Transcript ppt檔案 - 國立臺南大學

大學部 生態學與保育生物學學程 (必選)
2010 年 秋冬
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/
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 ).
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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
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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.
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Herbert Spencer
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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
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Darwin’s evolutionary framework
Charles Darwin’s The Origin of Species (1859) provided a
framework for developing the field of animal behavior:
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
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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
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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
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Romanes’ table of
emotions (情感)
 He listed
emotions in
order of their
historical or
evolutionary
appearance
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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
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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
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Jacques Loeb
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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
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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.
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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)
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 Three Pioneer Observers of Animal Behaviour
 Karl von Frisch, Konrad Lorenz and Nikolaas
Tinbergen, shared the 1973 Nobel Prize for
Physiology or Medicine.
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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.
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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.
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Nikolaas Tinbergen
 b. April 15, 1907, The Hague, Neth.—d.
Dec. 21, 1988, Oxford, Eng., Dutchborn 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
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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)
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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。
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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
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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
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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
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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
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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。
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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
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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)
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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).
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 A male European robin in breeding condition will attack
a tuft of red feathers (一叢紅色羽毛) placed in his
territory.
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 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.
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 The scratching
movements of dogs,
as well as many
other vertebrates, are
considered fixed
action patterns.
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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.
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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
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案例： the courtship ritual of the threespined 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。
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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
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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
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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
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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.
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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.
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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.
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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.
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A problem box
 Thorndike invented
many “problem
boxes” to measure
the learning ability
of animals.
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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
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 Upon presentation of a light, meat powder would be
blown into the dog's mouth, causing it to salivate.
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Operant conditioning (操作情境化)
 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.
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 a Skinner box, which is
used to demonstrate
operant conditioning.
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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。
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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。
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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
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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
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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.
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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).
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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.
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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.
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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
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 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.
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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
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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。
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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?
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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
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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
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 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.
 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.
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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應用動物行為學
 This subfield is divided into disciplines
 Many applied animal behaviorists focus on captive
animals or companion animals


Others study the positive effects of the human-animal
bond


i.e. training dogs and cats and solving behavioral problems
For example, pets improve the mental health of the elderly
Still other researchers work with laboratory, zoo and
farm animals
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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
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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.
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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.
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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
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問題與討論
[email protected]
 Ayo 台南 NUTN 站
http://myweb.nutn.edu.tw/~hycheng/
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