Transcript Chapter 1
2010 年 秋冬 大學部 生態學與保育生物學學程 ( 必選 )
學習與認知
(Learning and Cognition)
─ 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 ). Ayo 教材 ( 動物行為學 2010) 2
05
學習與認知 (Learning and cognition)
Definition of Learning Types of Learning Habituation ( 習慣 ) Classical conditioning Operant conditioning Latent Learning Social Learning Species differences in Learning Comparative Studies Other Evidence of Cognitive abilities in animals Ayo 教材 ( 動物行為學 2010) 3
People have
presuppositions
about the mental lives of animals
We
discount
the abilities of some species Especially those very different than us We
anthropomorphize
our pets and other primates We assume that they think like we do How do we know what animals know?
Does a chimpanzee plans its actions to catch termites? Does an ant lion( 蟻獅 ) understand its own behavior Ayo 教材 ( 動物行為學 2010) 4
Definition of Learning
It is “a
process
through which
experience changes
individual’s
behavior
” an Learning is a
change
in our
capacity
for
behavior
as a result of
experience
Excluding the effects of
fatigue
,
sensory adaptation
, or
maturation of the nervous system
Behavioral changes resulting from learning are
not
always expressed
immediately
We can’t know whether an animal has learned something just by seeing a change in its behavior Ayo 教材 ( 動物行為學 2010) 5
A type of learning:
habituation
In
habituation
, the animal
learns not to respond
to a particular stimulus Because the stimulus has proven to be harmless
Habituation
: the waning of a response after repeated presentation of a stimulus. Once it occurs, its effects are long lasting
Habituation
is everywhere, from protozoans to humans It is often considered the
simplest form
of learning Ayo 教材 ( 動物行為學 2010) 6
Habituation in the clamworm ( 沙蠶 )
雙齒圍沙蠶（ Nereis succinea ）是一種廣泛分佈 的多毛綱動物。 It partially emerges from its tube to filter tiny bits of food from the water withdraws into its tube when it senses danger (i.e. a shadow) In the lab, subsequent presentations decreased escape responses The clamworms had habituated The effects lasted several hours Ayo 教材 ( 動物行為學 2010) 7
Ayo 教材 ( 動物行為學 2010) Habituation of the withdrawal response to a shadow by the clamworm.
8
Habituation is not due to adaptation
The clamworms’ decline in responsiveness was not because the sense organs became adapted to the stimulus Nor was the decline due to muscle fatigue, because habituated worms still withdrew in response to prodding The clamworms had
learned
to stop responding to the shadow Ayo 教材 ( 動物行為學 2010) 9
Habituation is specific to a particular stimulus
Young turkeys, chickens, and pheasants innately show antipredator behaviors They crouch and give alarm calls at the sight of objects moving overhead When a model was presented frequently, it elicited fewer and fewer alarm calls from chicks As adults, they respond only to the image of a predator Such as a hawk flying overhead Ayo 教材 ( 動物行為學 2010) 10
Habituation also occurs in species interactions
Bird
species and
bullfrogs
of a
stranger’s call
respond aggressively to a recording but not to a familiar call Territorial species reduce aggressive responses toward neighbors, But still respond aggressively toward
unfamiliar intruders Habituation
mediates this process Animals stop responding to a call when its heard repeatedly The decline was specific to the calls Ayo 教材 ( 動物行為學 2010) 11
Habituation as an experimental tool
Habituation is a useful tool to study cognitive processes in animals, including humans A subject is habituated to a stimulus And then a new stimulus is presented If the subject’s response changes The experimenter knows it can detect the difference between the two stimuli Ayo 教材 ( 動物行為學 2010) 12
A type of learning: classical conditioning
Associative learning
: an animal makes an association between a stimulus and a response One type of associative learning:
classical conditioning (Pavlov dog)
A dog salivates when powdered food is blown into its mouth, but not when it hears a bell These two stimuli are paired immediately before food powder was presented The dog salivates in response to the bell alone Ayo 教材 ( 動物行為學 2010) 13
Classical conditioning explained
Unconditioned stimulus
(US): an animal has an inborn response to a certain stimulus The animal did not have to learn the response
Unconditioned response
(UR): the response to the US
Conditioned stimulus
(CS): a new stimulus is paired with the US until eventually it, too, can elicit the response
Conditioned response
conditioned stimulus (CR): the response to the Ayo 教材 ( 動物行為學 2010) 14
Classical conditioning is consistent
The order of the presentation of the US and CS is important The two stimuli must occur close together Useful signals are reliable They predict that a particular event or stimulus will follow An association between a CS and US can be lost If the CS is no longer reliable, the subject stops responding
Extinction
: the loss of the conditioned response Ayo 教材 ( 動物行為學 2010) 15
The adaptive value of classical conditioning
Learning through classical conditioning provides
fitness advantages
to wild animals Few studies have addressed the
value
of classical conditioning in the everyday life of an animal The
territorial
and
reproductive behaviors
in
blue gouramis
(fish that inhabit shallow pools and streams in Africa and Southeast Asia)
Sperm production
in male
field crickets
Feeding behavior
in
honeybees
Ayo 教材 ( 動物行為學 2010) 16
案例： the blue gourami ( 三星鬥魚 )
A male blue gourami defends its territory aggressively The intruder may respond with a submissive posture or retreat If not, the contest escalates into a battle that can result in serious injury Dangerous fights evolve when the value of the resource is great Success is crucial for males because females won’t mate with a male without a territory Ayo 教材 ( 動物行為學 2010) 17
Blue gourami ( 三星鬥魚 ) learning
With victory, a male
blue gourami
gains an immediate competitive edge and experience Increasing the probability of winning future battles If a male could learn the
signals
(visual, chemical, or mechanical) that indicate the approach of a rival He might be better prepared for battle and gain a competitive edge Males that had been classically conditioned to associate a
light
with the imminent appearance of a
rival
were superior in territorial defense Ayo 教材 ( 動物行為學 2010) 18
The
“
winner effect
” Successful males are more likely to attract females But
excessive aggressiveness
harms mating success A male
conditioned
to expect the arrival of a female is less likely to attack her ( 研究者使用「光」 ) Classical conditioning pays off in reproductive success In nature, other
cues
(i.e. the shape of a gravid belly) for may be a reliable cue of a willing female’s approach.
Ayo 教材 ( 動物行為學 2010) 19
Ayo 教材 ( 動物行為學 2010) Conditioned
blue gourami
males have
higher reproductive success
20
案例： male field crickets
Female
field crickets
mate more than once Sperm from different males compete inside her body to fertilize her eggs A male increases his chances of fathering more offspring by transferring more sperm to the female Males can learn to associate
environmental cues
with the presence of
male competitors
Ayo 教材 ( 動物行為學 2010) 21
Field
crickets
learned about spatial cues that signaled the presence of a competitor. Male produced larger sperm packets in the environment associated with a competitor.
Lego bricks Ayo 教材 ( 動物行為學 2010) 22
案例： feeding honeybees
A
foraging Bee
can be rapidly conditioned to respond to odor When the antennae of a bee are touched with a sucrose solution (the US), the bee extends its proboscis to lick it (the UR) When an odor is presented just before the sucrose solution is presented, the bee rapidly forms an association between the odor (the CS) and the sucrose And begins to extend its proboscis to the odor alone Ayo 教材 ( 動物行為學 2010) 23
Operant (instrumental) conditioning
Another form of associative learning It is also called
trial-and-error learning
When cats inside a “
puzzle box
” accidentally hit a lever in the correct way, the door would open, and the cat would get to eat Over successive trials, a cat would get faster and faster at performing the correct behavior to release the latch This type of learning emphasizes that the animal
operates
on the environment to produce consequences Ayo 教材 ( 動物行為學 2010) 24
The Skinner box
Easier to use than Thorndike’s puzzle box It’s still used today A hungry animal inside the Skinner box must learn to manipulate a mechanism (pressing a lever or pecking a key) To get a food reward Ayo 教材 ( 動物行為學 2010) 25
Reinforcers
Reinforcer
: a stimulus, such as a bit of food, that changes the probability that an animal will repeat its behavior
Positive reinforcer
: increases the probability of a behavior being repeated, such as food offered to a hungry rat or a drink to a thirsty one
Negative reinforcer
it is removed : increases the probability of a response once If an unpleasant or painful stimulus stops when an animal performs a certain act, it is likely to repeat that action It’s different than
punishment
: a decrease in a response due to the presentation of an aversive stimulus Ayo 教材 ( 動物行為學 2010) 26
Timing of events is critical
In operant conditioning and classical conditioning When the animal spontaneously performs a behavior, reinforcement
must follow closely
When
reinforcement
is
withheld
, the response rate
gradually declines
and become extinguished Just as the strength of the conditioned reflex decreases when the CS is presented many times without the US Ayo 教材 ( 動物行為學 2010) 27
Shaping ( 導引化 )
Operant conditioning can be used to teach animals to perform novel and complex acts
Shaping
: used by Hollywood animal trainers At first, the trainer reinforces any approximation of the desired act Later, it requires better performances to get a reward To train a sea lion to jump through a hoop, first reward it for approaching the hoop Then reward it only when it swims through the hoop Raise the hoop on successive trials Offer the sea lion a fish only when it makes the leap Ayo 教材 ( 動物行為學 2010) 28
Reinforcement( 增強作用 ) schedules
Realistically, the reward does not follow every performance of an act
Reinforcement schedule
rewards are offered : the frequency with which
Reinforcement schedules
Each can change
reinforcement schedule
has
predictable effects
On the rate of response and On how long the animal continues responding when it is no longer rewarded Ayo 教材 ( 動物行為學 2010) 29
Reinforcement schedules
A
continuous reinforcement schedule
: each occurrence of the behavior is rewarded Is best during the initial training to establish a response A
fixed ratio schedule
: the animal must respond a set number of times before being rewarded Very high response rates because the individual determines how quickly it will be rewarded A
variable ratio schedule
: the number of responses required for reinforcement varies randomly The individual is rewarded for fast responses The response persists even if the reward is withheld Ayo 教材 ( 動物行為學 2010) 30
Latent( 潛伏 ) learning
Sometimes animals learn
without any obvious reward
When animals learn important characteristics of their environment during unrewarded explorations, they can use this information later Familiarity with the terrain( 地形 ) improves survival Knowledge of the environment helps them evade predators Even ants gather information for
later use
Ants evaluate prospective nest sites based on a range of criteria: floor area, headroom, entrance size, darkness, hygiene, and the proximity of hostile neighbors Ayo 教材 ( 動物行為學 2010) 31
Ants keep track of local housing options
If a nest is destroyed, ants must quickly find another one Ants ignore a familiar but unattractive option in favor of exploring for a better one Even if they ultimately settle for something that is also unattractive In latent learning, animals didn’t learn an appropriate behavior
,
but they learned something about their environment So they could respond appropriately in a new situation Ayo 教材 ( 動物行為學 2010) 32
Ayo 教材 ( 動物行為學 2010) The experimental design for a study of latent learning in ants.
Alternative nest #1 was added for a week. Then a second nest (#2) was added.
Ants choose #2.
33
Social learning
Learning from others is not part of every animal’s behavioral repertoire Social species have a greater opportunity for
social learning
than do solitary species
Social learning
encompasses a broad range of phenomena Some of which suggest a higher level of
cognitive skill
能力 ) on the part of the animal than do others ( 認知 Animals may inadvertently provide information to other animals Or, individuals share information through specific signals Ayo 教材 ( 動物行為學 2010) 34
Social learning: stimulus enhancement
Information may not be actively communicated by one animal to another In
stimulus enhancement
: an animal may be attracted to a particular
object
because a conspecific is near it or interacting with it For example, rats learn dietary preferences( 飲食嗜好 ) from other rats by smelling their breath Ayo 教材 ( 動物行為學 2010) 35
Social learning: local enhancement
In
local enhancement
: an animal may be attracted to a particular
location
because a conspecific is there In nature, local and stimulus enhancement occur frequently during
foraging
Bumblebees (
大黃蜂
)
land on flowers already occupied by other bees Other animals also use conspecifics as cues to good foraging patches Ayo 教材 ( 動物行為學 2010) 36
Observational conditioning
A type of classical conditioning that occurs in social situations Animals learn to avoid dangerous situations by
watching
conspecifics
Rhesus monkeys
learn to fear and avoid
snakes
monkeys show their fear by watching other
Minnows (
米諾魚 ) learn to show fear responses to
pike (
狗魚 ) odor when they are paired with minnows that had experience with
pike Observational learning
does not assume that observers understand anything about the mental state of the animals they are learning from Ayo 教材 ( 動物行為學 2010) 37
Goal-directed
emulation
( 仿效 )
An observer seems to learn from observation
what goal
is to be achieved, but does not copy precisely what the demonstrator does Chimpanzees and children both watched an adult human retrieve(
回收
) artificial fruit from a clear plastic box Chimps showed
goal-directed emulation
: they directed their attention at the correct part of the box but did not imitate the action of the demonstrator Children imitated ( 模仿 ) the actions of the human exactly Ayo 教材 ( 動物行為學 2010) 38
Imitation ( 模仿 )
An observer copies exactly what a demonstrator does In the
two-action test
, the subject is presented with a task that has
two equally easy solutions
If subjects choose the solution they have seen demonstrated, it is evidence of
imitation
Observer
budgerigars
(
鸚鵡
, budgies) watched demonstrator budgies open a dish The observer used the same technique it had just witnessed Ayo 教材 ( 動物行為學 2010) 39
The adaptive value of social learning
The
adaptive value
of social learning is clear It
saves time and energy
that might be wasted as an individual learns the business of survival by trial and error Each individual may have the capacity to learn appropriate responses But it is more efficient and less dangerous to learn about the world from others Ayo 教材 ( 動物行為學 2010) 40
Animals learn from each other
Rats
learn about food by smelling each other’s breath and by observation A rat will try novel food if it observes another rat eating it Other species may learn routes to food from conspecifics
Guppies (
孔雀魚
)
learn a safe route by swimming in groups Animals also learn from
other species
Group-foraging
doves (
野鴿
)
in Barbados learn from other
doves
But territorial
doves
learn from
Carib grackles
, the species they feed with in mixed flocks Ayo 教材 ( 動物行為學 2010) 41
Traditions ( 傳統 )
Traditions
: spread through a group and are stable over time In England around 1921,
blue tits
learned to break into milk bottles to steal the cream, which floated to the top This spread through Great Britain as other birds acquired the habit Ayo 教材 ( 動物行為學 2010) 42
Traditions spread through groups of primates
Young
chimpanzees
in nature learn to use sticks and stems to gather termites after watching their mothers or other adults A young female
snow monkey
discovered that washing
sweet potatoes
(provided by researchers) in the sea cleaned them and enhanced the flavor by lightly salting it The tradition spread to others Ayo 教材 ( 動物行為學 2010) 43
Individual vs. social learning
It is hard to tell between individual learning (learning through one’s own experience) and social learning They may occur simultaneously For example, sweet potato washing may occur through stimulus enhancement A monkey picks up a dropped potato that has been washed, like the taste, and then is primed to learn to wash potatoes Differential reinforcement may also maintain the behavior Human caretakers give more sweet potatoes to members of the troop that wash them Ayo 教材 ( 動物行為學 2010) 44
Comparative studies of species differences
For decades, the dominant view in studying learning was that it occurred in essentially the same way across mammal species Many characteristics of learning are similar in many species In recent years, researchers have been intrigued not just by
similarities
across species, but also their
differences
Several studies documenting
differences
across species correlates with the
ecological conditions
they face Ayo 教材 ( 動物行為學 2010) 45
Learned knowledge is
not
genetically heritable
In some species, offspring learn from watching their parents A
jumping spider (
跳蛛
) (
椿象
)
can learn that
milkweed bugs
are not good to eat and ignores them This knowledge is not passed onto to the spider’s offspring Spiders must learn this for themselves Ayo 教材 ( 動物行為學 2010) 46
The ability to learn as a heritable trait
What is heritable, and subject to natural selection, is the
capacity
to learn The heritability of the
ability to learn
has been demonstrated
Fruit flies
that had learned the association between quinine and a particular flavor avoided that flavor, and laid their eggs on a neutral flavor After 15 generations, flies from these selected lines learned the task faster and remember it longer than were control flies Ayo 教材 ( 動物行為學 2010) 47
Learning has costs
Learning allows an animal to adjust its behavior to new situations. This ability may not always be advantageous - it has its
costs
It takes
time
to learn Innate behaviors save the time and trouble of making mistakes The ability to learn requires
neurons dedicated to the task
Neurons could be devoted to something else (i.e. large olfactory centers to detect and interpret scents left by prey) Learning has an “
operating cost
”—it takes energy to collect, process and store information Ayo 教材 ( 動物行為學 2010) 48
Species differ in learning ability
The environment and evolutionary history of a species influences how
learning
increases
Darwinian fitness
The relative number of offspring left by an individual There are
biological constraints
on learning Members of a particular species may be able to learn certain things and not others Ayo 教材 ( 動物行為學 2010) 49
Species differences in learning ability
Three related species of birds -
Clark’s nutcrackers
,
pinyon jays
, and
scrub jays
- cache (store) seeds The birds recover and eat the seeds during winter and spring when food is scarce They remember the exact locations, months after they’ve hidden them Ayo 教材 ( 動物行為學 2010) 50
←The Clark‘s Nutcracker (
Nucifraga columbiana
), is a large passerine bird, in the family Corvidae (
鴉科
).
↓The Pinyon Jay (
Gymnorhinus cyanocephalus
) is a jay between the North American Blue Jay and the Eurasian Jay in size.
← The Florida Scrub-Jay (
Aphelocoma coerulescens
) is one of the species of scrub-jay native to North America..
Ayo 教材 ( 動物行為學 2010) 51
Bird ecology influences seed storage
Clark’s nutcrackers
cache thousands of pine seeds in thousands of locations They live at high elevations with harsh, long winters They survive almost entirely on stored seeds
Pinyon jays
cache fewer seeds, closer to the collecting site Most of their winter diet consists of cached seeds
Scrub jays
store the fewest seeds Seeds comprise less than 60% of the winter diet Ayo 教材 ( 動物行為學 2010) 52
Seed caching
Ecological differences among caching species predict that: Species that rely on caching to survive the winter have a better spatial memory
Nutcrackers
and
pinyon jays
Depend most on finding stored seeds to survive the winter Did better than
scrub jays
in finding stored seeds Ayo 教材 ( 動物行為學 2010) 53
Histograms that show the accuracy with which
scrub jays
,
pinyon jays
, and
Clark’s nutcrackers
find their caches.
The solid bar, 15 holes were available for caching.
The striped bar, 90 holes were available.
Ayo 教材 ( 動物行為學 2010) 54
Learning is shaped by natural selection
•
Clark’s nutcracker
was the champion on a
spatial task
• There were no species differences in memory on a
nonspatial task
• Remembering
the color of a circle
was the species’ dependence on stored food
not
related to Ayo 教材 ( 動物行為學 2010) 55
Comparison of learning abilities among food-storing on spatial and nonspatial tasks. (a) location of circle (b) the color of circle.
Ayo 教材 ( 動物行為學 2010) 56
Comparison of learning abilities among food-storing on spatial and nonspatial tasks. (a) location of circle (b) the color of circle.
Ayo 教材 ( 動物行為學 2010) 57
Food storing in other species
supports the hypothesis that evolution shapes learning ability is found among other groups of related species
Titmice
and
chickadees
, a family of birds distinct from
nutcrackers
and
jays
, store seeds and insects They performed better than non-food-storing species in tests
Mammals
show a relationship between ecology and spatial skills The
Great Basin kangaroo rat Merriam’s kangaroo rat
does does not store food, while The
Merriam’s kangaroo rat
performed better on a spatial test than the
Great Basin rat
Ayo 教材 ( 動物行為學 2010) 58
The tits, chickadees, and titmice constitute Paridae(
山雀科
), a large family of small passerine birds which occur in the northern hemisphere and Africa. Most were formerly classified in the genus Parus.
The Black-capped Chickadee (
Poecile atricapillus
) is a small, common songbird, a passerine bird in the tit family (
山雀科
). It is the state bird of both Maine and Massachusetts, and the provincial Ayo 教材 ( 動物行為學 2010) bird of New Brunswick in Canada.
59
Kangaroo rats, genus Dipodomys, are small rodents native to North America.
Dipodomys microps
Chisel-toothed Kangaroo Rat
Dipodomys merriami
Mearns Ayo 教材 ( 動物行為學 2010) 60
Evidence of cognitive abilities in animals
Some scientists wonder whether animals have
mental experiences
i.e. thoughts and feelings How can we know whether other animals think or whether they are self-aware?
Evidence for cognitive abilities(
認知能力
) includes: Tool use ( 工具使用 ) Insight ( 洞察力 ) Detours ( 繞道 ) Understanding abstract concepts( 抽象概念 ), including self-awareness ( 自我意識 ) Ayo 教材 ( 動物行為學 2010) 61
Tool use ( 工具使用 )
Tool use
: the use of an object in order to obtain a goal Once considered a hallmark trait that separated humans from other animals, tool use is now known in many species For example, a
sea otter
( 海獺 ) uses a rock to break a clam shell A
vulture (
禿鷹
)
open drops a rock on an egg, which cracks Ayo 教材 ( 動物行為學 2010) 62
Does an animal have to understand how a
tool
works in order to use it?
Tool
use seems to demonstrate a high level of cognition ( 認知 ) Animals using a
tool
to solve a problem often appear to be thinking it through Ayo 教材 ( 動物行為學 2010) 63
Sultan
’
s tool use
Sultan learned to use a stick as a tool to rake in a banana from outside his cage He learned to join two sticks to reach a banana When the sticks separated, he immediately rejoined them This is evidence that he understood that joining two poles increased his reach Ayo 教材 ( 動物行為學 2010) 64
Insight ( 洞察力 )
Insight:
a situation where a flash of understanding occurs As seen in the
suddenness
of Sultan’s solution to obtaining a banana The animal may see
new relationships among events
And consider the problem
as a whole
Ayo 教材 ( 動物行為學 2010) 65
Animals may form a mental representation of the problem and then
mentally work through solutions
But, all the details of an animal’s prior experience need to be known to understand what it knows when it manipulates objects Seemingly insightful behavior might be specific stimulus–response relationships
learned
through
operant conditioning
To reach meat suspended on a string, a
raven
(
烏鴉
) had to repeatedly pull up a loop of string, step on the loop, and then reach down and pull up another loop Some birds can be taught by
operant conditioning
Ayo 教材 ( 動物行為學 2010) 66
Raven ( 烏鴉 )
Raven is the common name given to several larger-bodied members of the genus Corvus (
烏 鴉
).
New Caledonia crows→
Ayo 教材 ( 動物行為學 2010) 67
Insightful behavior in common ravens (
烏鴉
)
But, some ravens solved the problem immediately Without going through a learning process It is
unlikely
that this complicated behavior was learned, was genetically programmed, or occurred by chance The ravens apparently have the ability to find
insightful solutions
to
new problems
, using string as a tool Ayo 教材 ( 動物行為學 2010) 68
Insight alone may not always be enough
Familiarity with a simpler task may be required to succeed on a new task If a string with food was looped up and through the cage, then down again The birds had to pull the string
down
to raise the meat
Ravens
familiar with the
pull-up task
could quickly do the
pull-down task
But
naive birds could not
Ayo 教材 ( 動物行為學 2010) 69
Animals may understand
causal relationships
New Caledonia crows
make and use tools out of twigs and leaves In the lab, one crow made a
hook
of wire to extract a
bucket (
提桶
)
from a straight piece from a tube
Crows
may understand that tools can be used to obtain out-of-reach objects, even other tools Video cameras attached to wild crows revealed that crows
keep good tools
for future use Ayo 教材 ( 動物行為學 2010) 70
Species differ in their ability to understand
Capuchin monkeys (
卷尾猴
)
learned to push a stick into a tube to push a
peanut
out When a trap was placed into the tube, the monkey had to push the stick into the correct end
No monkeys
fully grasped the task
Chimpanzees
, in contrast, showed more understanding of the task
Human children
under three years old behave more like
capuchins
Ayo 教材 ( 動物行為學 2010) 71
Capuchin monkeys
卷尾猴屬（學名：
Cebus
），是卷尾猴 科的一屬，也叫懸猴， 分佈於北起中美宏都 拉斯，南到南美中部 （巴西中部、秘魯東 部和巴拉圭）的狹長 地帶。
Ayo 教材 ( 動物行為學 2010) 72
The tube task but with a trap added. The stick must be inserted in a particular end of the tube. Here, a capuchin monkey is about to make an error.
Ayo 教材 ( 動物行為學 2010) 73
黑猩猩
(
Chimpanzees)
Ayo 教材 ( 動物行為學 2010) 74
Tool use in humans
Tools
are of great importance in human evolution We can gain some
insight
into our own past by examining the behavior of our close relatives
Chimpanzees
are accomplished tool users They use
sticks
to forage for termites and
rocks
as a hammer and anvil to pound open nuts
Foraging
and
hunting
were probably the first contexts in which our ancestors used
tools
Ayo 教材 ( 動物行為學 2010) 75
Detours ( 繞道 )
Detouring
: the ability to identify an alternative route to a reward when the direct route is blocked Animals can improve on detour tests with
experience
Dogs
are not very good at solving detour problems often
dig under a fence
instead of
going around
it
Squirrels
are very good at solving detour problems They choose certain branches to get from tree to tree Ayo 教材 ( 動物行為學 2010) 76
Quail (
鵪鶉
)
and
herring gulls
easily solved a detour problem that required them to walk around a barrier, but
canaries (
金絲雀
)
could not
Canaries
can fly around an object Ayo 教材 ( 動物行為學 2010) 77
How do animals solve detour tasks?
Portia
jumping spiders (
跳蛛
) excel (
勝過
) at detours Their large anterior eyes are specialized for acute vision They hunt other spiders by climbing into their webs and luring( 誘惑 ) them with signals similar to those of prey
Portia
can spot spider webs from some distance away To reach its prey it must perform a
detour
Ayo 教材 ( 動物行為學 2010) 78
Portia
jumping spiders
Portia is a genus of jumping spider which feeds on other spiders (araneophagic). They are remarkable for their hunting behaviour which suggests they are capable of learning and problem solving, traits normally attributed to much larger animals.
Ayo 教材 ( 動物行為學 2010) 79
Portia
s
piders use detours
Before setting out,
Portia
can Choose correctly between detours that lead to prey versus those that do not
Portia
looks at the lure(
目標
), then slowly scans along the features of the potential route If the route ends, the spider turns back to look at the lure again, and begins again Ayo 教材 ( 動物行為學 2010) 80
Understanding abstract concepts
Many animals can discriminate relative numbers i.e. is there more fruit on this tree or that tree?
The ability to count is more difficult To assign a tag such as “1, 2, 3,” to individual quantities The ability to count things demonstrates some understanding of the
abstract concept
of numbers Ayo 教材 ( 動物行為學 2010) 81
Alex, a remarkable parrot
Alex, an African grey parrot, learned labels (names) for over 35 different objects He could identify, request, refuse, or comment on more than 100 different objects Ayo 教材 ( 動物行為學 2010) 82
Alex understood abstract concepts
He used language to show he understood abstract concepts He could: Say how many items were in a group of up to
6 items
, even if the objects were scattered around a tray Count items in a confounded number set (items that vary in more than one characteristic) Add up the total from two sequentially presented collections Alex had some understanding of the concept of
zero
Ayo 教材 ( 動物行為學 2010) 83
Sheba
, an amazing chimpanzee
Using a touch screen,
Sheba
can indicate the Arabic numeral that describes a group of objects She can
add numbers
: If three small groups of objects are put in three separate places around the room, she can visit them in turn and then correctly choose the numeral that represents the
sum
If the three groups of objects are replaced with numeral cards she can still choose the numeral that represents the
correct total
Ayo 教材 ( 動物行為學 2010) 84
Animals understand concepts
Alex understood the
concepts of same and different Brain size
does
not
reliably indicate
the ability of animals
to do tasks such as concept learning Alex’s brain was the size of a walnut
Bees
can learn to distinguish between same and different
Pigeons
form concepts such as “tree” or “water” or “human” Pigeons can identify an example of a particular category, such as a person They recognize water in various forms: a droplet, a river, a lake Ayo 教材 ( 動物行為學 2010) 85
Self-recognition and perspective taking
The “
mark test
” examines if an animal recognizes itself After a mark is placed on its face, the animal is shown a mirror If the subject recognizes itself, it sees that it has an odd mark, and touches and grooms toward the marked area Species that have passed the mark test
Chimpanzees
Dolphins
turn their bodies to inspect marks in the mirror
Asian elephants
touch marks with their trunks Ayo 教材 ( 動物行為學 2010) 86
Do animals have a concept of self?
We can ask whether animals can take the perspective of other individuals And understand what others know and do not know When given a choice, subordinate
chimpanzees
selected a piece of food not visible to the dominant individual The subordinate was aware of which piece of food the dominant could see Ayo 教材 ( 動物行為學 2010) 87
The
“
theory of mind
” There is a growing literature on “seeing and knowing” and the attribution of knowledge and mental states Many species have been tested with a range of
clever experiments.
A compelling idea about the evolution of the “
theory of mind
:” is that it is driven by
social complexity
The social environment creates new selection pressures for the evolution of “
social intelligence
” The ability to learn and keep track of relationships among other individuals may be evolutionarily advantageous Ayo 教材 ( 動物行為學 2010) 88
Summary
Learning
: a change in behavior as a result of experience Excluding maturation of the nervous system, fatigue, or sensory adaptation Learning is divided into:
habituation
,
classical conditioning
,
operant conditioning
,
latent and social learning
The ability to learn has a
genetic basis
Animals demonstrate
cognitive skills
in tasks besides learning Ayo 教材 ( 動物行為學 2010) 89
Tools
are objects that an animal uses to reach a goal
Detour behavior
: an animal takes an indirect route to a goal Many animals understand
abstract concepts
Some animals
recognize themselves
Some understand that others do not have the same knowledge Ayo 教材 ( 動物行為學 2010) 90
問題與討論
Ayo
台南
NUTN
站
http://myweb.nutn.edu.tw/~hycheng/
Ayo 教材 ( 動物行為學 2010) 91