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歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立 臺南大學 環境與生態學院 生態科學與技術學系 教授 Ayo NUTN Web:

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Presentation on theme: "歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立 臺南大學 環境與生態學院 生態科學與技術學系 教授 Ayo NUTN Web:"— Presentation transcript:

1 歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立 臺南大學 環境與生態學院 生態科學與技術學系 教授 Ayo NUTN Web: 大學部 生態學與保育生物學學程 ( 必選 ) 2010 年 秋冬

2 Ayo 教材 ( 動物行為學 2010) 2 Part 1. 動物行為的研究途徑 ( 個體行為 ) 歷史背景 (History of the Study of Animal Behavior ). 基因分析 (Genetic Analysis of Behavior ). 天擇 (Natural Selection and Behavior ). 學習與認知 (Learning and Cognition.) 生理分析 (Physiological Analysis) ( 一 ) 神經細胞 (Nerve Cells and Behavior ). ( 二 ) 內分泌系統 (The Endocrine System). 發育 (The Development of Behavior ).

3 Ayo 教材 ( 動物行為學 2010) 3 02 歷史背景 (History) The beginnings Classical ethology Comparative psychology Sociobiology and Behavioral Ecology More Recent Trends Field study Cellular and molecular basis Behavioral biology Applied animal behavior

4 Ayo 教材 ( 動物行為學 2010) 4 動物行為學術研究的起源 It is hard to pinpoint the precise beginnings of the study of animal behavior 於西方, Its roots can be traced back to the ancient Greek philosophers ( 古 希臘哲學家 ) summarized in 1855 by Herbert Spencer in Principles of Psychology ( 心理學原理 ) Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, sociologist, and prominent classical liberal political theorist of the Victorian era. Herbert Spencer

5 Ayo 教材 ( 動物行為學 2010) 5 Intellectual continuity ( 理智連貫性 ) There is a continuity in mental states between “lower” and “higher” animals ( 低等和高等動物 ) Based on a picture of evolution similar to Aristotle’s scala naturae, the great chain of being A linear, continuous evolution of species Progression from sponges, fish, amphibians, reptiles, birds, nonhuman animals then humans The animal mind and the human mind were simply points on a continuum

6 Ayo 教材 ( 動物行為學 2010) 6 Darwin ’ s evolutionary framework 1. Variation exists among individuals, and some of this variation is inherited 2. Most offspring do not survive to reproduce 3. Some individuals survive and produce more offspring, as a consequence of inherited characteristics Charles Darwin’s The Origin of Species (1859) provided a framework for developing the field of animal behavior:

7 Ayo 教材 ( 動物行為學 2010) 7 Consequences of differential survival 4. Natural selection: the differential survival and reproduction of individuals resulting from genetically-based variation in their behavior, morphology, physiology, etc. 5. Evolutionary change occurs as the heritable traits of individuals that survive and reproduce are spread through the population - Traits of less successful individuals are lost

8 Ayo 教材 ( 動物行為學 2010) 8 George Romanes George John Romanes FRS (19 May 1848 – 23 May 1894) was a Canadian-born English evolutionary biologist and physiologist who laid the foundation of what he called comparative psychology, postulating a similarity of cognitive processes and mechanisms between humans and animals. George Romanes

9 Ayo 教材 ( 動物行為學 2010) 9 Romanes ’ table of emotions ( 情感 ) He listed emotions in order of their historical or evolutionary appearance

10 Ayo 教材 ( 動物行為學 2010) 10 Other notable scientists Jacques Loeb (1918): all patterns of behavior were simply “forced movements” or tropisms ( 趨性 ) Physiochemical reactions toward or away from stimuli Herbert Spencer Jennings wrote Behavior of the Lower Organisms (1906) Disagreed with Loeb Emphasized the variability and modifiability of behavior

11 Ayo 教材 ( 動物行為學 2010) 11 Herbert Spencer Jennings and Jacques Loeb Herbert Spencer Jennings (born in Tonica, Illinois, April 8, 1868; died in Santa Monica, California, April 14, 1947) was a zoologist, geneticist, and eugenicist( 優生學者 ). His research helped demonstrate the link between physical and chemical stimulation and automatic responses in lower orders of animals. Jacques Loeb (born April 7, 1859, in Mayen, Rhineland-Palatinate; died February 11, 1924, in Hamilton, Bermuda) was a German-born American physiologist and biologist. Jacques Loeb

12 Ayo 教材 ( 動物行為學 2010) 12 20th century: Ethology vs. Comparative psychology( 比較心理學 ) Differences in opinion led to two major disciplines: Ethology: centered in Europe Focused on the evolution and function of behavior Comparative psychology: centered in the United States Focused on the mechanism and development of behavior They asked different questions, and studied different types of behavior and organisms Ethologists studied innate behavior in birds, fish, and insects Comparative psychologists emphasized learned behavior in mammals such as the Norway rat

13 Ayo 教材 ( 動物行為學 2010) 13 Ethology vs. comparative psychology Describing the normal function of behavior Ethologists observed the animal in its natural habitat or in environments designed to simulate that habitat Comparative psychologists believed that learning was best studied in the laboratory where variables could be controlled Interests, Ethologists were interested in species’ differences Comparative psychologists searched for general “laws” of behavior.

14 Ayo 教材 ( 動物行為學 2010) 14 Classical ethology Its approach: evolutionary, comparative, descriptive, field oriented Asks “Why is that animal doing that?” Founded by Konrad Lorenz, Niko Tinbergen, and Karl von Frisch European zoologists who shared the Nobel Prize in 1973 Concentrated on the evolution of behavior Focused on inherited behavior Emphasized phylogeny (the evolutionary history of a species)

15 Ayo 教材 ( 動物行為學 2010) 15 Three Pioneer Observers of Animal Behaviour Karl von Frisch, Konrad Lorenz and Nikolaas Tinbergen, shared the 1973 Nobel Prize for Physiology or Medicine.

16 Ayo 教材 ( 動物行為學 2010) 16 Karl von Frisch b. Nov. 20, 1886, Vienna, Austria—d. June 12, 1982, Munich, W.Ger., zoologist whose studies of communication among bees added significantly to the knowledge of the chemical and visual sensors of insects.

17 Ayo 教材 ( 動物行為學 2010) 17 Konrad Lorenz b. Nov. 7, 1903, Vienna, Austria— d. Feb. 27, 1989, Altenburg, Austrian zoologist, founder of modern ethology, the study of animal behaviour by means of comparative zoological methods. His ideas contributed to an understanding of how behavioral patterns may be traced to an evolutionary past, and he was also known for his work on the roots of aggression.

18 Ayo 教材 ( 動物行為學 2010) 18 Nikolaas Tinbergen b. April 15, 1907, The Hague, Neth.— d. Dec. 21, 1988, Oxford, Eng., Dutch- born British zoologist and ethologist (specialist in animal behaviour) Tinbergen emphasized the importance of both instinctive and learned behaviour to survival and used animal behaviour as a basis for speculations about the nature of human violence and aggression. He is especially well known for his long-term observations of sea gulls, which led to important generalizations on courtship and mating behaviour.

19 Ayo 教材 ( 動物行為學 2010) 19 Ethologists compare closely related species Lorenz: to know why an organism looks and acts as it does, one must resort to the comparative method Comparative anatomists do this in examining morphology For example, to understand why male flies of the species Hilara sartor spins an elaborate silken balloon( 精緻絲 球 ) to present to a female before mating It is compared to other fly species in the family Empididae ( 舞虻科 )

20 Ayo 教材 ( 動物行為學 2010) 20 舞虻:雙翅目 Diptera 舞虻,又名舞蠅,是舞虻科( Empididae )的昆蟲, 已發現超過 3000 種,大部份分佈在全北區。牠們 主要是掠食性的,有多種不同的形態,一般都是 細小至中等身型的,非金屬色及有剛毛的。大部 份舞虻幼蟲都是掠食性的,並棲息在不同的環境, 包括水中或陸上。 有些舞虻,如歐洲的 Hilara maura ,雄虻會將獵物 用絲包裹,用來向雌虻求愛。

21 Ayo 教材 ( 動物行為學 2010) 21 Gift-giving behavior of H. sartor Empidid flies species display almost every evolutionary step in the progression toward the balloon display Many species show a wide variety of behaviors when approaching a female To avoid being eaten by the predacious female Without a comparison of the behavior with that of other species It would be hard to explain why males offer silken balloons to females

22 Ayo 教材 ( 動物行為學 2010) 22 Progression of gift giving by male Empis flies ( 舞虻 ) A male E. trigram approaches the female while she is eating In E. poplitea, the male captures a prey (i.e. a fly), and gives it to the female, before attempting to copulate Male H. quadrivittata gift-wrap the meal in a silky cocoon before offering it to the female In H. thoracica, the large, elaborate cocoon contains food that is small and of little value In H. maura, only some males place food inside cocoons; others enclose something meaningless, such as a daisy petal ( 雛菊花瓣 ) A male H. sartor presents a female with an empty gauze case ( 空的薄紗盒 ) that turns off the predatory behavior of the female

23 Ayo 教材 ( 動物行為學 2010) 23 The fixed action pattern (FAP) In the early 1900s, work by Charles Otis Whitman and Oskar Heinroth Each independently concluded that the displays of different species are often exceptionally constant These patterns of movement were as reliable as morphological characters in defining a particular group A fixed action pattern (FAP): a motor response that is initiated by some environmental stimulus but that can continue to completion without the influence of external stimuli Stereotyped patterns ( 固定型態 ) of behavior

24 Ayo 教材 ( 動物行為學 2010) 24 Greylag geese ( 灰雁 ) 灰雁棲息於湖泊、河灘水域地帶。 本物種是古北區代表鳥種之一, 繁殖於西伯利亞南部、歐洲北部、 東部、冰島以及中國新疆西部北 部、青海柴達木盆地青海湖、東 北的內蒙古、黑龍江等省份; 越冬於歐洲南部、地中海沿岸、伊拉克、印度西北部、中 國南方的江蘇、湖南、廣東、福建等地也有本物種越冬。

25 Ayo 教材 ( 動物行為學 2010) 25 Greylag geese ( 灰雁 ) 的 FAP A female greylag goose will retrieve an egg that has rolled just outside her nest She reaches beyond it with her bill and rolls it toward her with the underside of the bill If the egg is removed once the rolling behavior has begun The goose continues the retrieval response until the now imaginary egg is safely returned to the nest

26 Ayo 教材 ( 動物行為學 2010) 26 Once initiated, FAPs continue to completion Other characteristics of FAPs include: The sequence of component acts of an FAP is unalterable An FAP is not learned It may be triggered under inappropriate circumstances It is performed by all appropriate members of a species George Barlow (1968): most patterns of behavior are not as stereotyped as suggested by the idea of the FAP Most FAPs cannot easily be separated into fixed and orientation components He suggested the term modal action pattern (MAP)

27 Ayo 教材 ( 動物行為學 2010) 27 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) (IRM) that then cause the performance of the instinctive act, sometimes composed of very stereotyped movements (fixed action patterns) (FAP).

28 Ayo 教材 ( 動物行為學 2010) 28 A male European robin in breeding condition will attack a tuft of red feathers ( 一叢紅色羽毛 ) placed in his territory.

29 Ayo 教材 ( 動物行為學 2010) 29 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.

30 Ayo 教材 ( 動物行為學 2010) 30 The scratching movements of dogs, as well as many other vertebrates, are considered fixed action patterns.

31 Ayo 教材 ( 動物行為學 2010) 31 Learning ( 學習 ) Learning is a change in behavior, based on experience. Chimpanzees are highly social creatures that live in a complex, variable, and changing world. Intelligence is important under such circumstances.

32 Ayo 教材 ( 動物行為學 2010) 32 Chain of reactions ( 反應鏈 ) Behavioral complexity can occur by building FAP sequences Results in an intricate pattern called a chain of reactions ( 反應鏈 ) Each component FAP brings the animal into the situation that evokes the next FAP

33 Ayo 教材 ( 動物行為學 2010) 33 案例: the courtship ritual of the three- spined stickleback ( 三刺魚 ) A complex sequence of behaviors that culminates in the synchronization of gamete release Each female behavior is triggered by the preceding male behavior Which was triggered by the preceding female behavior 主要分佈在歐洲各河流、地 中海及黑海;北非的阿爾及 爾一帶;北太平洋的韓國、 白令海至下加利福尼亞;北 大西洋的卻沙比克灣、哈得 遜灣及巴芬島。

34 Ayo 教材 ( 動物行為學 2010) 34 三刺魚的求偶 (courtship) This complex sequence is a chain of FAPs, each triggered by its own sign stimulus (releaser) A male stickleback in reproductive condition may attack a female entering his territory After mating with a female The male chases her away and defends his territory against other males He entices( 吸引 ) three to five other females with his courtship routine The male guards the developing eggs from predators and fans water over them for aeration

35 Ayo 教材 ( 動物行為學 2010) 35 三刺魚的求偶反應鏈 The female’s head-up posture Releases the male’s zigzag dance Which releases the females’ approach The male turns and swims to the nest Enticing the female to follow At the nest, he lies on his side Releasing the female to enter the nest Which releases the male to prod the female Causing her to release her eggs She leaves nest, the male enters and fertilizes the eggs

36 Ayo 教材 ( 動物行為學 2010) 36 反應鏈,過程未必固定 There are many deviations in the precise order of the events in the stickleback courtship ritual Some actions must be repeated several times if one partner is less motivated than the other Despite some flexibility, the component behaviors do not occur randomly In any display, a particular behavior is more likely to be followed by certain actions than by others

37 Ayo 教材 ( 動物行為學 2010) 37 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. As animals become accustomed to a stimulus, habituation may occur.

38 Ayo 教材 ( 動物行為學 2010) 38 比較心理學的研究途徑 Physiological, developmental, quantitative, laboratory- oriented Emphasizes laboratory studies of observable, quantifiable patterns of behavior Good science ( 好科學 ) cannot be done under uncontrolled conditions Criticized ethologists for neglecting to quantify their results and analyze their data Early research focused on learning and the physiological basis of behavior.

39 Ayo 教材 ( 動物行為學 2010) 39 C. Lloyd Morgan C. Lloyd Morgan (Conwy Lloyd Morgan) (6 February March 1936) was a British psychologist. Morgan's canon ( 摩根原則 ) In no case is an animal activity to be interpreted in terms of higher psychological processes, if it can be fairly interpreted in terms of processes which stand lower in the scale of psychological evolution and development. (Morgan 1903, p. 59) when two explanations for a behavior appear equally valid, the simpler is preferred.

40 Ayo 教材 ( 動物行為學 2010) 40 E. I. Thorndike was a pioneer in research on what was called trial- and–error learning, now usually called operant conditioning. Edward Lee Thorndike (August 31, 1874 Williamsburg, Massachusetts[1] – August 9, 1949) was an American psychologist who spent nearly his entire career at Teachers College, Columbia University. His work on animal behavior and the learning process led to the theory of connectionism and helped lay the scientific foundation for modern educational psychology.

41 Ayo 教材 ( 動物行為學 2010) 41 A problem box Thorndike invented many “problem boxes” to measure the learning ability of animals.

42 Ayo 教材 ( 動物行為學 2010) 42 The conditioned reflex ( 情境反應 ) Described by Ivan Pavlov, a Russian physiologist (1927) A dog begins to salivate at the sight of food The sight of food signals the presence of food Pavlov’s dog: Pavlov rang a bell immediately before feeding a dog In time, the dog salivated at the sound of the bell alone

43 Ayo 教材 ( 動物行為學 2010) 43 Upon presentation of a light, meat powder would be blown into the dog's mouth, causing it to salivate.

44 Ayo 教材 ( 動物行為學 2010) 44 Operant conditioning ( 操作情境化 ) In the 1930s, psychologist B. F. Skinner demonstrated operant conditioning by employing a device now called a Skinner box. In operant conditioning, the reinforcement follows the behavioral response. In other words, the animal must do something.

45 Ayo 教材 ( 動物行為學 2010) 45 a Skinner box, which is used to demonstrate operant conditioning.

46 Ayo 教材 ( 動物行為學 2010) 46 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。

47 Ayo 教材 ( 動物行為學 2010) 47 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。

48 Ayo 教材 ( 動物行為學 2010) 48 情境 ( 制約 ) 引發反應 Comparative psychologists used operant and classical conditioning techniques to study learning abilities of species Thorndike examined learning in fish, chickens, cats, dogs, and monkeys He noted similarities in the learning processes of these species Consistent with the idea of intellectual continuity Animals might differ in what they learned or in how rapidly they learned it But the process of learning is the same in all species

49 Ayo 教材 ( 動物行為學 2010) 49 行為主張 (Behaviorism) Another important event that steered comparative psychology toward objectivity and laboratory analysis Behaviorism: a school of psychology that restricts the study of behavior to events that can be seen A description of the stimulus and the response it elicits Behaviorists identify the stimuli that elicit responses and the rewards and punishments that maintain them Designed experiments that yielded quantifiable data, invented equipment, and developed statistical techniques

50 Ayo 教材 ( 動物行為學 2010) 50 行為主張 關注 學習能力 Assumed that an animal’s mental capacity could not be measured directly, but its ability to solve a problem could Focused attention on learning ability as a popular research subject A learned response could be described objectively Experiments could be conducted under the controlled conditions of the laboratory. Use a simple and convenient animal i.e. a Norway rat Apply the results to other species.

51 Ayo 教材 ( 動物行為學 2010) 51 生理心理學的根源 The neurological mechanisms of behavior Pierre Flourens studied the relationship between behavior and brain structure He removed parts of the brain to look for the effect on the animal’s behavior Marie Jean Pierre Flourens (15 April 1794 – 6 December 1867), father of Gustave Flourens, was a French physiologist, the founder of experimental brain science and a pioneer in anesthesia ( 麻醉 ).

52 Ayo 教材 ( 動物行為學 2010) 52 Karl Lashley and Frank Beach Karl Lashley examined the role of the brain in emotion and vision and discovered that: Complex problem solving involved the entire cerebral cortex Learned responses do not depend on a fixed pattern of muscle movements Frank Beach studied the effects of brain lesions on rat maternal behavior Also studied the roles of nerves, hormones, and experience on the sexual behavior of fishes, amphibians, reptiles, birds, and mammals.

53 Ayo 教材 ( 動物行為學 2010) 53 Karl Lashley and Frank Beach Karl Spencer Lashley (1890–1958), was an American psychologist and behaviorist well-remembered for his influential contributions to the study of learning and memory. Frank Ambrose Beach, April 13, 1911 — June 15, 1988, believing that learned behavior was too complex for detailed analysis, he shifted the focus of the field toward the study of instinctive, or as he preferred, species-specific behavioral patterns, such as mating and parental behavior.

54 Ayo 教材 ( 動物行為學 2010) 54 比較心理學,擴展領域 Robert Yerkes studied primate behavior C. R. Carpenter studied howler monkeys, spider monkeys and gibbons, each in their natural settings T. C. Schneirla used both field observation and laboratory experimentation to investigate the social behavior in army ants He applied the rigorous methodology of laboratory researchers to his field studies These pioneering studies helped weave the sciences of ethology and comparative psychology together

55 Ayo 教材 ( 動物行為學 2010) 55 Robert Mearns Yerkes (May 26, 1876 – February 3, 1956) was an American psychologist, ethologist, and primatologist best known for his work in intelligence testing and in the field of comparative psychology. Clarence Ray Carpenter (usually credited as C. R. Carpenter) (1906? – March 1, 1975) was an American primatologist who was one of the first scientific investigators to film and videotape the behavior of primates in their natural environments.

56 Ayo 教材 ( 動物行為學 2010) 56 社會生物學 (Sociobiology) In the 1960s and early 1970s, field researchers such as John Crook and John Eisenberg suggested that ecological context could be a better correlate of social behavior than phylogeny Because ethologists often focus on phylogenetic analyses of behavior A new discipline – sociobiology, focused on the application of evolutionary theory to social behavior

57 Ayo 教材 ( 動物行為學 2010) 57 Behavioral Ecology ( 行為生態學 ) The key element of this approach: behavior should, on average, maximize fitness of individuals Behavioral ecologists try to identify the payoffs and costs that play a role in the evolution of behavior 按本益分析,研擬最佳化行為模式。再以適當的物 種,驗證這個模式。

58 Ayo 教材 ( 動物行為學 2010) 58 History of sociobiology During the late 1960s and early 1970s, most scientists believed that natural selection acted primarily on individuals Inconsistencies with selection at the level of the individual remained How do sterile castes in species of ants, bees, and wasps evolve? How is the evolution of nonreproducing individuals consistent with Darwinian selection? Why did altruistic behavior (which benefits others but is costly to the performer) evolve?

59 Ayo 教材 ( 動物行為學 2010) 59 Inclusive fitness Hamilton’s term(1964) inclusive fitness describes an individual’s collective genetic success as a combination of: Direct fitness (own reproduction) and Indirect fitness (effects on reproduction by nondescendant kin) An individual’s inclusive fitness includes all the offspring, (personal or of relatives), that are alive because of the actions of that individual

60 Ayo 教材 ( 動物行為學 2010) 60 E.O. Wilson E. O. Wilson’s 1975 text, Sociobiology, integrates ideas from ethology, ecology, and population biology Sociobiology = the systematic study of the biological basis of all social behavior To understand the evolution of social behavior, one must know Demography (e.g., information on population growth and age structure) and The genetic structure of populations

61 Ayo 教材 ( 動物行為學 2010) 61 Wilson is a two-time winner of the Pulitzer Prize for General Non-Fiction. He is known for his career as a scientist, his advocacy for environmentalism, and his secular-humanist and deist ideas pertaining to religious and ethical matters. Edward Osborne Wilson (born June 10, 1929) is an American biologist, researcher (sociobiology, biodiversity), theorist (consilience, biophilia), naturalist (conservationist) and author. His biological specialty is myrmecology, the study of ants.

62 Ayo 教材 ( 動物行為學 2010) 62 The field of animal behavior is revitalized During the 1970s and the early 1980s, research on sociobiological topics in animal behavior flourished Sociobiology provided a framework to test hypotheses about the adaptiveness or survival value of behavior For a time almost all research in animal behavior was done under the banner of sociobiology. Of all questions about animal behavior, one—its function, or survival value —dominated the field

63 Ayo 教材 ( 動物行為學 2010) 63 Ethology comes full circle By the end of the 1980s, it was apparent that understanding animal behavior would be fuller if both its immediate and evolutionary causes are considered The study of animal behavior has returned to research that considers all of Tinbergen’s four questions Ethology has regained its balance and addresses questions of mechanism as well as function

64 Ayo 教材 ( 動物行為學 2010) 64 Changes in focus: field studies Early field studies were purely descriptive and may have included an explanation of what was described Today’s field studies begin with a clearly stated hypothesis to be tested by data collection and analysis

65 Ayo 教材 ( 動物行為學 2010) 65 Field studies focus on costs and benefits Field studies focus on the costs and benefits of a particular behavior The common currency is reproductive success Natural selection shapes an efficient and optimal form of behavior Researchers might determine the fitness consequences of natural variation in the expression of behavior in a population

66 Ayo 教材 ( 動物行為學 2010) 66 Evolutionary stable strategies ( 演化穩定策略 ) An individual’s best choice of action often depends on what other members of the population are doing Consider whether an individual’s choice of action is an evolutionarily stable strategy (ESS) An ESS = a strategy that cannot be invaded by the spread of any rare alternative strategy when it is adopted by most members of the population The concept of ESS has been applied in studies of mating systems, communication, conflict, and cooperation

67 Ayo 教材 ( 動物行為學 2010) 67 The cellular and molecular basis of behavior New tools and techniques are increasing understanding of the mechanisms of behavior Sensory receptors and detection neurons respond to specific features of a stimulus We can identify neurons in circuits that underlie FAPs For example, it is now possible to map the nervous systems of several invertebrates

68 Ayo 教材 ( 動物行為學 2010) 68 Specific neurons can be linked to behavior In the grasshopper three different hind leg movements (FAPs) are involved in producing the courtship sound signals A specific type of brain nerve cell is responsible for each FAP During courtship, these nerve cell types are activated in a specific sequence Fixed action patterns are discussed in terms of neural networks, command neurons, or central pattern generators

69 Ayo 教材 ( 動物行為學 2010) 69 Research can focus on the molecular level Behavioral genomics: the study of the role of all an organism’s genetic material in behavior The biggest challenge will be to figure out how the environment and genome work together to direct the structure and behavior of an individual

70 Ayo 教材 ( 動物行為學 2010) 70 A reductionist approach to animal behavior Tries to understand an organism’s behavior by understanding its components But by the early 21 st century there were appeals to “return to the whole organism” If you want to know why an animal behaves a certain way, you must look at the whole organism, not just neurons, genes, and molecules

71 Ayo 教材 ( 動物行為學 2010) 71 Behavioral biology ( 行為生物學 ) Describes behavioral research that includes more than one of Tinbergen’s four questions An approach to animal behavior that integrates Tinbergen’s four questions ‑ cause, development, survival value, and evolution To provide a complete, correct understanding of behavior Tinabergen's four questions 的整合

72 Ayo 教材 ( 動物行為學 2010) 72 動物認知 (cognition) ( 認知行為學 ) This field began with Donald Griffin’s 1976 controversial book, The Question of Animal Awareness Animal cognition (cognitive ethology): an interdisciplinary area of research that brings Tinbergen’s four questions to bear on the study of animals’ mental experiences

73 Ayo 教材 ( 動物行為學 2010) 73 應用動物行為學 This subfield is divided into disciplines Many applied animal behaviorists focus on captive animals or companion animals i.e. training dogs and cats and solving behavioral problems Others study the positive effects of the human-animal bond For example, pets improve the mental health of the elderly Still other researchers work with laboratory, zoo and farm animals

74 Ayo 教材 ( 動物行為學 2010) 74 Applied animal behaviorists work to improve the welfare of captive animals Consider the “Five Freedoms” for captive animals proposed by The Farm Animal Welfare Council in the United Kingdom: (1) Freedom from thirst, hunger and malnutrition (2) Freedom from discomfort due to environment (3) Freedom from pain, injury and disease (4) Freedom to express the normal behavior of the species (5) Freedom from fear and distress

75 Ayo 教材 ( 動物行為學 2010) 75 Other applied animal behaviorists work with wild animals Professionals work in wildlife management (e.g., increasing the population of game species) and pest management Conservation behavior: principals of animal behavior are used in efforts to conserve biodiversity We need behavioral data about habitat preferences, migratory routes, territory size, social organization, food requirements, the risk of predation, mating habits etc.

76 Ayo 教材 ( 動物行為學 2010) 76 The study of animal behavior has been rejuvenated Many disciplines are now contributing to its study New techniques and interactions among disciplines allow us to ask and answer many questions about behavior that could not be addressed previously Our efforts will be most fruitful if we keep a clear focus on behavior as the driving interest of research.

77 Ayo 教材 ( 動物行為學 2010) 77

78 Ayo 教材 ( 動物行為學 2010) 78

79 Ayo 教材 ( 動物行為學 2010) 79 Summary The most important concept in the study of animal behavior was Darwin’s idea of evolution through natural selection Ethology and comparative psychology were two early approaches to the study of animal behavior Sociobiology focuses on the application of evolutionary theory to social behavior Researchers use new technologies to explore the mechanisms of behavior on a molecular or cellular level Today, the study of animal behavior has returned to a more balanced approach that considers mechanism and function Information on animal behavior is being applied to assist the welfare of captive animals and in conservation biology

80 Ayo 教材 ( 動物行為學 2010) 80 問題與討論 Ayo 台南 NUTN 站

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