The Royal Society of Edinburgh

2013 James Black Prize Lecture

The Cultures of Apes and Other Animals
Professor Andrew Whiten FRSE FBA,
Professor of Evolutionary and Development Psychology
and Wardlaw Professor of Psychology,
University of St Andrews
Monday 19 May 2014
Report by Jeremy Watson

On presenting Professor Whiten with the Sir James Black Senior Prize, Professor Alan
Alexander, General Secretary of the RSE, said the award was for his outstanding
contribution to the understanding of the importance of a secondary inheritance system and
the role it plays in concert with genetic inheritance and individual learning.
Beginning his lecture, Professor Whiten asked, what do we mean by culture? For instance,
The Sunday Times has a section called Culture, which covers theatre, books, cinema and
the arts generally. That could be termed High Culture, Professor Whiten said. But is that a
broad enough definition? When scientists talk about culture, it means everything that is
learned through social learning and persists long enough to be called tradition. Such
traditions make up cultures, of which there are many different examples around the world,
involving what we wear, what we eat, how we eat and how we communicate through oral
and written language.
However, although human beings are so richly cultural, does that mean that chimpanzees do
not have cultures as defined above? For that matter, what about bees, crocodiles or rats?
Answering either yes or no is too simplistic, Professor Whiten said. To help us decide,
culture could be split into three main aspects.
Population-level patterning of traditions in time and space is the first. Human traditions vary
by region and also change over time, but so do animal examples such as birdsong, in which
dialects vary from area to area, even amongst species. The second aspect is cultural
content; for example, both humans and chimpanzees use tools and technologies and the
fact that they do so allows useful comparisons to be made. The third aspect is social
transmission, or how culture is passed on, through such processes as imitation or teaching.
So what is chimpanzee culture, Professor Whiten asked? Fifty years ago, next to nothing
was known about wild chimpanzee life, but subsequent research has discovered an
extraordinary amount. Between 10 and 20 years ago, it became apparent that chimpanzees
behave in different ways across Africa, as people do.
To build on this, Professor Whiten and his team have carried out a two-phase investigation,
based on an accumulation of 150 years across research sites. They first simply asked what
were common chimpanzee behaviour patterns in some areas but not in others, and this
yielded 65 candidate cultural variants.
In phase two, these variants were then filtered to identify 39 behaviours such as food
processing, tool use, social behaviour, grooming techniques and courtship that were
customary or habitual in some areas but absent in others, without any apparent ecological or
genetic explanation, and were thus inferred to be traditions. This was quite a revelation,
Professor Whiten said. Here we had a very rich picture of chimpanzee traditions that you
can call culture.
One example is using a natural hammer, made of wood or stone, to crack open a nut; a
behaviour widespread in West Africa but absent in areas elsewhere. Successful nut cracking

has to be learned, Professor Whiten explained. The shell cannot just be bashed with a stone
or the nut will be crushed as well, and the technique can take up to a decade to perfect.
So how is this behaviour passed on? To find out, researchers studied chimps on an island
sanctuary in Lake Victoria, in East Africa, where nut cracking does not occur in the wild. One
older chimp was trained to crack nuts while younger animals watched. These younger
animals learned to crack, whilst those who saw no model did not.
Two general conclusions have been drawn from the research, Professor Whiten said. The
first is that wild chimpanzees have multiple and diverse traditions a propensity that has
probably been there since humans and chimps, our closest animal relatives, shared a
common ancestry. The second conclusion is that each community is unique in its behaviour.
Professor Whiten said that in the same way that it is possible to tell where in the world
somebody lives because of their cultural repertoire, he can now tell which part of Africa a
chimpanzee comes from, by its behavioural profile.
But, he added, there were still nagging doubts about whether all the differences are due to
social learning. What scientists would ideally do is translocate chimps from one area to
another to see if their behaviour spread. However, as this would be unethical, researchers
have carried out studies in captive chimp centres.
In one, in each of two groups of chimpanzees a high-ranking female was shown a different
technique to extract food from an artificial foraging device. When they had learned their
technique, they were reunited with their group and the other chimps could watch. The
different behaviours duly spread within each of these groups, forming a local tradition.
This posed the question, can techniques spread from group to group? A second experiment
was conducted in which chimp groups seeded with different techniques could watch each
other as they acquired a new skill. The researchers found these chimpanzees also had the
capacity to spread behaviours from group to group.
This led Professor Whiten to ask what the process is by which culture is transmitted. Are
chimpanzees imitators copying what others do or emulators who see what effect the tool
can have then work out how to do it for themselves? They may appear to be imitating
aping others, Professor Whiten said, but are they, instead, working out how the world
works and then reinventing the necessary technique, which is quite a clever approach?
Further experimentation, however, in which chimps saw only the results of an action without
being shown how to do it, found the animals were unlikely to be purely emulators. They
could not work it out for themselves, Professor Whiten explained. By contrast they can watch
another of their species perform a task and then copy it.
Despite this, could there be an element of both imitation and emulation in chimpanzee
behaviour? Professor Whiten said he believed there could be a form of intelligent imitation
that sat between the two ends of the spectrum. An emulation/imitation switching experiment
was carried out with a group of chimpanzees and a group of young children.
The first step was showing both groups how to extract a reward food for the chimp, a
sticker for the children from a black box. Both groups were given a tool a stick and
went through a ritual that involved first poking the stick into the top of the black box and then
into the front to extract the reward. The children and the chimps watched this and copied
both steps. Two other groups then experienced the same, except that the box was
transparent, so now it could be seen that the action in the top hole was ineffective.
Surprisingly, the children continued to use the original technique, even though it seemed
clear that poking the stick in the top was irrelevant. However, the chimps stopped copying
the first of the two actions, realising it was irrelevant. This showed an element of intelligent
imitation, Professor Whiten said.

What the children did has now been called over-imitation, Professor Whiten said. As a
species, we are perhaps so deeply cultural that it pays to to operate a rule of thumb in
which we copy most adult behaviour faithfully.
A similar rule of thumb is conformity doing what a majority of others are doing simply
because everybody is doing it. Professor Whiten outlined an example from an experiment
with wild vervet monkeys, in which some groups were given nice-tasting pink food and
horrible-tasting blue food. Other groups were given the opposite. The monkeys learned to
eat the nice food and leave the horrible food.
Four months later, the coloured foods were brought back, but now with no nasty-tasting
additive, and the new crop of infants all copied their mothers in eating only what they
perceived to be good food, even though both blue and pink foods were now perfectly edible
an example of strong vertical cultural transmission.
However, male vervet monkeys can, and do, switch groups as they mature. When they did
this, despite what they had learned previously, they also switched their preferred colour of
foods an example of how the conformity rule of thumb works. When in Rome, do as the
Romans, particularly if you are a vervet monkey, said Professor Whiten. Thats conformity.
So what is the difference between animal and human culture, Professor Whiten asked?
Human culture is demonstrably cumulative and passed down through the generations, he
said, building on what has gone before. Humans have progressed from stone tools
thousands of years ago to 500 different types of hammer in Birmingham during the Industrial
Revolution. Thats what we can uniquely achieve, Professor Whiten added.
But although humans may be at the pinnacle of culture, can any animals also accumulate
culture? There is evidence that they can, Professor Whiten said. When chimpanzees hunt
for termites, the elaborate tool set they use to extract the insects from deep subterranean
nests cannot have been developed in one generation. There is the likelihood of cumulative
culture, but it is minimal compared to the human case.
In conclusion, Professor Whiten asked why all this is important for animals and humans.
First, it shows that as well as an inheritance system based on genetics, there is a secondary
system, based on social learning. It also shows that although the culture of humans may be
distinctive, quite rich foundations are shared with our closest relative the chimpanzee
and other big-brained animals. In fact, more basic foundations of social learning are
demonstrably widespread across the animal kingdom.



Q&A

Q: In the last 20 years of science, there has been an emphasis on the influence of genetics
on development. You have shown that there is cultural learning too. In terms of animals, will
this lead to a change of policy away from conservancy in zoos towards conservancy of
natural habitats?
A: One of the tragedies is that the chimpanzee species is being extinguished by our own.
We are not only losing chimpanzees but chimpanzee cultures. What will our research lead
to? There is a project run by the Smithsonian National Zoo in the USA with golden lion
tamarins. They tried to reintroduce them to the wild but it was disastrous, the tamarins didnt
know what to do. Now they are putting animals experienced in coping in the wild next to
these animals. In effect, they have to be apprenticed.


Q: We know animals make tools. But animals do it for a direct and immediate reward.
Humans make tools for future use. Have you shown that that notion may no longer be valid?
A: Yes. Chimpanzees, for example, bring tools from a distance to a foraging site. But there is
still a huge gulf between animals and humans. They make tools through destruction; we
create tools constructively.

Q: You have shown us learning in many ways. Do chimps actually teach?
A: There is a little evidence regarding behaviours such as nut cracking. Researchers have
described a young chimp making a mess of it. The mother came over and did it properly
then let him have the hammer. But this is very different from a community that actively
teaches. This was not intentional teaching. Predatory animals may teach more, however.
How else does a young cheetah go from suckling on its mother to catching and killing an
antelope? But slow-growing primates dont need to do this. They have years to learn what to
do.

Q: If humans are top of the cumulative cultural pyramid, is there evidence that this process is
accelerating? Is this acceleration also happening in animal culture?
A: It is accelerating. Culture changes very fast; just look at digital technology. But we have
big brains, three times the size of those of chimpanzees. We have been able to make our
cumulative cultural changes over many thousands of years. We may need to study
chimpanzees for thousands of years to see if anything like this process is happening.


VOTE OF THANKS
The Vote of Thanks was given by Professor Aubrey Manning OBE FRSE, who said he was
grateful to Professor Whiten for not taking an anthropocentric line. His superb lecture had
shown that we share many qualities with our fellow inhabitants of the planet.













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