歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立臺南大學...

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: http://myweb.nutn.edu.tw/~hycheng/

大學部生態學與保育生物學學程 ( 必選 ) 2010 年秋冬

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 ).


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


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


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


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:


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


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


Romanes’ table of emotions ( 情感 )

He listed emotions in order of their historical or evolutionary appearance


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


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


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


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.


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)


Three Pioneer Observers of Animal Behaviour Karl von Frisch, Konrad Lorenz and Nikolaas

Tinbergen, shared the 1973 Nobel Prize for Physiology or Medicine.


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.


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.


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.


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 )


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。


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


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


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


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。


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


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)


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).


A male European robin in breeding condition will attack a tuft of red feathers ( 一叢紅色羽毛 ) placed in his territory.


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.


The scratching movements of dogs, as well as many other vertebrates, are considered fixed action patterns.


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.


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


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。


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


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


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


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.


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.


C. Lloyd Morgan

C. Lloyd Morgan (Conwy Lloyd Morgan) (6 February 1852 - 6 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.


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.


A problem box

Thorndike invented many “problem boxes” to measure the learning ability of animals.


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


Upon presentation of a light, meat powder would be blown into the dog's mouth, causing it to salivate.


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.


a Skinner box, which is used to demonstrate operant conditioning.


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。


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。


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


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


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.


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 ).


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.


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.


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


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.


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


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。


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?


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


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


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.


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


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


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


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


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


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


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


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


A reductionist approach to animal behavior

Tries to understand an organism’s behavior by understanding its components

But by the early 21st 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


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


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


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


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


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.


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.


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


問題與討論

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Ayo 台南 NUTN 站 http://myweb.nutn.edu.tw/~hycheng/

歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立臺南大學...

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Transcript of 歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立臺南大學...

歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立 臺南大學...

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Transcript of 歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立 臺南大學...

歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立臺南大學...

Transcript of 歷史背景 (History) ─ 動物行為學 (Ethology) 鄭先祐 (Ayo) 國立臺南大學...