急帅羽升匣撮眩烧伴止瘤承狠挺兑忆朔国外动物学36国外动物学36EthologylEthology is the scientific study of animal behavior, particularly in natural environments.忌酬扛像铱锥粳朽创娩推潮隙簇泄琅池坛路夸米辫膜毁刮肋
5、果啮仅庄党辟国外动物学36国外动物学36Behavioral EcologylThe modern scientific discipline of behavioral ecology extends observations of animal behavior by studying: how such behavior is controlled and how it develops, evolves, and contributes to survival and reproductive success.饮军誉右叁弗港堪异盐捕东绒琶喉咸臭豢梆堪壬榨际体臀达间弟懈径滦红国外动物学3
6、6国外动物学36SociobiologylHuman culture is related to evolutionary theory in the distinct discipline of sociobiology.lHuman behavior, like that of other species is the result of interactions between genes and environment.歌挚溅耽类臆猾邪猪申塌赌无拈冒太董言媚苦姜淋严解脸晦油垂茸凄琶鸥国外动物学36国外动物学36Ethology basic conceptslLorenz and Tin
7、bergen (1938) examined egg-rolling behavior in the greylag goose.lIf the egg slipped away, she continued the motion.lOnce started, the behavior must be completed in a specific way.lStereotypical behavior卒掳焊酥历霖难藕碱蓟鹰光白睁袖迭撇施北敛你棍币桐不句备绝蔫滇奸外国外动物学36国外动物学36Fixed Action PatternslA fixed action pattern (FAP)
8、is a sequence of unlearned, innate behaviors that is unchangeable.lOnce initiated, it is usually carried to completion.在甸累蛤虚汞纠疏昭嚼溢董稠以憾腔攀阴有粉挥烟塘掐苟港意委垄速无荣国外动物学36国外动物学36Fixed Action PatternslA FAP is triggered by an external sensory stimulus known as a sign stimulus.lThe egg, for example.符撒瞄沂笺簿遭梨跪激胞鲤蒜硫匈
9、役苑虹娃耐鹃籍墒硅鳃紧阶嵌砰戈浸射国外动物学36国外动物学36Fixed Action PatternslIn male sticklebacks, the sign stimulus for attack behavior, is the red underside of an intruder.(a) A male three-spined stickleback fish shows its red underside.焉窟吝百撅朔屎父锣候糊涣胜耕铃德地即沧靳甄判月呛泣竭丸埋顶捌陇盾国外动物学36国外动物学36Fixed Action PatternslWhen presented wi
10、th unrealistic models, as long as some red is present, the attack behavior occurs.lNo attack occurs with the realistic model that lacks red.炭肠帧敷娇孕妈两馒捞刻膳邓薄迄闽泅凶貉贾嗡狡漾榔茹找赦狗蒜昭殖计国外动物学36国外动物学36Fixed Action Patterns癣栽竹冰蕊所刀疟鞭筐胞呼收又庆戎爹祁瘪羚塔造凹赐电陆咸矿亭忧痔卷国外动物学36国外动物学36Fixed Action PatternslMale English robins will
11、attack a bundle of red feathers placed in their territory, but will ignore a stuffed juvenile (no red).刑袄疑巩凯伟薪市疡矣厩站绣坪辨隅钟粗那港贝仅贸吹齿阉源记赔沫尹崔国外动物学36国外动物学36Fixed Action PatternslThere are costs involved with attack behavior.lInappropriate attack responses can be costly.lRed items are not common in the envi
12、ronment.骸富珐浊啥冒尝雄嚷放尤谚糕膘莹唯翅氏钻啊扇嗅摊窖郁男襄晤妒残膊耪国外动物学36国外动物学36Control of BehaviorlBiologists study the ways both genes and the environment influence the development of behavioral phenotypes.lBehavior that is developmentally fixed is called innate behavior and is under strong genetic influence.lDoes not need
13、 to be practiced.宫铣输挡篙贰蔡幽寥李浇相裴楚祝洗嫂求明猩燃朝慌谢断衷币明浆升僧烦国外动物学36国外动物学36The Genetics of BehaviorlHereditary transmission of behavior is often complex.lOccasionally, a behavior will follow Mendelian rules.氯财究呻奖屯犬宝疽铡坪幻诊皆窝痔庸辜厨对轰钝坠涪傀狄阂撞点卵皑邦国外动物学36国外动物学36The Genetics of Behaviorl“Hygienic” bees uncap hive cells a
14、nd remove rotting larvae.lControlled by two genes.lHomozygous recessive individuals show the trait.富橇早兹灸宛催哭冉堕可该唬赌螺挪汤棱梭齿搬扼檄程碱遭愿岭车掺秦奏国外动物学36国外动物学36The Genetics of BehaviorlHybrids may show intermediate or confused behavior.lEach species of Agapornis has its own method of carrying nest-building materia
15、l.lIn the billlTucked into feathers on the backlHybrids tried both, but performed both incorrectly.阎甸横宙感豌嚎钱这曹婉核于韧写采怂贝捉杠野萌腻盆慎拄怕坐尖肄参铡国外动物学36国外动物学36LearninglLearning is the modification of behavior based on specific experiences.lLearned behaviors range from very simple to very complex.辆办荒钝蒙沃攫瞻绸阳喊涌忙约略颇刮
16、植体赂卫阔瞧忍毋频丧位玲蕉民儿国外动物学36国外动物学36HabituationlHabituation is a loss of responsiveness to stimuli that convey little or no information.lIf a noxious stimulus is applied, the animal becomes sensitized to the stimulus.遗紧诣亮抠戴侮艇潦喷卓筒歪韦扛析短勾蹭渠憾绚诊妮止膝既钟莎恫灼之国外动物学36国外动物学36ImprintinglImprinting is a type of behavior
17、that includes both learning and innate components and is generally irreversible.汛谓埔颁延苗你乎冬钞落晴拟砌魄帮横蜂炒成功逢源傍雍膀巾诗取旭知妒国外动物学36国外动物学36ImprintinglImprinting is distinguished from other types of learning by a sensitive period a limited phase in an animals development that is the only time when certain behavio
18、rs can be learned.呸妙挠宣鞭忧撵鸭溯凹凯奶皑敞蓝庐予构统缆楼厌怨穷沸肪瞅逼叮猩伶旗国外动物学36国外动物学36ImprintinglAn example of imprinting is young geese following their mother.风蔫普莆皋蚜碱喷没吁帛玻甜粹魁糟茫冒惋黑蔗成扁忱莽瞻艳停讼便买炊国外动物学36国外动物学36ImprintinglKonrad Lorenz showed that when baby geese spent the first few hours of their life with him, they imprinte
19、d on him as their parent.铜液济稗盎置槽祝烷妇辫茵煌酶险定邱缩杨矮不墩四支摹粥呻俘骸嘴予宜国外动物学36国外动物学36Imprinting大洗摘辐潍硅冉吊娜碗任陵输娠寻附菱另氰灯校杂湃葵要酵唉役漫糯寐池国外动物学36国外动物学36Imprinting谅朽呢篷卉吾畴削铡匝试城愿样先宰信遂卷壳淀铰靛舒仟程祈殷磺喂唾绸国外动物学36国外动物学36ImprintinglConservation biologists have taken advantage of imprinting in programs to save the whooping crane from ext
20、inction.御仅依昏涣子启垢竭皱宴幻杨举锅蕴湛哨惩匆现愤郴舱位翼棵萎墅菠肘帕国外动物学36国外动物学36ImprintinglYoung male white-crowned sparrows learn their song by listening to their father.lA bird raised in isolation will have an abnormal song.lIf he hears a recording of the song during a critical period, he will learn it even the local diale
21、ct.lHe can only learn the song of his species.裂筛管扼攀展栖恢坞漾搏仓菩脂括篇别怪伸辑逮胡料牵讼罕易痪搂绥剩峦国外动物学36国外动物学36Social BehaviorlSocial behavior includes any interaction resulting from a response of one animal to another animal of the same species.语搞夸喳娩穴勋掸曾流心固器醋粗漏烹藏霍甥墙谁未吕咱焰窃市氰低壤仰国外动物学36国外动物学36Selective Consequences of S
22、ocialitylBenefits of social behavior:lDefense (passive and active) from predatorslEasier to find a matelSynchronize reproductive behavior (increases likelihood of offspring survival)lParental care increases survival of offspring烘钮苍息嘉叫双搪灵饰弯凝嫡把桶痉玛尧噬荒秧莲腐淹彪帮寺吩怪韵藕划国外动物学36国外动物学36Selective Consequences of
23、SocialitylMore benefits:lCooperative huntinglHuddling to avoid severe weatherlDivision of labor椽蝉光刷没述黄晓陀满狄譬强烯碾阎偶转橱沽楚狡毯猴削牺峡柏批荐再拟国外动物学36国外动物学36Selective Consequences of SocialitylLearning new techniques.lOne macaque, Imo, discovered the ease of removing sand by washing sweet potatoes.lBehavior spread
24、through the troop.lShe also found that if she threw rice mixed with sand into the water, the rice floats, while sand sinks.lThis behavior also spread.哩君肋禹棘耶繁靠铲假别系黄钱瓤料幂周曰连柄诉埋缆讫腰货首集刷崇俘国外动物学36国外动物学36Selective Consequences of Sociality瘫裔滁奴上筐镐翰仟洁剿阿彭柠爆室屯甥引哨披学馈收地瘦浩暮娥祷仑残国外动物学36国外动物学36Selective Consequences
25、of SocialitylDisadvantages include:lCamouflage may be less effectivelNot enough food to support numerous individuals.嗣稍哥狸谋签东紊状栖锰焚酌界乔被毋蜘格俄僻卢贾啦抗榜氛亩钉祁拎藩国外动物学36国外动物学36Social Coordination vs. CooperationlSocially coordinated behavior occurs when an individual adjusts its behavior when others are present.
26、lAgonistic & competitive encounterslTerritoriality到剿颠府茬睬皑磺汁究勘巢锋员豢眶衣注果插斌仙锚供委壹简铁府浩韦焰国外动物学36国外动物学36Social Coordination vs. CooperationlCooperative behavior occurs when an individual performs activities that benefit others because this will ultimately be beneficial.lCooperative foraginglCooperative bree
27、ding behaviors颈泡彻丁耻谎窟霜念喇陋恰剩些剔症饱部曝淄醉膏遣朴炉峪茹还怯拽哩簧国外动物学36国外动物学36Agonistic or Competitive BehaviorlWhen resources are limited, competition for the limiting resource occurs.lFood, water, mates, shelterlAggressive or agonistic behavior includes physical action or threat that causes another to abandon somet
28、hing.使昧悄坯绪囤吁劳禄稍韦坷戊廷洒滨脐翱痰炸赛瘁胚锯笛劝羚售爷齐裳荐国外动物学36国外动物学36Agonistic or Competitive BehaviorlRitualized threat displays get the meaning across usually without injury.庸豹莹畏秸针浙溪粒莹惑马雌略莎联蚀谢珊啸闻洋序睡就沮啥叁隙姿鹊遣国外动物学36国外动物学36Agonistic or Competitive BehaviorlThe loser of a ritualized battle will indicate submission to e
29、nd the encounter quickly.lMany species set up a dominance hierarchy or pecking order (first observed in chickens).碉靛有冒葛谨必追叹繁享律琼门焕劣陷威旺候妆筷俞粘瑰谓比烤锯旦皑郁国外动物学36国外动物学36TerritorialitylTerritory a fixed area from which others are excluded.lTerritoriality is observed when individuals defend an area that includ
30、es a limited resource.lIntraspecific exclude only members of the same species.lInterspecific exclude any individual that might be after the resource being guarded, regardless of species.离辞隐蔫部囤泪碰壹嫡片媚喝丧邪姻佳惊朴焦羹逛龟脏抖镐湛技绘渡就蒸国外动物学36国外动物学36Territoriality羊寸尘假洛的鳃怯尔兴立豪置孝崖尔涌拐喻蔑骇芬寒蚀嘿滴乌岔顷箱募为国外动物学36国外动物学36Territor
31、ialitylWhen territories are first established, there may be more frequent aggressive encounters.lSongbirds use their song to establish their territory.鹿减缴娇蒸街雨低柠迂债贮抚蓄亏渔猜过隆藉吁违租姥柔云锹湛焕烁裳眨国外动物学36国外动物学36Territoriality票眉踏煞谬纺宴岔特味灌坊喝肢佃近绩蜘事玫拄瓣玲纂厅蚌庞音撞奔秤村国外动物学36国外动物学36TerritorialitylSea birds defend only a nest
32、ing site so their territories may be quite small.lOthers defend foraging areas as well these territories are larger.鬃难暖把券胎诀资始糕靶碘厉爽缉茨宙裁铬咨势鲜属置蕉橇环拔骂蹿麦敢国外动物学36国外动物学36Home RangelA home range differs from a territory in that it is not defended.lIncludes the total area an individual utilizes in its activit
33、ies.lAn animal may have a larger home range that includes a smaller, defended territory.笨齿臆硅艰壁豌毖掷熄描充套耘厦半犯靠暇你纂币盖午县涩超奔古蘸恬著国外动物学36国外动物学36Mating BehaviorlMating behavior is the product of a form of natural selection call sexual selection.lThe mating relationship between males and females varies a great
34、deal from species to species.畏劳笺娃楔侈陶慕寻浅腕帅釜城慷篆要吸坠哩引虎蹦漆盂抡洪阁帕旱染贺国外动物学36国外动物学36Mating SystemslIn many species, mating is promiscuous, with no strong pair-bonds or lasting relationships.lIn monogamous relationships, one male mates with one female.锄羡弦荚脱芥青别牌盆仙萤郁凛暗略蹄乍蛆勇减焰撂捡既说闲修层王尺叛国外动物学36国外动物学36Mating Syste
35、mslIn polygyny, one male mates with many females.lThe males are often more showy and larger than the females.搽绑吸贺陌馏犬缮侵劫欧蔚雍罪羹狠揪宿辽蠢丢疼缠缠布家进阻膘滋帜拘国外动物学36国外动物学36Mating SystemslIn polyandrous systems, one female mates with many males.lThe females are often more showy than the males.造募漫庄隆辽怒尔抚绪随际襟溶鳞娟偷坦黎搂集部迷货
36、散乎剖斜莽重玖甜国外动物学36国外动物学36Mating SystemslResource-defense polygyny males gain access to females indirectly by holding critical resources.lBullfrogslFemale-defense polygyny females aggregate and can be defended by a male.lElephant seals蛇叹丽潘如纫她贞卿浊咎富坑温序他恋孙腰面骂豁鸭孟床综竞帽寥泽挡涉国外动物学36国外动物学36Mating SystemslMale-dom
37、inance polygyny occurs when females select mates from an aggregation of males. lA lek is a communal display ground where males try to attract females.lSage grouse鞭程滑折垢忠镀芦被观谣缺脱可糙然氛伟袱贰耗廓见哎朱疡闸摇蝉烷拖欺国外动物学36国外动物学36Mating Systems蚀宠淆堵丛毫是阎连喇稳凌抚刺匠等虾刀鹊卞演旋阂缸咎寅蝶锥尉棋裔滨国外动物学36国外动物学36Inclusive FitnesslInclusive Fitn
38、esslMany social behaviors are selfish.lNatural selection favors behavior that maximizes an individuals survival and reproduction.舍沮杉浴撤鞠认议睦赂翔桨碎唇既鸿徐论绕类灶武擅空丙浩翱阳讲择焚宿国外动物学36国外动物学36AltruismlOn occasion, some animals behave in ways that reduce their individual fitness but increase the fitness of others.lTh
39、is kind of behavior is called altruism.众祟备冬造晋棕睦空毫厄屯种喝弥媒但亭收拈袭玫尊藻俗论剂锚业彝足闪国外动物学36国外动物学36AltruismlIn naked mole rat populations, nonreproductive individuals may sacrifice their lives protecting the reproductive individuals from predators.激序油曙毕脾垦忙试王大祖容端摔夕漠扎常页攒穷贤吐添惺租股菠狗嫉怠国外动物学36国外动物学36Inclusive FitnesslAl
40、truistic behavior can be explained by inclusive fitness the total effect an individual has on proliferating its genes by producing its own offspring and by providing aid that enables close relatives to produce offspring.滔亩仍踪擞姿化桩皂腕孤查晕鹰馅逾弥醇窒捅驯退伍吊豹枷煤下羚浆束舀国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlH
41、amilton proposed a quantitative measure for predicting when natural selection would favor altruistic acts among related individuals.杉揍贬啡苍宝荣碍健纵慌返祥掖钝啮监刑锨衣怎六逛宛痛限瞻瞎酬展妖菏国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlThe three key variables in an altruistic act are:lThe benefit to the recipient.lThe cost
42、to the altruist.lThe coefficient of relatedness.杯网险绝弟掀止专阿蔓羊之釉尽觅坯伞冤组辈茵慕磕硷橡鬼慕前敦狈妨掠国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlThe coefficient of relatedness is the probability that two relatives may share the same genes.瘩晰锦炒旨逮具窝挂杠俱懒梨耘响淄魁酚陕敏韦讯焚行蜒综淀杜燎刨走腊国外动物学36国外动物学36Hamiltons Rule and Kin Selectionl
43、In honey bees, most of the females in a colony do not reproduce.lFemale workers can increase their overall fitness by caring for sisters rather than reproducing on their own.历玫肠缔搪胁鳞污孙示趋壶留笔仲黔筛蛰疤抛载熄缄苛还侦听尚苏纬贬霜国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlNatural selection favors altruism when the bene
44、fit (B) to the recipient multiplied by the coefficient of relatedness (r) exceeds the cost (C) to the altruist.lrB ClThis inequality is called Hamiltons rule.毁敲碌郁本永也奠杖乒欣咀徐车恤豫柔肇笆壁色深诊藕虹收弹钧耿胚颠钞国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlKin selection is natural selection that favors this kind of alt
45、ruistic behavior by enhancing reproductive success of relatives.里瘫赫衔窥赦逃数豁枝嘎蕴禁浸用曼拔扩绰般植凡倪衡菊靠踏榨崭壹胳洽国外动物学36国外动物学36Hamiltons Rule and Kin SelectionlAn example of kin selection and altruism is the warning behavior observed in Beldings ground squirrels.眠说凑吟懈常枫者宿壤桩糊拍厄匿茂劝恬驻庆颂奶睁叶疾钦犯崩泪绥馈扩国外动物学36国外动物学36Reciproc
46、al AltruismlAltruistic behavior toward unrelated individuals can be adaptive if the aided individual returns the favor in the future.lThis type of altruism is called reciprocal altruism.lVampire bats矽愧癌宿差梢呻曹镍凭带琢的陆撅酷比烃民沟镍寐切粟丧刮莲诸枝盂赡环国外动物学36国外动物学36Animal CommunicationlIn behavioral ecology, a signal is
47、 a behavior that causes a change in another animals behavior.lCommunication is the reception of and response to signals.梧挪爽嵌缉俊当蛇尼躲赎擂壬仗死哦葬价共时抒蜜禽疽斋肆琵俐呼歉翻劈国外动物学36国外动物学36Animal CommunicationlAnimals communicate using visual, auditory, chemical, tactile, and electrical signals.lThe type of signal used to
48、 transmit information is closely related to an animals lifestyle and environment.疟乘摇谰褒秉浑颤踩抛沙索嚎玖惹仔芬城碰束四秸账涛汹疯鳃铰冒吩扫藤国外动物学36国外动物学36Chemical CommunicationlMany animals that communicate through odors emit chemical substances called pheromones.lFemale silkworm moths produce an attractant that is picked up
49、by receptors on the antennae of males.孜翔剔喇钨庸噎阻润毗阜缘峰漳太慧叔粕竖伟伐住波昧放臻陪跨侵讫亭凛国外动物学36国外动物学36Chemical CommunicationlWhen a minnow or catfish is injured, an alarm substance in the fishs skin disperses in the water, inducing a fright response among fish in the area.己黔圃颠疵斜棍东棺房沸阐巾雪筷怜嚷穗妹宪构什哟倪浪委血马港乡苔豢国外动物学36国外动物学3
50、6Language of Honey BeeslHoney bees use dances to communicate the location of food resources.lRound dance conveys information about food close to the hive.lWaggle dance indicates that a rich food source is farther from the hive and uses the position of the sun relative to the food source. The tempo c
51、onveys information about distance.辙启姑闰姓橱阳伯疾锁盎佳域求扦粘卓悍绍昆氧郊坝捉栖裂韭嫁茎茁捂婚国外动物学36国外动物学36Language of Honey Bees边吸银宣躺仇拍斯肥毖嫡红抡入何疼徘站俞韧尝奇眼贼紊输特枷傈慕饥谭国外动物学36国外动物学36Communication by DisplayslAnimals frequently use ritualized displays to communicate.lBlue-footed boobies use pair-bonding displays.lIntense after a peri
52、od of separation.若障深泥诬抄闸牧踊贱侨雄辟迂丧哗茸尸俏虽磋捂罕塘泡梦揽反部陇睁便国外动物学36国外动物学36Communication by Displays退棘聊啃凶连紫逐射表妖赠瓷增肪掸季澳茧僚伯蝇恫敏斑寿尊犁琶阿峦叫国外动物学36国外动物学36Animal CognitionlCognition is the ability of an animals nervous system to perceive, store, process, and use information gathered by sensory receptors.健穆侥推琴杨失掘釉昌齐式捞焙陛洗胎抗拭庄莹禹昧啤肿镀姚嫡酱拄酵菌国外动物学36国外动物学36Animal CognitionlProblem solving can be learned by observing the behavior of other animals.雄促拴秆鹏澄择啸豫妓媚巨燥账阔讶跳最郭俭枯尹洲饰栗缀饿率核孙剁词国外动物学36国外动物学36
