Table of Content
Module 1
Chapter 1: Introduction …………………………………………………………………… 5
Chapter 2: Behaviorism …………………………………………………………………. 23
Chapter 3: Cognitive Social Learning …………………………………………………..35
Chapter 4: Memory & Planning ………………………………………………………….59
Focus Terms for Module 1 ………………………………………………….73
2
LECTURE NOTES
3
4
Introduction
Brief History of Study of Animal Psychology

o
Charles Darwin
o
George Romanes

Founder of comparative psychology

Methodology –

& laboratory experimental method
o
Behaviorists seek to
by placing
animals in barren environments in which they can do little else than what the
experimenters want
o
Edward Thorndike

5

o
John Watson

o
, not observations & collection of anecdotes
Behaviorism – eschews reference to
terms
B.F. Skinner

o
Ivan Pavlov


Animal

Wolfgang Kohler

Animals’ inner thoughts (
(
) & feelings
)
6

(Behavioral Biology) – study of animal behaviors
o
Most ethologists are
o
Konrad Lorenz
, not psychologists
7

Combination of
&
+ neuroanatomy, ecology, & evolution

o
More interested in
Methodology:

Fieldwork

The days of a few scribbled observations in a notebook are gone nowadays field work is very

Data collection is
and systematic, typed
into handheld digital devices, and complemented with fecal & urine
samples that allow DNA analysis & hormone assays

They offer hints & suggestions but rarely allow firm
- e.g. One may encounter wild chimpanzees who crack nuts with
stones, but it is impossible to know how they discovered this
technique or how they learn it from one another
8

Carefully-controlled experiments

Captive animals (e.g. apes) under enlightened conditions (e.g. a
sizeable group in a spacious outdoor area) have the advantage of
providing a close-up look at
that one can't get in the field

Here, animals can be watched and videotaped
than is possible in the wild where animals
often disappear

of the two methods

Observations in the field have often inspired experiments in the lab

Observations in captivity (e.g. chimpanzees reconcile after fights)
have
stimulated
observations
in
the
field
on
the
same
phenomenon

Cognitive ethology
o
of evolutionary psychology, animal cognitive
psychology, & ethology
o
Systematic description + observation of behavior
o
Every species has its own
, its
own Umvelt (see next section) which dictates what it needs to know in order
to make a living
9
The Importance of "Umvelt" in Psychology of Animals

Jakob von Uexküll:

Not "
- animal point of view
" - the habitat that an organism needs
for survival

An organism's
not available,
comprehended, & discernible to other species
10
o
E.g. some animals perceive ultraviolet light, while others live in a world of
smells or like the star-nosed mole, feel their way around underground

Nagel: we can never fully understand other species's Umvelt
o
Nagel: "What is it like to be a bat?" (very famous)
o
Not trying to know how a human would feel as a bat, but how a bat feels like
a bat (whose major sense is
which is beyond our
comprehension)
o
Nagel concluded that we would
b/c we have no
way of entering the subjective life of another species

de Waal: Humans can try to
o
the Umvelt of other organisms
Even though we can't feel what other species feel, we can still try to
our own narrow Umvelt and apply our imagination to theirs
11
o
Humans inhabit the same Umwelt as
as we
process our surroundings in similar ways - stereoscopic vision, grasping
hands, ability to climb & jump, & emotional communication via facial muscles
o
But
? Every species
deals flexibly with the environment & develops solutions to the problems it
poses; each one does it differently
Anecdotes: The Importance of Stories


Some scientists dislike or ignore anecdotes b/c:
o
“Merely stories”, not “
”
o
Not reproducible & too tainted with
But systematic analyses of anecdotes can lead to data that are reproducible by
that mimic the anecdotal situations

Beckoff: anecdotes are central to the study of animal behavior and especially
animal emotions b/c the latter

As more and more stories are accumulated, a solid behavior data can be
developed and this can be used to

The more stories gathered showing the same thing, the less likely it will be that
has influenced the collection of data or the
conclusions drawn

What makes a “good” anecdotal account? (Nussbaum)
o
The author pursues
12
o
She displays
to a good
theoretical end
o
She neither withholds
,
nor uses it loosely
o
She is keenly aware that the animal has
but is
o
She gives an account of the emotions of a specific animal, and not a fanciful
of emotion
Anthropomorphism & Misrepresentation

Anthropomorphism
o
Tendency to attribute
to
nonhuman animals

,
&
everyday
way
of
considering animals

And as humans who study other animals, we can only describe &
explain their behavior using words with which we’re familiar from a
point of view

o
Anthropomorphism is
But have to bear in mind that animals are not little people – maybe
emotionally satisfying to treat them that way, but must consider psychology of
animals
13

When a scientist tries to figure out what’s happening in a dog’s head,
she has to be anthropomorphic, but she tries to do it from a

Just b/c she says a dog is happy or jealous, this doesn’t mean he’s
happy or jealous as humans are
o
Anthropocentrism is problematic only when the human-animal comparison is
a stretch, such as with regards to species
 Kissing

Kissing gouramis don't really kiss in the same way & for the same
reasons that humans do - adult fish sometimes lock their
protruding mouths together to

Apes, however, do
after
a separation by placing their lips gently on each other's mouth or
shoulder & hence kiss in a way & under circumstances that greatly
resemble humans kissing

Tickling & laughing

When young apes are being tickled, they make breathy sounds
with a rhythm of inhalation & exhalation that

One
the term laughter for
this behavior as too anthropomorphic
14
o
Anthropomorphism is not always as problematic as people think - to rail
against it for the sake of scientific objectivity often hides one's discomfort with
the notion of

When we are considering species like the apes, which are known as
"
" (humanlike), however, anthropomorphism
is in fact a

Dubbing an ape's kiss "mouth-to-mouth" contact so as to avoid
anthropomorphism deliberately obfuscates the

Our terminology should honor the obvious
o
(Burghardt) – Anthropomorphism’s place in the scientific studies of animals

Being anthropomorphic is a
to make
the thoughts and feelings of other animals accessible to humans

Anthropomorphic accounts can be an
to behavior

Anthropomorphic accounts not necessarily projection of human values
or inserting something human into animals, but simply be recognition of
& using
human language to communicate what we observe
15

E.g. Saying that animals "plan" for the future or "reconcile" after fights is
more than anthropomorphic language: these terms propose

If primates are capable of planning, they should hold on to a tool
that they can use only in the future

If primates reconcile after fights, we should see a reduction of
tensions as well as improved social relationships after opponents
have made up by means of friendly contact

Critical anthropomorphism is a valuable source of
if it serves as a means rather than an end

Misrepresentation
o
A form of anthropomorphism but one that focuses on an animal’s
human qualities as an excuse for killing it,
brutalizing it, or being indifferent to its welfare
o
The animal becomes a projection of our own
o
This sort of
is particularly
prevalent in human attitudes to predators, pests, and scavengers – species
that either compete with us directly or which survive off the surpluses of
human cultures
o
E.g. wolves – malevolent, merciless, insatiable killer, the archetypal bête
noire – extermination and torturing of wolves, leading to their near extinction
16
Anthropocentrism, Human Exceptionalism, Anthropodenial

o
Seeing animals
o
Start with human minds &
E.g. Are they intelligent or conscious? Do they think or
feel?
o
than anthropomorphism – since we
More
are human, we can never be entirely free from anthropocentrism
o
Anthropocentrism sometimes makes us forget that we, humans, are fellow
animals too – our species is different,

o
"Human is the only animal that .... " - most of these claims are
o
A lot of times when we fail to find a capacity in a given species, instead of
congratulating ourselves on our human exceptionalism, we should ask
ourselves if (1) we have
o
something & (2)
Examples:
 Gibbons

Once considered backward primates (the "
")
even though they belong to the same large-brained family as
humans & apes
17

When presented with problems that required them to choose
between various cups, strings, & sticks, they
compared to other species

Tool use was tested by dropping a banana outside their cage &
placing a stick nearby: all they had to do to get the banana was
pick up the stick to move it closer, but they
(unlike chimpanzees & many monkeys)

Reason why they can't do it? They are exclusively
they are
;
that propel themselves through trees
by hanging by their arms & hands

Gibbon hands, specialized for this kind of locomotion, act more like
hooks than the versatile grasping & feeling organs of most other
primates

Beck: Instead of presenting objects lying on a surface to them, he
elevated them to the animal's
them easier to grasp

The gibbons solved all these problems
demonstrating the same intelligence as other apes

Their earlier poor performance had had more to do with
than with their mental powers.

Elephants

For years elephants were believed to be
using tools
18
, making

The pachyderms
the same aforementioned out-
of-reach banana test, leaving the stick alone

Their failure could not be attributed to an
from a flat surface b/c elephants are ground
dwellers & pick up items all the time, sometimes tiny ones

Conclusion - elephants didn't get the problem

Problem:

Elephant's grasping organ is also its
- elephants
use their trunks not only to reach food but also to sniff &
touch it

But picking up a stick
; even when they bring the stick close to the
food, it impedes their feeding & smelling it

Alternative experiment (Foerder & Reiss)

Hang food
above an elephant's
enclosure just out of his reach

Several sticks & a sturdy square box were presented to the
elephant

The elephant
but began
with his foot many times in a
straight line until it was right underneath the fruit

He then stood on the box with his front legs, which enabled
him to reach the food with his trunk
19

Elephants
- if they
are the right ones

Anthropodenial (de Waal)
o
- a priori rejection of humanlike traits in other
animals or animallike traits in us
o
Anthropomorphism & anthropodenial have an

The
:
another species is to us, the
anthropomorphism will assist our understanding of this species and the
will be the danger of anthropodenial

Conversely, the more
a species is from us, the
that
anthropomorphism
will
propose
questionable similarities that have come about independently e.g.
saying that ants have "queens," “soldiers," and "slaves"
Biocentrism & Anthroharmonic Approach

Shift of paradigm from anthropocentrism to biocentrism
20
o
Biocentrism

Humans are members of a
along with all other species, & on equal terms.

This community consists of a
between all members, both physically, & in terms of
relationships with other species

Every organism is a "
",
i.e., each organism has a purpose and a reason for being, which is
inherently "good" or "valuable"

Humans are not
to other species
21

Anthroharmonic Approach (Scharper)
o
An approach to the study of
o
Acknowledges the importance of the human and makes the human
fundamental but
o
Humans and animals are
,
and we complete one another
22
Behaviorism
1.
(Pavlovian/Reflex/Respondent/Type
I conditioning)
2.

(Instrumental/Type II conditioning)
1 and 2 =
o
“
”
o
: Stimulus-Response

Animals’ ability to recognize
in
the world around them

Powerful mechanism that animals use extensively to steer themselves
through life

Animals may or may not understand the relation between cause &
effect; only
, not
understand

Classical Conditioning

Pavlov's original experiment
23
o
In his initial experiments, Pavlov presented a
(bell
ring) and then gave the dog food; after a few repetitions, the dogs started to
salivate in
o
to the stimulus.
3 Stages of Classical Conditioning
24

Pavlov called:
o
The food -
(US) b/c its effects did
not depend on previous experience
o
The response to the US (salivate) –
(UR)
o
The stimulus (bell ring) -
(CS) b/c
its effects depend on its association with food
o
The response to the CS (salivate) (CR)

Definitions
o
Classical conditioning occurs when a conditioned stimulus (
paired with an unconditioned stimulus (
o
The conditioned stimulus (CS) is usually a
sound of a tuning fork)
o
The unconditioned stimulus (US) is
(e.g., the taste of food)
25
) is
)
stimulus (e.g., the
o
The unconditioned response (UR) to the US is an
(e.g., salivation)
o
After
is repeated (some learning may occur already
after only one pairing), the organism exhibits a conditioned response (CR) to
the CS when the CS is presented alone
26
Operant Conditioning (Instrumental Conditioning)

Introduction
o
Operant Conditioning deals with operants –
that have an effect on the surrounding environment
o
Thorndike’s

Behavior which is reinforced tends to be
(i.e. strengthened)

Behavior which is not reinforced tends to be
(i.e. weakened)
o
B.F. Skinner - father of operant conditioning
27

Studied operant conditioning by conducting experiments using animals
which he placed in a "

"
Concepts and procedures
o
Core tools to modify operant behavior:

Reinforcement - stimulus that
probability that a
preceding behavior will be repeated
reinforcement


When a behavior (response) is followed by the
(hence "positive") of a stimulus ("positive" does not
necessariily mean "good" or "favorable")

This

E.g. if a rat in a Skinner box gets food when it presses a
the frequency of that behavior
lever, its rate of pressing will go up
28
reinforcement (escape)


When a behavior (response) is followed by the
of a stimulus (which is aversive, hence its removal is
"rewarding" to the animal or person)
("negative" – removal of a stimulus)

This

E.g. In the Skinner box experiment, the aversive stimulus
that behavior's frequency
might be a
continuously
sounding inside the box; negative reinforcement would
happen when the rat
turning off the noise
29
,

Punishment - stimulus that
probability that a
preceding behavior will be repeated
punishment


When a behavior (response) is followed by the
of a stimulus (e.g. a shock or loud noise) ("addition" - hence
positive)

E.g. Every time a dog barks, his owner squirts water on his
face.

This results in a
n that behavior
(penalty

or
punishment by contingent withdrawal)

When a behavior (response) is followed by the
of a stimulus (removal, hence negative)
e.g. taking away a dog's toy following an undesired behavior

This results in a
in that behavior
30
Stimulus Presented
Stimulus Removed/Withheld
Behavior
Encouraged
Behavior
Suppressed
Stimulus Presented
Behavior
Encouraged
Behavior
Suppressed
31
Stimulus Removed/Withheld


When a previously reinforced behavior is
with either positive or negative reinforcement


During extinction the behavior becomes
E.g. a rat is first given food many times for lever presses; in "extinction",
no food is given → rat continues to press
and eventually stops (lever pressing is said to be
"extinguished")
32
Practical applications of operant conditioning

(successive approximation)
o
Conditioning method used in
& in teaching non-
verbal humans
o
The trainer starts by identifying the desired final (or "
behavior
o
Next, the trainer chooses a behavior that the animal or person
with some probability
33
")
o
The form of this behavior is then gradually changed across
by reinforcing behaviors that approximate the target
behavior more and more closely.
o
When the target behavior is finally emitted, it may be strengthened &
maintained by the use of a
34
Cognitive Social Learning
Problems with behaviorism

Only concerned with
, as opposed to internal
events like thinking and emotion (e.g. Skinner did not say that the rats learned to
press a lever b/c they wanted food)

Behaviorism fails to take into account the role of
in learning → incomplete explanation of the learning
process in humans & animals

Despite the range of potential problems that can confront an animal, behaviorists
argue that they are all solved in the same manner – animals are assumed to
until by
the
correct response is made and reward is forthcoming
Cognitive-social learning

Learning is
- it is a cognitive
process that takes place in a social context

Cognitive processes –
processes (e.g. thinking, knowing,
problem-solving, remembering)

Reinforcement plays a role in learning but is
for learning

Examples of cognitive-social learning
o
35
o
o
o
o
Insight

Definition:
o An
that leads to a
problem being solved – the change in behavior is sometimes attributed to a
period of thought followed by a flash of inspiration
o Insight – “what leads to what”
; confirmation of expectancies –
leads to learning
o Therefore learning is not just about simple conditioning, but
generated by confirmed expectations (e.g. knowing how to
navigate the maze or how to use tools after exploring them)
 Characteristics of insight learning
o Transition from pre-solution to solution is
o Insight based on
performances
o Solutions gained by insight are
o Solutions gained by insights are
(
to other problems
)
36
 Wolfgang Kohler:
by chimpanzees – not just
o Insight plays a role in
stumble on solutions through trial and error, but a
of problems
o Examples:


A banana was hung from the roof of an enclosure, out of reach of the
chimps; the enclosure contained three packing boxes

Initially a chimp (named Sultan) would jump or throw things at the banana
or drag humans by the hand toward it in the hope that they’d help him out,
or at least be willing to serve as a footstool.

If this failed, he would sit around for a while without doing anything until he
might hit at a

All of a sudden, Sultan stacked three boxes on top of each other (the
heaviest one at the bottom and the smallest one on top) & reached the
banana
37

Sultan was put inside an enclosure & some pieces of fruit were placed out

of reach beyond the bars
Sultan was provided with 2 hollow bamboo sticks, each of which was too

short to reach the highly desired food items
At first Sultan tried to reach the fruit with his hands, then with each of the

sticks separately – all these efforts were futile & Sultan gave up
Then Sultan started playing with the sticks for a while before inserting the

thinner of the two sticks inside the thicker one – armed with this now
lengthened stick, Sultan rushed back to where the fruit was & raked it all in
rapidly

Insight:

A sudden insight may explain how Sultan put together what he knew
about bananas, boxes, and sticks to produce a
that would take care of his problem

Kohler ruled out imitation and trial-and-error learning, since Sultan had
had
with these
solutions nor ever been rewarded for them

The outcome was unwaveringly
in
which the ape kept trying to reach his goal despite the numerous
stacking errors resulting in the collapse of his towers

Once a solution was discovered, the ape found it easier to solve similar
problems, as if they had learned something about the
38

Other examples of insight o Younger female chimpanzees collect water in their mouths for an aging female
chimpanzee who can barely walk anymore, spitting it into her open mouth so that
she doesn’t have to walk all the way to the spigot
o Goodall: Madame Bee, a wild chimpanzee, had become too old and weak to
climb into fruiting trees; she would patiently wait at the bottom for her daughter to
carry down fruits, upon which the two of them would contentedly munch together
o In such cases, apes grasp a problem and come up with a fresh solution, but the
striking part here is that they perceive another ape’s problem
(
)
Causal Inference & Folk Physics

Causal inferences
o The term “insight” is now
in discussions of
problem solving by animals
o As an alternative, it has been proposed that animals have some understanding of
&
that
they
can
draw
inferences based on this understanding to solve problems

Chimpanzees: Premack
o A single subject would be shown an array of objects as the following:

To
, the chimpanzee was required to replace the
strange shape in the upper row with the knife from the lower row
39

The choice of the knife was intended to reveal that the ape understood this
object causes an apple to be cut in half
o Results:
o Even when the apple was replaced by
& a ball
cut into pieces, the subjects still made the correct choice even though the
subjects had rarely seen a knife & a ball together

Bird
o Heinrich et al: raven

Experiment 1:

Hand-reared ravens were presented with a piece of meat hanging from a
perch

The meat could not be obtained by flying towards it & clasping it in the
beak; instead, to reach the meat, some birds settled on the perch where
the string was attached & grasped the string below the perch with their
beak &

To stop the meat falling back, they placed a foot on the string & then let it
drop from their beak whereupon they bent down to grasp again the string
below the perch
40

This operation was repeated until the meat was near enough to be
grasped directly with the beak
41

Experiment 2:

Meat could be retrieved by standing on the perch &

Birds who had mastered the
were also
adept at mastering this new task

Birds
of pulling string
never mastered this second task

Conclusion

Ravens have some kind of understanding of
relationships, i.e. an apprehension of a cause-effect relations between
string, food, and certain body parts (appreciation of folk physics)
Latent (Exploratory) Learning

Tolman: learning in the absence of an obvious reward
o Learning is
, not shown in behavior
o Rats can learn a route in a maze without obtaining reinforcement:

Group 1 (Control group) – rat was fed upon reaching goal

Group 2 – not rewarded at all

Group 3 – not rewarded for trials 1-10, but rewarded on trials 11-22

Group 4 – rewarded on trials 1-10, but not rewarded on trials 11-22
42
o Results

Group 1 –
running the maze than
Group 2 due to availability of the reward

Group 3 – key condition; almost immediately after receiving food, Group 3’s
errors dropped to the level of Group 1

The rats
how to navigate their way through
the maze all along, yet they weren’t sufficiently motivated to do so until a
reward was presented

They were able to use a cognitive map of the maze when the rewards
were introduced

Group 4 - Group 4’s errors increased quickly when the possibility of a reward
was removed, again illustrating the
of
rewards

Exploratory learning
o “
” – stimuli without patent reward
o Motivation in latent learning – get to know the
43
o Very important for organisms in real life – after establishing the
(e.g. food or a hole), the animal can reasonably
expect it to be there when it returns
Imprinting

Definition
o
learning (occurring at a particular age
or life stage) that is rapid & independent of the consequences of behavior
o Imprinting =
o “
” – during a particular short
period of its ontogenesis (development), an organism suddenly “learns” what to
do
o
are released in response to learned
stimulus
o Most imprinting promotes
animals &
shapes their future breeding activities

Filial imprinting
o When a young animal acquires several of its
from its parent.
o Nidifugous birds (e.g. chickens, ducks, geese) imprint on their parents &
44
o Lorenz demonstrated how incubator-hatched greylag geese would imprint on the
first suitable moving stimulus they saw within a
(between 13 to 16 hours shortly after hatching)
o The goslings would imprint on Lorenz himself (to be more specific, on his wading
boots)
o Filial imprinting of birds was a primary technique used to create the movie
Winged Migration (Le Peuple Migrateur), which contains a great deal of footage
of migratory birds in flight – the birds imprinted on handlers, who wore yellow
jackets and honked horns constantly; the birds were then trained to fly along with
a variety of aircraft, primarily ultralights
Social Learning

Diet Selection & Foraging
– an increase in the tendency
o
to approach an object as a consequence of observing another animal interact
with it

McQuoid & Galef (1992) – Burmese jungle fowl

Hungry fowl were allowed to explore an enclosure in which four bowls
were placed in fixed positions

Food was consistently available in one of these bowls and eventually all
the subjects approached this bowl whenever they entered the enclosure

Towards the end of this training, another group of fowl (
watched the first group (
)
) as they ate from the bowl
45

When the observers were themselves permitted into the enclosure, they
showed a
for the bowl from which the
demonstrators had been seen to eat
o Neophobia – a reluctance to approach something that is novel

Galef et al (1)

A
rat was allowed to eat food with a distinctive
flavor (either coca or cinnamon)

An
rat was then placed in the company of the
demonstrator for 30 minutes, but in the absence of any food, before being
allowed to choose between food flavored with either cocoa or cinnamon

The observers preferred the food that was of
as that consumed by the demonstrator

This preference has been shown to occur even if the demonstrator eats
the flavored diet up to 4 hours before its encounter with the observer

After a single interaction with a demonstrator, the
endures for about 12 hours

If the interaction between the observer and demonstrator is repeated on a
number of occasions, or the interaction is repeated with a number of
different demonstrators, then the preference lasts considerably

Once acquired, the flavor preference is
subsequently exposing the animal to other types of food
46
by

Galef et al (2)

An observer rat was placed into a bucket of the apparatus, and an
was placed into a wire-mesh basket

Some demonstrators had food dusted on their faces and others had food
placed directly into their stomachs through a tube

In both cases, the observers subsequently showed a
that
had
just
been
fed
to
the
demonstrator

However, if the
of the demonstrator was dusted with food,
and placed foremost in the basket, then only a
for the food was demonstrated

If a
, rather than a rat, was placed in the
basket, then despite being dusted with food, there was
that this resulted in a change in the attractiveness of the food

Conclusions:
 The demonstrator
conscious
if it is to encourage the development of a food preference in another rat
47
 The observer must
, and
preferably its front rather than its rear end
o Social influences on diet selection have been demonstrated in blackbirds, sheep,
goats, cats, hyenas, pigs, rabbits, mice, & monkeys

Choosing a Mate
o Female guppies were allowed to observe 2 males: one male was alone & the
other with a female companion – when the
observer was given a
choice between the 2 males, she preferred the one
o Similar findings for Japanese quail – however, if the procedure is reversed so
that a
is given a choice between a female he has seen in isolation
and a female he has seen courting & mating with another male, then the
observer will tend to prefer the female he had seen

Fear of Predators
o Monkeys’ fear of snake

When an adult free-ranging monkey suddenly encounter a snake, it will
attempt to flee from the snake, its facial expressions will indicate fear, it will
look at the snake, and it will make specific alarm calls

A monkey that has been reared in the laboratory will show virtually

Fear of snakes by monkeys is not innate but
48

Laboratory-reared monkeys
of a wild-
reared monkey to a snake – on seeing its reaction, the observer responds in
a similar way; if subsequently the observer is exposed to the snake by itself, it
will show the same reaction too
Copying Behavior

Imitation & mimicry vs. Contagious behavior
– result in animals performing
o Both
new responses, & these responses are arbitrary, rather than being closely tied to
species-typical reactions to certain stimuli
– occurs when the response of one
o
animal triggers the same response in another animal (e.g. chorusing in dogs &
roosters)

true imitation


Humans have
to imitate
each other without there being any need for deeper reason e.g. yawning &
laughing


Birds feeding in flocks
Imitation
o Old definition = doing an act from

Naturalistic evidence

Blue tits & great tits in Britain
49
 Break through the foil tops of milk bottles to drink the cream at the top
 This skill is believed to have originated in a small group & its spread to
the rest of the population has been attributed to imitation

Primates
 Japanese macaque monkeys wash sweet potatoes before eating them
 “Imo” was observed to wash potatoes in the ocean before eating it
– removed
from potato
 Over the following years, many more monkeys took up potato
washing habit
 Ivory Coast chimpanzees use stones to
 Examples of "
" - habits learned from
others

Laboratory studies: Bidirectional control procedure: to demonstrate true
imitation (Heyes & Dawson)

Two subjects (rats in original study) put into a box containing two
compartments separated by a transparent wall
50

Animal on one side of the partition (the “
”) has a
joystick, which it can operate by pushing to the left or the right

Some demonstrators are trained to push the joystick to the left, for which
they receive a reward, others are trained to push it to the right to obtain
their reward.

Once a demonstrator knows what it has to do, a second animal – the
– is placed in the other half of the box, where
it can watch the demonstrator

When the observer has had a few sessions of watching the demonstrator,
it is given its own chance in the demonstrator’s half of the box

Rats who observed other rats push the joystick to the left
to push it to the left themselves - & similarly for rats who observed the
joystick pushed to the right
o New definition of imitation - "

"?
Requires individuals to
copy another's
specific techniques to achieve a specific goal

Merely duplicating behavior is not enough - it has to be done with

While imitation is common in lots of animals according to the old definition,
true imitation is

Examples:

Apes & children were prompted to imitate an experimenter - they'd watch
a human model open a puzzle box or rake in food with a tool
51

While the children copied the demonstrated action,
hence the conclusion that other species lack imitative capacities and
cannot possibly have culture

Problem: anthropocentrism

Whether apes imitate us or not is wholly beside the point

For culture to arise in a species, all that matters is that its members pick
up habits

Two ways to make a fair comparison in this regard:
1. Raise apes in a human home so that they are as
as
children around a human experimenter
2.
- to test a species with
models of its own kind

Human-raised apes

As good at imitating humans as are young children - apes, like young
children, are
and prefer to copy the species
that raise them

Under most circumstances, this will be their own kind, but if reared by
another species, they are prepared to imitate that one as well

Using human as models, they
to
brush teeth, ride bikes, light fires, eat with knife & fork (see mimicry below)

Other examples: dogs raised by cats show feline behavior such as sitting
in boxes, crawling under tight spaces, licking their paws to clean their face
52

Horner & Whiten: working with human-raised orphan chimps
 Experimenter poked a stick into holes in a large plastic box, going
through a series of holes until a candy would roll out; but only one hole
mattered
 If the box was made of black plastic, it was impossible to tell that some
of the holes were just for show; a transparent box, on the other hand,
made it obvious where the candies came from
 Handed the stick and the box, young chimps mimicked only the
, at least with the transparent box
 The children, on the other hand, mimicked
that the experimenter had demonstrated, including
- they did so even with the transparent box, approaching the problem
more like a magic ritual than as a goal-directed task
 Chimps fit definition of true imitation better than children as they were
showing
- the sort that pays close
attention to goals and methods
 Children only showed dumb copying or indiscriminate copying or
- fits human species's purported reliance on
culture b/c it makes us imitate behavior regardless of what it is good
for; we transmit habits in full without every individual making their own
ill-informed decisions
53

The Conspecific approach (de Waal & Whiten)

When chimps were given a chance to watch one another - apes truly do
ape, allowing behavior to be
within the group

Katie imitating her mother, Georgia who had learned to flip open a little
door in a box, then stick a rod deep into the opening to retrieve reward

Katie had watched her mother do this five times, following her every move
and smelling her mouth every time she got a reward

After her mother was removed to another room, Katie could finally access
the box herself; and even before the experimenter had added any
rewards, she flipped open the door with one hand and inserted the rod
with the other

Mimicry
o
of a demonstrator when it
does not result in any tangible reward:

It seems some arbitrary responses can be acquired through
rather than by trial and error

Reward is
- i.e. not necessary for the response to be
performed repeatedly

"Habits" & "fashions" - spread without any associated rewards, social learning
is truly social; it is about conformity instead of payoffs
54
o Bonding- and Identification-based Observational Learning (

Primate social learning stems from


) (de Waal)
born from the desire to act like others & to fit in
Explains why apes imitate their own kind
than the
average human, and why, among humans, they imitate only those whom they
feel close to

Also explains why young chimps, especially females, learn so much from their
mothers, and why high-status individuals are favorite models (ref: the
Beckhams, Kardashians, & Biebers)
o Examples of intra-species mimicry

Primates:

Mimicry is common in
e.g. hairstyles, accents,
dance steps, hand gestures

Macaques on the Arashiyama mountaintop in Japan customarily
together - the young learn to do it without any reward
other than perhaps the noise associated with it

Kohler: apes invented
all the time e.g.
marching single file around & around a post, trotting in the same rhythm
with emphasis on one stamping foot, wagging their head in same rhythm,
all acting in synchrony as if in a trance
55

In Zambia, scientists followed the
- one female was the first to stick a straw of grass into her ear, letting it
hang out while walking around and grooming others; over the years, other
chimps followed her example with several of them adopting the same new
"look"
of

some
wild
chimpanzee communities, in which two individuals hold hands above their
heads while grooming each other's armpits with their other hands

An infant male chimp may mimic the
of
the alpha male who always bangs a specific metal door to accentuate his
performance

Horner: spread brightly colored plastic chips around in an enclosure,
which the chimps could collect & carry to a container in exchange for
rewards; exposed to the sight of a top-ranking group member trained to
drop tokens into one container & a bottom-ranking one trained to use a
different container, the colony massively followed in the footsteps of the
member

Other species

Imitations in monkeys, dogs, corvids, parrots, and dolphins

Dogs vs. wolves: conspecific approach

Instead of following human instructions, both dogs and wolves saw a
member of their own species manipulate a lever to open the lid of a
box with hidden food
56

Next, they were allowed to try the same box themselves

The wolves
the dogs -
wolves may be poor at following human pointing, but when it comes
to picking up hints from their own kind, they beat dogs

rather than cognition - wolves watch one
another more closely as they rely on the pack for survival, whereas
dogs rely on us
o Other examples of

mimicry
Chimpanzee learned to sharpen pencils, use sandpaper, wash dishes, wipe
its bottoms


Parrots & mynah birds mimicking human speech

Seals can mimic human speech
Mechanisms of imitation & mimicry
o Rizzolatti et al: discovery in monkeys of
, which
have the property of firing either when the animals makes a particular action, or
when it observes the same action being made
o When one animal
a response, a
mirror neuron will be activated and excite the neurons controlling the same
response in the observer
o Imitation is a bodily process facilitated by
o These neurons, significantly (and ironically), were discovered not in humans but
in macaques
57
58
Memory & Planning
Memory

Spatial memory

Short-term memory

Long-term memory

Prospective memory (planning)
Spatial Memory

(Olton) – for rats
o Description

A central area with a number of arms running off it; at the end of each arm a
small piece of food is hidden in a food cup

Animal’s task is to run down each arm and
as efficiently as possible
59

Normally the food, once taken, is not replaced by the experimenters, so the
rat must
it has been down in
order not to waste time & energy returning to arms in which the cup has
already been emptied

Different numbers of arms can be attached to the central area, but
is the number commonly used in memory research

Animal can be detained in the central area by means of small doors operated
by remote control by the experimenters - this enable them to test
the rats can remember which arms they have been down
o What can be tested?

Capacity

Capacity of a rat’s STM can be readily tested in the radial arm maze by
until the rat starts to
make errors

In a standard sized radial maze of 8 arms, rats
to an arm they have already visited

Numerous control experiments have demonstrated that rats really do use
memory to
which arms they have visited
(e.g. they do not scent-mark arms they have already been)

Rats do
simply go around the maze clockwise or
anticlockwise – to a human observer their activity seems fairly random

If more than 8 arms are used, accuracy starts to
can still perform above chance level within 17 or 24 arms
60
; they

But as the number of arms increases, they start to simplify the task by
such as always turning to
the right when leaving an arm

Duration

How long animals’ memory in those STM tasks can last?

Rats working in the 8-arm radial maze were
from the
maze after making their 1st 4 choices

After a delay of 4 minutes to 24 hours, they were brought back to the
maze to finish their task – after a gap of up to 4 hours,
but after 8 or more hours,

Water maze (Morris)
o A
(1.3 m in diameter); water
made opaque by addition of milk powder; hidden beneath the surface of the
water is an escape platform; no food is used to motivate the rat, the desire is to
escape from the water-filled vat is motivation enough
o Rats are
swimmers & are highly
motivated to find the hidden platform, which saves them from having to swim any
longer
61
o The ingenuity of this maze is that it gets around the problem of
(each time a rat is put in the water it can be assumed to be equally motivated to
find the hidden platform: in a more orthodox, food-motivated maze an animal
may become less motivated as its hunger abates)
o The water maze also prevents all possibility of the rat finding its way by using
(e.g. by scent marking the parts
of the maze it has already explored)
o The maze’s
ensures that the rats must be
navigating solely on the basis of whatever is visible outside the maze (door,
windows in the lab) & their memory for where they have already swum in the
maze
o Problem: The rat's memory is tested
Short-Term Memory: Matsuzawa & Inoue on Chimpanzee

The Test
o Test 1:

This test included 3 5-y.o. chimps who were taught the order of Arabic
numerals
, and a dozen human volunteers.
62

Participants saw nine numbers displayed on a computer screen – when they
touched the first number, the other eight turned into
the test was to touch all these squares in the order of the numbers that used
to be there

Results:

The chimps, while
than the people, could do
this

One chimp, Ayumu, did the best.
o Test 2:

Subjects: Ayumu & 9 college students

Five numbers flashed on the screen
were replaced by white squares.

The challenge, again, was to touch these squares

63
before they

Results:

When the numbers were displayed for about 7/10th of a second, Ayumu
and the college students were both able to do this correctly about
of the time.

But when the numbers were displayed for just 4/10th or 2/10th of a
second, Ayumu was

.
The briefer of those times is too short to allow a look around the screen,
and in those tests Ayumu still scored about 80%, while the humans'
scores

to 40%.
That indicates Ayumu was better at taking in the
of numbers at a glance

Even with six months of training, three students
to the three young chimps

Explanation
o Human ancestors
much of this skill over evolutionary
time to make room in the brain for gaining language abilities
o The youth of Ayumu and his peers – the memory for images that is needed for
the tests resembles a skill found in
, but which dissipates with
age

The young chimps performed
than older chimps in the new
study

Ayumu's mom did
than the college students
64
o Implications

Ayumu has violated the dictum that, without exception, tests of intelligence
ought to confirm

Old thinking: humans command
whereas animals command very few abilities (all of which are adaptations
restricted to a single goal or activity)

Hauser: “
” – the gap between
human and animal cognition, even a chimpanzee, is greater than the
gap between a chimp and a beetle!

Evolution stops at the human head
 We descend from the apes in body but

- something major must have
happened after we split off from the apes; an abrupt change in the
last few million years
 This idea remains
in much of the social
sciences, philosophy, and the humanities
 It views our mind as so original that there is no point comparing it to
other minds except to confirm its
 However,
in
biology,
neuroscience,
medical
increasingly psychology,
is now the default assumption
65
science,
and

The case of bipedalism:

Previously, we made a big deal of human’s
while ignoring the many animals (from chickens to hopping kangaroos)
that move the same way

At some savanna sites, bonobos walk entire distances upright through
tall grasses, making confident strides like humans

Bipedalism is really
as it has been made
out to be

The distress Ayumu’s
caused
in the scientific community was of the same order as when, half a century
ago, DNA studies revealed that humans barely differ enough from bonobos
and chimpanzees to deserve their own genus
Long-Term Memory

Episodic memory
– the recall of what happened at
o
which place and at what time – the three W’s of events: what, when, and where
o Rats running radial arm maze & water maze
o Tinklepaugh on macaque monkey

A macaque watched either a lettuce leaf or a banana being placed under a
cup; as soon as the monkey was given access, she ran to the baited cup
66

If she found the food that she had seen being hidden, everything proceeded
smoothly; but if the experimenter had replaced the banana with lettuce, the
the reward; she’d frantically look
monkey only
around, inspecting the location over and over, while angrily shrieking at the
experimenter
o Martin-Ordas on chimpanzees and orangutans

She gave the apes a task that required them to find the
to fetch either a banana or frozen yogurt – apes had watched tools being
hidden in boxes, after which they needed to pick the right box to get a tool for
the task

, after the apes had gone through
scores of other events and tests, they all of a sudden encountered the same
person, presenting the same setup in the same rooms of the building

Those who had the previous experience knew right away what tool to use
and where to look for it – they
and
they solved the problem in a matter of seconds

Food-caching birds
o 2 key features of food-caching birds’ natural behavior

Birds rely on memory to

In the wild, some food-storing species (e.g. scrub jays) cache insects & other
perishable items in the wild – therefore useful for them to encode & recall
information about
has been cached
67
, as well as
o Clayton & Dickinson (1998)

“wax worms”
Jays were allowed to cache & recover
(wax worm larvae) &

peanuts
Jays show a
for caching, recovering, &
eating fresh wax worms when given both worms and peanuts

Worms
over time, so if they are left for a
period of 5 days or so, they become rotten and unpalatable

If jays remember when they cached as well as what they cached and where,
then they should
worms when they were cached just a
few hours ago

However, they should
the worms if the worms were cached
several days ago & have had time to rot

Findings – jays
from worms to
peanuts after long retention interval (124 hours), but
occurred after short retention interval (4 hours)
68
such reversal

Implication

The switch in preference from worms to peanuts after the long retention
interval required the birds to recognize a particular cache site in terms of
both its
& the
that had
elapsed between caching & recovery

This demonstrates recall of information about “
(peanuts & worms) were cached, “
stored, and “

” items
” each type of item was
” (short vs. long interval) the worms had been cached.
Pigeons (Vaughan & Green)
o Pigeons were trained in Skinner boxes to discriminate between
(“squiggles”)
o In one experiment, the slides had
drawn on them; in another, the slides were
taken
around the Cambridge, Massachusetts, area
o In each experiment, the pigeons were trained using a
procedure
to respond to one stimulus in each pair (the positive stimulus), & to refrain from
responding to the other stimulus – the choice of which stimulus in each pair was
positive and which was negative was completely arbitrary
o With sufficient training (nearly 1000 daily sessions for the squiggles, about 850
for the photographs), the birds reliably chose the
in each pair
69
o They were then given a break for
in the case of the
squiggles &
in the case of the photographs before
being retested
o On retesting they showed a
of which were the
positive and which were the negative stimuli
Prospective Memory (Planning) 
Behaviors
Naturalistic Observations: Bonobos at Lola ya Bonobo (jungle sanctuary in
Kinshasa)
o Lisala the Bonobo picked up an enormous 15-pound rock and lifted it onto her
back; then carried it on her shoulders while her baby clung to her lower back
o Lisala walked all told almost 10 minutes, only interrupted once when she put
down the rock and picked up some palm nuts that are extremely tough - before
she reached her destination, which was a large slab of hard rock
o She cleared if of debris with a few wipes of her hand, then put down her rock, her
infant, and the palm nuts
o She set out to crack the palm nuts, placing them on the large anvil while banging
them with her 15-pound rock as a hammer
o She spent about 15 minutes on this activity, then left her tool behind
o Purposeful & future-oriented behavior

It is hard to imagine that Lisala had gone through all this trouble
, which she must have had well before she picked up
the nuts
70

She probably knew where to find those, hence planned her route via this
location, to end up at a point that she knew had a hard enough surface for
successful cracking

This planning is different from, say squirrels collecting nuts in the fall & hiding
them for retrieval in winter - because it is based on
, not evolved tendency found in all members of the species

Experiments
o Scrub jays (Raby)

Experiment 1:

Raby offered jays an opportunity to store food in two compartments of
their cage that would be

during the night
The next morning they would get a chance to visit only one of the two
compartments
 One compartment had become associated with
,
since the birds had spent mornings there without breakfast
 The second compartment was known as the "breakfast room" because
it was

every morning
Given a chance in the evening to cache pine nuts, the birds put three
times as many nuts in the first room as in the second, thus
they might suffer there

Experiment 2

The birds had learned to associate both compartments with different
kinds of food
71

Once they knew what kind to expect, they tended to store a
in each compartment in the evening

This guaranteed a
breakfast if they
ended up in one of those compartments next morning

Conclusions - when scrub jays stash away food, they do not seem guided
by their present needs and desires but rather by the ones they
in the future
72
FOCUS TERMS
73
74
Psychology of Animals: Module 1 Focus Terms
1. Introduction

Romanes (comparative psychology); Pavlov & Skinner (behaviorism); Kohler
(cognitive psychology); Lorenz (ethology)

Ethology: most ethologists are zoologists (e.g. Conrad Lorenz), use laboratory &
field science

Fieldwork: systematic & scientific; continuous data collection; offer hints &
suggestions but rarely firm conclusions; not better or worse than experiments

Carefully-controlled experiments: close-up look at naturalistic behavior; animals
watched more fully; stimulate observations in the field; not better or worse than
fieldwork

Umvelt: cognitive ethology; Nagel – we can never fully understand other
species’s Umvelt; de Waal – we can try to understand other organism’s Umvelt

Anecdotes: most scientists who study the psychology of animals no longer
believe that anecdotes should be avoided at all costs.

Anthropomorphism - inevitable & involuntary

Anthropocentrism: seeing animals from humans' perspective; opposite of
biocentrism

Anthroharmonic approach: refuses to acknowledge the fundamental importance
of human beings in the world.

Human exceptionalism: e.g. “Human is the only animal that …. “
2. Behaviorism

Behaviorism: black box approach; associative learning; reductionistic

Classical conditioning: - reflex conditioning
o
Unconditioned stimulus (US), unconditioned response (UR), conditioned
stimulus (CS), conditioned response (CR):

US - biologically potent (e.g. the taste of food)

CS - usually a neutral stimulus (e.g. the sound of a tuning fork)
75


UR – e.g. Pavlov’s dog salivated to sight & smell of food
Operant conditioning
o Positive reinforcement, negative reinforcement (escape learning), positive
punishment, negative punishment
o Shaping (successive approximation)
3. Cognitive Learning & Social Learning

Problem with behaviorism: (1) only concerned with observable behaviors (2) fails
to take into account role of inherited & cognitive factors in learning

Insight
o Solutions gained by insights retained longer than those gained by trial-&error.
o Kohler: block stacking chimpanzee
o Goodall: aging wild chimp “Madame Bee” got her food delivered by her
daughter
o Premack’s chimpanzee: causal inference
o Heinrich’s raven
 Folk physics.
 Means-end relationships.
 May be explained by trial-&-error learning.

Latent (exploratory) learning:
 Tolman’s experiment: Group 3

Rats were not rewarded for trials 1 to 10, but rewarded for trials 11 to 22

Showed least number of errors in running the maze at the end of the 22day period
 Exploratory learning


"Indifferent stimuli"

Enables an animal to know its surrounding

To locate "commodity" (e.g. food)
Imprinting
 Conditioning, insight, instinct
76
 Phase sensitive learning
 Ontogenic insight: abilities to learn to do things during a short period
immediately after birth

Social Learning
o
Galef's experiments:

Social learning can overcome neophobia

Experiment 1:
 Observer rat's food preference still occurs even if demonstrator eats
flavored diet up to 4 hours before encounter with observer
 After single interaction with demonstrator, acquired preference
endures for about 12 hours.
 Once acquired, flavor preference hard to disrupt by subsequent
exposure to other types of food

Experiment 2:

Observer must interact with demonstrator.

Demonstrator can be unconscious.

More effective if observer interacts with front, not rear, end of
demonstrator.
o
o
Social learning in mate selection in guppies & Japanese quails

Female guppy observers prefer males seen with female companion.

Male Japanese quail observers prefer female seen in isolation.
Social learning of fear of snakes in monkey


Monkeys reared in laboratory show no fear of snakes
Copying behaviors
o Contagious behavior

Species-typical reaction – chorusing in rooster

Social facilitation – birds feeding in flock
o Imitation

Arbitrary responses
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
Examples: blue tits breaking through foil top of milk bottles to drink cream;
Japanese macaque monkeys washing sweet potatoes before eating;
Heyes & Dawson’s bidirectonal experiment

“True imitation” – intentionality, insight, comprehension; very rare

Human-raised young chimpanzees: as capable as imitating human adults
as young children; only mimicked necessary moves (Horner & Whiten)
unlike human children who mimicked everything

De
Waal
&
Whiten’s
conspecific
approach:
faithfully-transmitted
behaviors between Katie & Georgia
o Mimicry
 All about conformity; reward is secondary, not based on trial-&-error
 De Waal’s BIOL (Bonding- and Identification-based Observational
Learning) can explain:

why apes imitate their own kind far better than the average human.

why, among humans, apes imitate only those whom they feel close to.

why young chimps, especially females, learn so much from their
mothers

Demonstrated by Horner’s brightly colored plastic chips experiment
o Mirror neurons (Rizzolatti): neurological mechanism for imitation & mimicry
4. Memory & Planning

Spatial memory
o Radial maze:
 Invented by Olton, most common - 8 arms
 Problem of motivation esp. when rat not hungry
 Measures capacity & duration of STM
o Water maze: advantages
 Gets around problem of motivation
 Devoid of local cues (e.g. scent)

Short-term memory
o Matsuzawa & Inoue experiment
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 Young chimpanzees > College students
 Young chimpanzees > Old chimpanzees
 College students > Old chimpanzees
 Young children > College students
o Ayumu’s photographic memory stunned the scientific world – b/c it challenges
the notion of humaniqueness

Long-term memory (LTM)
o Episodic memory: demonstrated by Tinklepaugh’s experiments on macaque
monkeys & Martin-Ordas’s experiments on chimpanzees & orangutans
o Clayton & Dickinson experiment: jays’ LTM for 3 “wh” information
 What – perishable worm vs. non-perishable peanuts
 When – recently cached or cached long time ago
 Where

Prospective memory
o E.g. Lisala (bonobo in Kinshasa) carrying a heavy stone on her shoulders
o Raby’s scrub jay food-caching experiments: scrub jays can anticipate hunger
& prefer more varied meals
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