Animal

Behavior
Introduction
What is behavior?

- Animal behavior ¤×ÍÊÔ觷ÕèÊѵǏ·Óà¾×è͵ͺʹͧµ‹ÍÊÔè§àÌÒã¹ÊÔè§áǴŌÍÁ

- Individual behavior ¤×Í¡ÒáÃзӷÕè㪌¡ÅŒÒÁà¹×éÍÀÒÂ㵌¡ÒäǺ¤ØÁ¢Í§Ãкº»ÃÐÊÒ·
- ÁÕ¤ÇÒÁàª×èÍÁ⧵‹Í physiology
¤ÇÒÁÊӤѭ
1. ª‹ÇÂÊѵNJãˌ䴌ÍÒËÒÃ
2. ¡ÒÃÊ׺¾Ñ¹¸ØáººÍÒÈÑÂà¾È
(´Ö§´Ù´ à¡ÕéÂǾÒÃÒÊÕ àÅÕé§ÅÙ¡)
3. ¡ÒÃÃÑ¡ÉÒÀÒÇиÓç´ØÅ (homeostasis)
What is behavior?

ÊÔè§àÌÒÀÒ¹͡áÅÐ
ÀÒÂã¹ (Stimulus)

¤ÓÊÑè§ (Signal)
¾ÄµÔ¡ÃÃÁ
(ÚBehavior)
What is behavior?

- ÁÕ¤ÇÒÁÊӤѭµ‹Í¡Ãкǹ¡Òà natural selection

- ʋ§¼Åµ‹Í anatomy à¾ÃÒÐ recognition áÅÐ communication ¢Öé¹ÍÂÙ¡‹ ѺËҧ¡ÒÂ

behavior is adaptive COST vs BENEFIT

Benefits contribute Reproductive Success Fitness

to direct fitness
What is behavior?

- Discrete sensory inputs can stimulate both simple and complex behaviors

Êԧ⵨‹Ò½Ù§¦‹ÒÊÔ§âµà´ç¡·ÓäÁ?
Tinbergen’s Four Questions

1. Function à¾×èÍÍÐäà (˹ŒÒ·Õè¢Í§¾ÄµÔ¡ÃÃÁ)

2. Mechanism à¡Ô´ÂÑ§ä§ (¡Åä¡·Ò§ physiology ·Õè·Óãˌà¡Ô´¾ÄµÔ¡ÃÃÁ)
3. Development µÑé§áµ‹à¡Ô´¨¹âµ (¾Ñ²¹Ò¡ÒÃ)
4. Evolution ÇÔÇѲ¹Ò¡Òà (¨Ø´¡Óà¹Ô´ ÊÒà˵Ø)

Why is a common shrew selective in

what it accepts as prey?

Common shrew (Sorex araneus)

Ultimate & Proximate questions
1. Ultimate questions = Functional explanations (WHY?)
à¡ÕèÂǡѺ Function Evolution
2. Proximate questions = Mechanical explanations (HOW?)
à¡ÕèÂǡѺ Mechanism Development
Why do many songbirds breed in
spring and early summer?
Behavior and
Innate behavior
Behavior concept
lovebird
- ¡ÒÃáÊ´§¾ÄµÔ¡ÃÃÁ
= genetic + environment

¾ÄµÔ¡ÃÃÁÁÕ phenotyoic
variation ¨Ò¡ genetic ·Õµè ‹Ò§¡Ñ¹

- physiological rediness
= µŒÍ§ÁÕ¤ÇҾÌÍÁ·Ò§´ŒÒ¹Ã‹Ò§¡Ò¡‹Í¹¶Ö§¨Ð
áÊ´§¾ÄµÔ¡ÃÃÁ䴌
Behavior concept
male mice
bee cross-fostering experiments

twin
Behavior concept
1. Innate behavior
- developmentally fixed = all individuals exhibit virtually the
same behavior despite the inevitable environmental differences
within and outside their bodies during development and
throughout life.
- Without role models and opportunities for learning—as in
species with nonoverlapping generations— individuals might fail
to acquire the appropriate behavior, or acquire inappropriate
behavior, if genes did not exert strong influences on the
development of the behavior.
Behavior concept

2. Learning behavior
- the modification of behavior resulting from specific experiences.

Learning vs maturation
maturation = the situation in which
a behavior may improve because of
ongoing developmental changes in
neuromuscular systems
not true learning
Fixed Action Pattern (FAP)

- A sequence of behavioral acts that is essentially unchangeable

(usually occurs in a series of actions the same way every time)
- usually carried to completion once initiated.

Goose seeing egg-like object, then moving them to the egg pik.
Fixed Action Pattern (FAP)

The territorial response of male sticklebacks

Fixed Action Pattern (FAP)

- The FAP is triggered by an external sensory stimulus known as a sign

stimulus (stimuli are usually obvious).
- When these stimuli are communication signals exchanged between
members of a species, they are known as releasers.

IRM = Innate Releasing Mechanism

Fixed Action Pattern (FAP)

moths instantly fold their wings and drop to

the ground in response to the ultrasonic
signals sent out by predator bats
Orientation

Orientation ໚¹¡ÒÃÇÒ§µÓá˹‹§¢Í§µ¹ãˌÊÍ´¤ÅŒÍ§¡ÑºÊÔè§àÌÒÀÒ¹͡

external cues = ÊÔè§àÌҺҧÍ‹ҧ·Õªè ‹ÇÂãˌÊÒÁÒöà¤Å×è͹·Õèä»ã¹·Ò§·Õè¶Ù¡µŒÍ§

Movement
Paramecium

1. Kinesis = random movement

patterns in response to stimuli
in which there is no orientation
of the organism’s body to the
source of stimulation
Movement
Sow bugs, or woodlice
= Thigmokinesis
1. Kinesis
- äÁ‹ä´Œ·ÓÁØÁÊÍ´¤ÅŒÍ§¡ÑºÊÔè§àÌÒ
- ÍѵÃÒàÃçǢͧ¡ÒÃà¤Å×è͹·ÕèáÅÐ
à»ÅÕè¹·ÔÈ·Ò§ ÊÍ´¤ÅŒÍ§¡ÑºÊÔè§àÌÒ
Movement
Sea Turtle
2. Taxis = directed reactions
involving (in a single-stimulus
situation) an orientation of the
long axis of the body in line
with the stimulus source.
Movement
stream fish (trout)
2. Taxis
- ÁÕ·Ôȷҧṋ¹Í¹¡ÑºÊÔè§àÌÒ
(positive, negative)

áÁŧÊÒº = thigmotaxis
Migration

Migration = the regular

movement of animals over
relatively long distances
Migration
Wilderbeast Christmas red crab

Monarch butterfly
Navigation

Navigation = is the process by which an animal uses various

cues to determine its position in reference to a goal as it
migrates or homes. µŒÍ§Ãкبش·ÕèÍÂً
Navigation
1. Piloting: an animal moves from one familiar landmark to
another until it reaches its destination.
§‹Ò à¾ÃÒФ،¹ªÔ¹¡Ñº landmark áÅÐÃٌ·ÔÈ·Ò§

Gray whale
Navigation
2. Compass orientation: animals can detect directions and travel in
particular paths until reaching destination.
ÂÒ¡¡Ç‹Ò ᵋÁÕ·ÔÈ·Ò§·ÕÃè ٌÍÂÙá‹ ÅŒÇ ¼‹Ò¹ÊÔè§àÌÒ㹸ÃÃÁªÒµÔ ઋ¹ ʹÒÁáÁ‹àËÅç¡âÅ¡
Loggerhead sea turtles / Salmon
Behavioral Rhythms

- ¡ÒÃáÊ´§¾ÄµÔ¡ÃÃÁµÒÁª‹Ç§àÇÅÒµ‹Ò§æ à¡ÕèÂǡѺ light and dark cycles

- ÊѵǏºÒ§µÑÇÁÕ circadian rhythms ËÃ×Í circannual rhythms ã¹µÑÇÍÂÙዠŌÇ
Behavioral Rhythms

Lunar Cycle = ¢ŒÒ§¢Öé¹¢ŒÒ§áÃÁ ¡‹Íãˌà¡Ô´¹éÓ¢Öé¹¹éÓŧ

ᵋäÁ‹ä´Œà¡ÕèÂǡѺ light and dark cycles

fiddler crab
Animal Signals and Communication

- Signal = ¾ÄµÔ¡ÃÃÁ·Õè·ÓãˌÊѵǏ͹×è à»ÅÕ蹾ĵԡÃÃÁ

- Communication µŒÍ§»ÃСͺ´ŒÇ ¡ÒÃÃѺ ʋ§ áÅеͺʹͧµ‹ÍÊÑ­­Ò³
ÊÒÁÒöà¡Ô´·Ñé§ã¹áÅй͡ʻ‚ªÊÕ 

• Visual signals
• Tactile signals
• Chemical signals
• Auditory signals
• Electrical signals ઋ¹ »ÅÒäËÅä¿¿‡Ò
Animal Signals and Communication

Stimulus-response chain = ÊÔè§àÌÒ˹Ö觨Ð仡Ãе،¹¾ÄµÔ¡ÃÃÁµ‹Íä»àÃ×èÍÂæ

»ÅÒËÅѧ˹ÒÁ áÁŧËÇÕè
Animal Signals and Communication
¡ÒÃÊ×èÍÊÒÃà¡ÕèÂǡѺ lifestyle + environment
• Auditory signals

¨Ô駨͡ ªÐÁ´ ËÁÒã¹

Animal Signals and Communication

• Auditory signals
Bird Calls VS Songs fallow deer bucks
Animal Signals and Communication

• Tactile signals
Animal Signals and Communication
The dance of the honeybee
The information content of
animal communication varies
considerably.

• Visual communication
Animal Signals and Communication
bees and ants
• Chemical communication

- Pheromone = chemicals
released by an individual that
bring about mating and other Male silkworm moths
behaviors
㪌à¾×èÍÊ׺¾Ñ¹¸Ø àµ×͹ÀÑ ·Óãˌ໚¹ËÁѹ
Animal Signals and Communication
Minnows
• Chemical communication

- Pheromone = chemicals
released by an individual that
bring about mating and other honeybee colony
behaviors
㪌à¾×èÍÊ׺¾Ñ¹¸Ø àµ×͹ÀÑ ·Óãˌ໚¹ËÁѹ
Learning Behavior
Some behaviors develop more gradually

white-crowned sparrow (Zonotrichia leucophrys)

Habituation

- animal learns to ignore a repeated irrelevant stimulus

crows
Imprinting

- the recognition, response, and attachment of young to a

particular adult or object
duck Konrad Lorenz
Imprinting

- sensitive period = a limited phase in an individual animal’s

development when learning particular behaviors can take place
- critical period = the attachment response performance and
reinforcement are greatest

¡Ù»ÃÕ
Imprinting

1. filial imprinting 2. sexual imprinting

Spatial Learning

- using landmark or location indicator

- conitive map = a representation in an animal’s nervous system of
the spatial relationships between objects in its surroundings
Bee Clark’s nutcracker
Associative learning
1. Classical conditioning
= learning to associate an arbitrary stimulus with a particular outcome
associate its behavior with events
Pavlov’s dog
Associative learning

2. Operant conditioning (trial and error learning)

= learning to associate one of its own behaviors with a reward or
punishment and then tends to repeat or avoid that behavior
associate its behavior with the consequences of that behavior
Skinner box Fox Blue jay
Associative learning
àª×èÍÁâ§äÁ‹ä´Œ·Ñé§ËÁ´
Bird pelican chick

àÊÕ§¡ÑºÍѹµÃÒÂ

ÊաѺÍÒËÒÃ
Associative learning

1. Classical conditioning 2. Operant conditioning

Play

- a behavior has no apparent external goal, but may facilitate social

development or practice of certain behaviors and provide exercise.

º‹§ºÍ¡ÍÓ¹Ò¨ã¹Í¹Ò¤µä´Œ
Insight learning

- Cognition = the ability of an animal’s nervous system to perceive,

store, process, and use information gathered by sensory receptors
ravens abstract thinking by honeybees
Insight learning

- insight learning = ability to adapt past experiences that may involve

different stimuli to solve a new problem.
Insight learning

- Cognitive ethology = consciousness or awareness

Grass snakes
Social Learning
- learning through observing others
- forms the roots of culture = a system of information transfer through
social learning or teaching that influences the behavior of individuals in
a population
Foraging Behavior
Foraging behavior

- food-obtaining behavior
- To obtain food, any predator must expend time and energy, first in
searching for its prey and then in handling it (i.e., pursuing, subduing,
and consuming it)
optimal foraging theory

- natural selection will benefit animals that maximize their energy

intake-to-expenditure ratio
- ´ÙÇ‹Ò cost (reard) ¡Ñº benefit (risk) Íѹä˹ÁÕ¤‹ÒÊÙ§¡Ç‹Ò
¹¡¾ÔÃÒº ¡ÇÒ§
optimal foraging theory
Diet width

1. Generalists ¡Ô¹ä´ŒËÅÒÂÍ‹ҧ ᵋ¡çàÅ×Í¡ÍÒËÒ÷Õèãˌ¾Åѧ§Ò¹ÊÙ§

Diet width

2. Specialists ¡Ô¹ä´Œà©¾ÒкҧÍ‹ҧ
Reproductive behavior
Courtship behavior

patterns that lead to copulation and consists of a series of displays

and movements by the male or female

µÑǼٌ·ÕèËÒ§ÊÇÂÍÒ¨º‹§ºÍ¡¶Ö§¤ÇÒÁá¢ç§áç
¤ÇÒÁÍØ´ÁÊÁºÙó¢Í§Ã‹Ò§¡ÒÂ
Mating systems

1. Promiscuity
- no strong bond pairs between males and females
Mating systems
1. Promiscuity 2. Monogamy
- no strong bond pairs between - one male mating with
males and females one female
Mating systems

3. Polygamy
- an individual of one sex mating with several of the other sex.

3.1. Polygyny ª 1 + ­ àÂÍÐ 3.2. Polyandry ­ 1 + ª àÂÍÐ

¹¡ÍÕ¡á¨Ç
sexual dimorphism

males and females differ in appearance, a characteristic known as

sexual dimorphism, typically varies with the type of mating system
Parental care

any form of parental behavior that appears likely to increase the

fitness of a parents’ offspring
Parental care

¢Ö鹡Ѻ the needs of the young

Pheasant

Quail

monogamy
polygyny
Parental care

- certainty of paternity
African Jacana Mouthbrooders
high when egg laying and mating occur
together, as in external fertilization

internal äÁ‹Ãٌ äÁ‹àËç¹ÅÙ¡

Sexual Selection

- A form of selection in which individuals with certain inherited

characteristics are more likely than other individuals to obtain mates
- ÁÑ¡¨Ð·Óãˌà¡Ô´ sexual dimorphism
Sexual Selection

- direct competition between - choices made between them by

members of one sex for mates the opposite sex
(intrasexual selection) (intersexual selection)

striking colours

large antlers
Sexual Selection

- influenced by imprinting

àÅ×Í¡µÒÁ¾‹Í
Sexual Selection
- Mate-choice copying = individuals in a population copy
the mate choice of others
Agonistic behavior

- a system of behavior patterns with the common function of

adjustment to situations of conflict among conspecifics
- threats, submissions, chases, and physical combat

1. Threatening behavior
Agonistic behavior
2. Physical combat

3. Ritual
Game Theory

- the fitness of a particular behavioral phenotype is

influenced by other behavioral phenotypes in the population
fitness ¢Í§¾ÄµÔ¡ÃÁ䴌ÃѺÍÔ·¸Ô¾Å¨Ò¡¾ÄµÔ¡ÃÃÁÍ×蹢ͧÍÕ¡½†ÒÂ

¾ÄµÔ¡ÃÃÁà¡ÕèÂÇ¢ŒÍ§ËÅÒ»˜¨¨ÑÂ
äÁ‹ÁÕµÑÇã´·Õè¢Á‹ ¡Ñ¹ä´ŒÊÁºÙó
Game Theory

Side-blotched lizard

ÍÒ³Òࢵ
µÑÇàÁÕ·ÕèÁÒ¼ÊÁ
Genetic variation
Garter snake

à»ÅÕè¹¾×é¹·ÕèÍÒÈÑ ¡çà»ÅÕè¹ÍÒËÒÃ
Altruistic behavior

- One individual appears to behave in a way that benefits

others rather than itself
1. Sentinel behavior ÃÐÇѧÀÑÂ
Belding ground squirrel prairie dog
Altruistic behavior

2. Helpers at the nest

Naked mole rats
Florida scrub jay
Altruistic behavior

individual fitness
Inclusive fitness
µŒÍ§¡ÒÃàÂÍÐæ
= + ¢Í§µ¹àͧ

relatives’ fitness
¢Í§­ÒµÔ
Altruistic behavior
- genetic distance Áռŵ‹Í¡Òê‹ÇÂàËÅ×Í
- ¤Ó¹Ç³ä´Œ¨Ò¡ Coefficient of relatedness =
the proportion of gene that are identical in two individuals
because of common ancestors

r = n(0.5)L n = ¨Ó¹Ç¹ route

L = ¨Ó¹Ç¹ meiosis
Altruistic behavior

r = n(0.5)L n = ¨Ó¹Ç¹ route

L = ¨Ó¹Ç¹ meiosis
Altruistic behavior

r = ¤ÇÒÁã¡ÅŒªÔ´
- Hamilton's rule rB > C B = ¼Å»ÃÐ⪹·Õè­ÒµÔ¨Ð䴌
C = ÊÔ觷Õèŧ·Ø¹ä»
- Kin selection = Natural selection that favors altruism by
enhancing the reproductive success of relatives
µÑÇ·Õè fitness ÊÙ§ àÊÕÂÊÅÐ
à¾×èÍãˌ genetic material ¢Í§¡Ñ¹áÅСѹÂѧ¤§ÍÂً
Altruistic behavior

- Sentinel or selfish behavior

Social grooming
Vampire bat in primates
- Reciprocal altruism
= ª‹ÇµÑÇ·ÕèäÁ‹ãª‹­ÒµÔ
àÁè×ͼٌÃѺÁÕ¡Òà returns the favor
Social behavior
- Social behavior is any kind of interaction between two or
more animals, usually of the same species

- The discipline of sociobiology applies evolutionary theory

to the study and interpretation of social behavior.

Male dedge warblers

Cooperation

ËÇÁÁ×͵‹ÍµŒÒ¹ predator
Aggressive behavior

involve conflict among conspecifics and so wouldalso be

included under agonistic behavior

1. Territorial - Spraying behavior

2. Dominance - Pecking order
3. Sexual
4. Parental
5. Parent-offspring