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Senior Freshman Animal
Behaviour Lecture 5 :
Cultural Transmission
Part 2: Imitation
How do animals do cultural
transmission?
We do it through imitation, but that’s actually
quite difficult.
Imagine a dog wanting to open a gate by
imitating owner’s actions
Dog needs to:
Know that the same actions would get the same result
Know the owner’s moves made the gate open
Picture himself in the owner’s position
Imagine the action from a different viewing angle
Understand the mechanics of the latch at least a bit.
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Four mechanisms mistaken for true imitation:
1)  Stimulus enhancement
2)  Social faciitation
3) Contagion (or observational learning)
4)  Emulation
1) Stimulus enhancement = learning by watching others
that a particular object or area is of interest, leading to
investigation of that object or area.
Early 70’s milk delivered to the door in bottles with foil caps.
Blue and great tits started to peck open the lids and drink the
cream on top of the milk.
This behaviour spread all across the UK surprisingly quickly.
It was assumed the birds watched
each other and learned the behaviour.
Held up as a good case of imitation
in the wild.
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Sherry and Galef 1984
investigated bottle opening
by blue tits.
3 groups of birds
1) Watch demonstrator bird open bottle
2) Just get bottle, opened previously by bird
3) Given closed bottle
Result: groups 1 and 2 equally fast to learn to
open the bottle, group 3 slower
Conclusion: Not imitation but Stimulus enhancement
= attention of observer drawn to an object or area by demonstrator.
Bird’s attention drawn to importance of the bottle; method to
open it trial and error.
Imo the Japanese Macaque,
studied by Kawamura 1959
Scientists put out rice and
potatoes for macaques, onto sand.
One day Imo took potato and put it
into the water and the sand fell off.
Later discovered could sort rice from sand in same way.
Both behaviours spread through the group quickly, so
researchers thought it good evidence of imitation.
But to study Imo’s behaviour the researchers put out more food
whenever the behaviour was seen.
Effectively rewarding the behaviour. Others learned through
stimulus enhancement (the sea is important) and
instrumental learning (carry food to the sea and you get more
food).
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Rewards are a common problem with almost all
anecdotal evidence.
Must avoid rewarding behaviour or you are simply training the
animal to do the behaviour, it’s not learning through cultural
transmission.
Reward may be subtle:
e.g. Baby dolphin in a zoo “imitated” sealion in same
enclosure in postures for sleep, swimming style and grooming.
BUT there are interpretations other than
imitation.
Also saw a man smoking, went to mum
and took a mouthful of milk then spat
out the milk to make same effect.
Interpretation of dolphin’s behaviour
1) It was indeed imitating what it saw, OR
2) It did random moves, some of which were seen by the
public as being imitation.
When they saw this, the public responded by clapping.
Dolphins like an audience, so he learned to do those
particular moves more often.
So possible that dolphin was trained through instrumental
learning to do particular movements which look to us like
imitation of others in its environment.
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2) Social Facilitation
= carrying out an already known
behaviour when others in the social
group are doing it too.
Advantageous because safer and
need less vigilance.
If it’s safe for someone to do an
activity, chances are it’s safe for
you to do it too.
Just timing of behaviours, not
acquisition of new ones.
3) Contagion or Observational Conditioning
= the observer “catches the mood” of the
demonstrator, and copies its general attitude to the
novel object.
e.g. learning to mob a predator by observing the response
of a demonstrator.
Humans are very
prone to this,
particularly when
young.
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Bandura et al. 1961 did Bobo Doll Experiment to
demonstrate contagion:
3 stages to expt
1) Preschool children did art project while:
For Group 1 – adult calm playing with quiet toy; ignored
nearby Bobo doll
For Group 2 – adult attacked Bobo doll with physical attack
while making violent comments
2) Each child alone had their aggression raised by being
shown some toys, then told “the other kids can play with
these but not you”.
3) Each child was taken into a new room where the bobo doll
was, and given choice of
Non-aggressive toys : Tea set, crayons, teddies, farm animals
Aggressive toys: Mallet and peg board, dart gun,
Results:
Children who had seen a calm
adult chose mainly nonaggressive toys and were nice
to the bobo doll.
Children who had seen an
aggressive adult chose
aggressive toys and attacked
the doll violently.
Conclusion: Children pick up
the attitude of adults by
contagion.
(Seeing aggression does NOT
give it a safe outlet, it breeds it!)
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4) Emulation= when an animal understands the goal of a
behavioural sequence, but gets to that goal by its own means,
discovered by trial and error, not by imitating the exact moves
of the demonstrator.
Need a task that can be done two ways. e.g. “mechanical
fruit”
Fruit opened by 1) twisting and pulling or
2) poking out the fastening rods.
If emulation, we expect no correlation
between demonstrator’s choice of
opening technique and observer’s.
If true imitation, expect good
correlation; observer should do what
demonstrator did.
Whiten 1996 used mechanical fruit to investigate emulation vs
imitation in chimps and human children aged 2, 3 and 4 years.
Demonstrator was adult, familiar to each group.
Results: Both chimps and children were influenced by the
demonstrator, so some degree of imitation present.
Chimps much more likely to use own method - emulation.
As kids got older, they imitated more and emulated less.
In fact older kids
copied all
movements, even
irrelevant ones.
Good imitation,
but intelligent?
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True imitation = when an animal learns a new behaviour
or sequence of movements which lead to a particular goal, by
watching a demonstrator carry out that sequence of actions.
Perhaps the chimps would have been
better imitators if they had a chimp
demonstrator.
But maybe imitation is not so adaptive
anyway!
An experiment with budgies which used the
same species as demonstrator does seem
to show true imitation.
Galef et al. 1986 Budgies’ Imitation
Experiment
Birds were given new feeding dishes with
lids, which could be opened with foot or
beak by a demonstrator budgie. Observer
birds had never seen these food dishes
before.
Treatments:
Group 1 saw demonstrator use foot.
Group 2 saw demonstrator use beak.
Results:
Almost all Group 1 observer birds used foot.
Almost all Group 2 observer birds used beak.
Conclusion : Budgies can learn a new action by observation,
i.e. budgies can show true imitation.
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Similar experiments with pigeons and quail show the same
type of results.
Quail study had the dish opened by stepping on a lever, or
pecking the lever.
BUT not strictly imitation as movements already known
(stepping and pecking) so this is classed as emulation only.
Moore 1992 studied Okichoro the
parrot
Attempted unanswerable demonstration of
imitation between species.
Put Okichoro in a room alone with a video
camera.
Every now and then a keeper went in and
said “ciao”, “nod” or “look at my tongue” and
made a suitable gesture.
Absolutely no reward was given to Okichoro for any
response he might make – guarding against accidental
instrumental learning.
The researchers then watched the video…
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Okichoro, all on his own, practicing the moves with the
appropriate words.
True imitation as shows new moves, unrewarded and too
complex to be hit upon by accident.
So even if chimps can’t do it, parrots can!
Teaching = when the demonstrator modifies its behaviour
only in the presence of appropriate naïve observers and at
some cost to itself.
The observer should also learn the information taught earlier
or faster than they would have if they had not been taught.
Caro and Hauser 1992 showed teaching in cheetah
Mothers sometimes don’t
choke prey but bring back for
cubs to learn how to kill.
Older cubs allowed to
overtake mother and make
the kill.
Cost: danger of losing prey
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Similarly, in nut opening chimp troops, mother may leave
implements and nuts near anvil for young to try with, and
learn to open the nuts.
Cost: Wasted nuts if juvenile doesn’t find them
Fewer nuts for mother.
Just watching mother doesn’t count as teaching, since mother
not altering her behaviour or incurring any costs.
Outcomes from Cultural Learning
1) New ideas spread very quickly.
2) Local dialects and traditions (culture!).
e.g. primate local traditions such as termite fishing and
nut cracking.
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Huffman 1970s-80s discovered
“Stoneplay” in Japanese macaques
Invented by a young female called
Glance in 1974
Collected a set of stones, stacked them up, knocked them down
Became very protective of them if anyone approached.
Huffman returned after 4 years and a few
more of the troop were doing it.
In 10 years it spread to almost half the
troop; 27 males and 14 females.
Cultural transmission was only ever to
those younger than Glance
Outcomes from Cultural
Learning: 3) Speciation!
Peter and Rosemary Grant 1994 studied
two Galapagos Finches: Medium Ground
finch and Large Cactus finch
The two species are very similar indeed in
appearance.
Large Cactus finch has a more repetitive
song than the medium ground finch.
Medium ground finch
If the female is given a choice of the two
songs, choose that of their own species 95%
of time.
So females can recognise species on song.
Large
Cactus
Finch
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Experiment to see what keeps
the species separate.
Large Cactus Finch
Taught cactus finch male to sing
ground finch song (by cultural
transmission) and offered them as
mates to both species of female.
Medium ground finch
Result: female cactus finches not interested in him, but
ground finches mated with him; i.e. mated according to song
not species.
Repeated with male ground finches – got same result.
Offspring produced by these matings healthy and fertile.
Conclusion: this culturally learned song in the males,
and culturally learned preference in the females is all that
separates the species.
Memes and
the future
Ideas spreading through cultural transmission
were called “memes” by Richard Dawkins
1976 in “The Selfish Gene”.
Cultural differences, passed on as memes,
can be very long lasting (books, CDs, Data
bases)
And spread incredibly quickly worldwide
(broadcasting, internet)
Is this a good thing??
Old ideas may not be relevant / beneficial now.
New ideas are untested for safety.
Inability to forget isn’t healthy but can we lose
memes stored in databases?
Or just lose all our time sifting through them!
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Recommended reading:
NB Same as last week!
Alcock J. Animal Behaviour 8th Edition Chapter 2 “Understanding the
proximate and ultimate causes of bird song”
Dugatkin L.A. 2004 Principles of Animal Behaviour
pp 15-18 cultural transmission of diet in rats
pp 145-175 Chapter on all aspects of cultural transmission
pp.198 – 205 – Cultural transmission of mate choice
Goodenough et al. 2001 Perspectives on Animal Behaviour
pp 95-98 Cultural transmission in a variety of contexts
Background reading:
New Scientist 24th March 2001, Vol 169 p26 “Culture Shock” –evidence for
culture in humpback whales.
New Scientist 3rd Aug 2002 Vol 175 p 41 “The golden meme” – the evolution
of ideas passed on by human culture.
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