Animal Behavior (動物行為) 教科書:Wallace, R. A. (1997) Biology: the world of life. Addison Wesley Longman, Inc. 鄭先祐 (Ayo) 國立臺南大學 環境生態研究所 Japalura@hotmail.com Ethology and comparative psychology The modern study of behavior has taken two distinct routes in this century -- one European, one American. Fig 24.1 Two of the bounders of modern ethology (a) Konrad Lorenz Animal Behavior 2 The European approach was called ethology. Its goal was to understand behavior by study its cause, development, evolution, and function through the observation of animals in the wild, or under somewhat natural conditions. Animal Behavior Fig 24.1 Two of the bounders of modern ethology (b) Niko Tinbergen 3 American approach The American approach was called comparative psychology. Its goal was to understand behavior by studying animals in the laboratory under carefully controlled conditions. Its focus was learning, and its primary animal was the Norway rat. Animal Behavior 4 The definition of Instincts (本能) The perception of something in the environment (a releaser) triggers a reaction in a center in the central nervous system(the innate releasing mechanism) that then cause the performance of the instinctive act, sometimes composed of very stereotyped movements (fixed action patterns). Animal Behavior 5 Fig. 24.2 A male European robin in breeding condition will attack a tuft of red feathers (一叢紅色羽毛) placed in his territory. Animal Behavior 6 Fig. 24.4 Simplified diagram of how a fixed action pattern can be triggered. The releaser is perceived by some sort of receptor, which triggers the IRM to activate certain muscles, thereby producing an instinctive movement that usually involves fixed action patterns. Animal Behavior 7 Fig. 24.3 The scratching movements of dogs, as well as many other vertebrates, are considered fixed action patterns. Animal Behavior 8 Learning (學習) Learning is a change in behavior, based on experience. Fig. 24.5 Chimpanzees are highly social creatures that live in a complex, variable, and changing world. Intelligence is important under such circumstances. Animal Behavior 9 Essay 24.1 The advantage of forgetting Savants, a special class of retardates(智力退化), have very low IQs, but some are able to accomplish incredible mathematical feats(技藝), such as multiplying 2 five-figure numbers in their heads. Others can immediately tell you the day of the week on which Christmas day fell in 1492 or any day in any year. (記憶) Animal Behavior 10 A mnemoist A mnemoist (a Russian profession) would sometimes memorize lines of 50 words. Once, in just a few moments, he memorized the nonsense formula Nd2(853/vx)(2762/n2v)(86x/273)(n2b)(rd) = sv (1625/322)(r2s5) Fifteen years later, upon request, he repeated the entire formula without a single mistake. Animal Behavior 11 Essay 24.1 Why haven't been selected for so that by now we can all, more or less, perform such feats? Animal Behavior 12 Habituation (習慣) Habituation is , in a sense, learning not to respond to a stimulus. The first time an animal encounters a stimulus, it may respond vigorously. But if the stimulus is presented over and over without consequence, the response to it gradually lessens and may finally disappear altogether. (Fig. 24.6) Animal Behavior 13 Fig. 24.6 As animals become accustomed to a stimulus, habituation may occur. Animal Behavior 14 Classical conditioning (制約,情境化) Classical conditioning was first described by the famed Russian biologist Ivan Pavlov. The response to a normal stimulus comes to be elicited by a substitute stimulus. Animal Behavior 15 Fig. 24.7 Upon presentation of a light, meat powder would be blown into the dog's mouth, causing it to salivate. Animal Behavior 16 Fig.24.7 (b) in the first graph that the dog salivated at maximal levels after only eight trails. When the experiment was reversed and food no longer followed the light, the dog stopped salivating Animal Behavior after only nine trails. 17 Operant conditioning In operant conditioning, the reinforcement follows the behavioral response. In other words, the animal must do something. In the 1930s, psychologist B. F. Skinner demonstrated operant conditioning by employing a device now called a Skinner box (Fig. 24.8) Animal Behavior 18 Fig. 24.8 (a) B. F. Skinner, one of the most important twentieth-century psychologists. Animal Behavior 19 Fig. 24.8 a Skinner box, which is used to demonstrate operant conditioning. Animal Behavior 20 Skinner box An animal placed inside a Skinner box must learn to press a small bar in order to receive a pellet of food from an automatic dispenser. 開始的時候是隨機動作,當觸動到按鈕,有 食物下來,動物開始學習到觸動按鈕有食物 下來的情境。動物就會常常去觸動。 倘若每次觸動都會有,且非常的穩定。動物 會有愈來愈放心的現像。觸動頻率會減少。 Animal Behavior 21 Operant conditioning 倘若食物的供應轉變成不穩定,有時有, 但有時沒有。 動物的心情似乎變成有點緊張。動物會 增加去觸動按鈕的頻率。 甚至會隨時一再的觸動。沒有時間作其 它事情,譬如出外散步。 Animal Behavior 22 Fig. 24.9 In classical conditioning, a desirable commodity, such as food, comes to be associated with an irrelevant signal. Animal Behavior 23 Fig. 24.9(b) In operant conditioning, the animal can act when given a signal, but only one action is rewarded. Animal Behavior 24 Fig. 24.10 Although young birds have the innate ability to fly, they canAnimal improve with practice. Behavior 25 Fig. 24.10(b) A younger bird crashes headfirst into the ground. Animal Behavior 26 Imprinting (印痕) Critical period, a window of time when the young are particularly sensitive to certain aspects of their environment. At this time, the goslings, and the young of many other species as well, learn the traits of whatever is around them. Imprinting:the animal learns to make a specific response to certain aspects of its environment. Animal Behavior 27 Fig. 24.11 Konrad Lorenz leading a group of goslings (小鵝) that had imprinted on him. Animal Behavior 28 Fig. 24.12 A crane (鶴) imprinted on humans. The female whooping crane has been handreared and therefore had imprinted on humans. She rejected the mate provided for her but could be enticed to lay eggs (artificially fertilized) by "dancing" with humans. Animal Behavior 29 Orientation and Navigation Orientation is simply facing in the right direction navigation involves finding one's way from point A to point B. 這是隨季節遷移的動物必備的能力。 於籠子內的候鳥於遷移季節到來時,會 顯現出不斷的向遷移方向衝擊的現像。 Animal Behavior 30 Kramer experiments Kramer surmised that they were orienting according to the position of the sun. To test this idea, he blocked their view of the sun and used mirrors to change its apparent position. The birds oriented with respect to the position of the new "sun". Animal Behavior 31 Fig. 24.15 Kramer's orientation cage. The birds can see only sky through the cage roof. The direction of the sun can be shifted with mirrors. Animal Behavior 32 Kramer put identical food boxes around the cage, with food in only one of the boxes. The boxes were stationary, and the one containing food was always at the same point of the compass. 無論如何轉換位置,birds went directly to the correct food box. 但若是在陰天,the birds were disoriented and had trouble locating their food box. Animal Behavior 33 Biological clock If the artificial sun remained stationary, the birds would shift their direction with respect to it at a rate of about 15 degrees per hour, the sun's rate of movement across the sky. This meant that some sort of biological clock was operating, and a very precise clock at that. Animal Behavior 34 Essay 24.2 Biological clocks 將倉鼠放在微亮的 暗室,讓其無法得 知日夜運行。 但倉鼠仍就會在夜 晚的時間,出來運 動。 觀察多日,每日情 況都一樣。 Circadian rhythms Animal Behavior 35 調整周期 Although biological rhythms continue without environmental cues, they are not completely independent of such cues. Light-dark cycle will set, or entrain, the rhythm so that its period length matches that of the environment. Animal Behavior 36 Fig. 24.14 A pigeon with electrical coils on its head. Pigeons can detect slight magnetic fields and that magnetism can influence their orientation. Animal Behavior 37 Social Behavior The famous student of animal behavior, Jane Goodall (Fig. 24.15), who has spent much of her life among the chimpanzees of East Africa's Gombe Stream Preserve. Animal Behavior 38 Fig. 24.16 Fighting can be a dangerous activity among some species. Animal Behavior 39 Fig. 24.17 (a) a goldenfronted woodpecker (b) a red- bellied woodpecker 兩種很相近,但相互並不 interbreed Animal Behavior 40 Fig. 24.18 a raccoon approached a hare in the darkness and the hare leaped over the raccoon . They may have known each other. There is evidence that animals of different species that share the same area may react to each other individually. Animal Behavior 41 Fig. 24.19 Male rattlesnakes fighting. Each could kill the other, yet they do not bite. The fight is more a test of will and strength as the snakes press against each other, belly to belly. Animal Behavior 42 Fig. 24.20 Hyenas may kill any stranger they find on their territory. Animal Behavior 43 Fig. 24.21 Porpoises are intelligent creatures that often cooperate. Here a young porpoise swims beneath its mother. Animal Behavior 44 Fig. 24.22 Musk oxen live above the Arctic cycle and are preyed upon by wolves. If attacked, they immediately form a circle with the adults facing outward and calves inside. Animal Behavior 45 Fig. 24.23 Wolves often cooperate to bring down prey, but then the dominant individuals feed first. Animal Behavior 46 Fig. 24.24 Leaf cutter ants of the tropics carefully excise section of leaves, which they then take to the colony where the plant material is used to grow Behavior 47 fungi on which the ants Animal feed. Altruism An activity that benefits another organism at the individual's own expense. It seems to be common among animals. How did it evolve? Animal Behavior Fig. 24.25 救難犬 48 Fig. 24.26 Kin selection is the process by which an individual increases its kinds of genes in the population by helping relatives. Animal Behavior 49 ← Since they show altruistic behavior to each other's offspring, more offspring from each population survive. ↑Since they do not show altruistic behavior to each other's offspring, fewer offspring from each population survive. Animal Behavior 50 Fig. 24.28 A male baboon will sometimes threaten a predator, thereby placing himself at risk for the sake of the group. Animal Behavior 51 Essay 24.3 How to recognize kin (1) an animal comes to recognize those individuals it grew up with and to treat them as if they were related. (2) through maternal "labeling". (3) through "genetic markers". Animal Behavior 52 Reciprocal altruism Reciprocal altruism is selfless behavior that is extended when it is likely that the favor will be returned. Reciprocal altruism would be expected in those groups that are highly social and relatively intelligent and their behavior can be remembered. Animal Behavior 53 Fig. 24.29 In a formalized reciprocal altruism, society at large pays a small cost in order to offer help. Here, relief is distributed to hurricane victims at public expense. Animal Behavior 54 問題與討論 Animal Behavior 55