The Evolution of Culture in Animals

The Evolution of Culture

in Animals

John Tyler Bonner

Original drawings by Margaret La Farge

PRINCETON UNIVERSITY PRESS

P R I N C E T O N , NEW JERSEY
CHAPTER 1

Philosophy and Less Grand Matters

This is a book on evolutionary biology. It stresses certain aspects

in a way that will perhaps provide some new insights into old
problems. I say “perhaps” because there are many sophisticated
evolutionary biologists who will point out that there are really no
new or revolutionary ideas; what I have to say is essentially what
they knew all along. That is most likely the case; yet they have not
put it all quite this way. The difference will be in the grouping and
arrangement of facts and ideas. It is often true that, as a subject
advances (especially when it advances rapidly), we do not always
appreciate immediately all the riches we have before us.
While it is my hope that I can convince the professional biologist
that there are new things to learn from old facts, I also want to
reach the anthropologist, the sociologist, and the well-read layman.
Therefore I will couch my arguments in as little jargon as possible.
This is not to mean that I subscribe to vagueness or that I court
imprecision. To the contrary, I want to make my exact meaning
immediately clear.
My purpose is to trace back into the origins of the human cultural
capacity back into early biological evolution. It will soon be
evident that I am not a catastrophist and do not believe that, like
the great good, culture suddenly appeared out of the blue at a
restricted moment in the early history of man. Rather, to continue
to borrow from the nineteenth-century geologists, I am a
uniformitarian and believe that all evolutionary changes were
relatively gradual and that we can find the seeds of human culture
in very early biological evolution.

A BRIEF ABSTRACT OF THE BOOK

There is a main point, a principal conclusion to this book, and I

should like to state it in the beginning so that it will be easier to
follow the thread of my argument. It is that even though culture
itself does not involve genetic inheritance or, therefore, Darwinian
4 · Philosophy and Less Grand Matters

evolution by natural selection, the ability of any animal to have

culture is a direct product of such an evolutionary mechanism.
Passing information by behavioral rather than genetic means has
made it possible in some cases to pass kinds of information that
either cannot be transmitted genetically at all or are less effectively
transmitted by genes. Natural selection operates on the genes and
only involves gene transmission; yet the power to transmit by
behavioral means is as a method adaptively advantageous.
Therefore, there has arisen a genetically determined behavioral
capacity to transmit information by signs, by language, by
imitation.
This kind of nongenetic transmission is mediated by the brain,
and so there has been a selection pressure for a larger and more
complex brain. The advantage of culture both in its present form
as seen in man, and in its more primitive forms as seen in other
animals, has continuously exerted pressure for brain expansion.
As a result there was first an increase in the ability to learn and
later in the ability to teach. A related trend can be seen in the
increase of the flexibility of responses ranging from single, rigid,
genetically determined responses to multiple responses that can
either be learned or even invented. These trends form a
progression ultimately leading to culture. I shall also examine the
early origins of these trends and follow the complete sequence of
biological events that are the antecedents of culture.
Culture is transmitted by behavioral rather than genetic means,
and it will be important to keep this distinction clear. The problem
is that any pattern of behavior could have both a genetic and a
learned or acquired component, which involves the old intractable
question of nature versus nurture. Many reasons can be given for
the intractability, but none of them makes the question any less
interesting, and human nature is such that we shall continue to try
and find ways of identifying what is inherited and what is learned.
However, for various reasons, involving both its difficulty of
analysis and the intellectually destructive political emotions it
generates, this is a subject I shall be careful to avoid in this book.
Instead I shall ask why culture exists at all, a question that can be
answered in the straightforward Darwinian manner just indicated.

REDUCTIONISM AND HOLISM

The convict that has arisen between biology and the social sciences
can, in large measure, be seen in terms of the conflict between
 Philosophy and Less Grand Matters · 5
reductionism and holism. By reductionism we mean a science (or
a hierarchical level) can be understood in terms of its component
parts from the level below; for instance, the symmetrical structure
of a crystal can be interpreted largely from the properties of its
constituent molecules. By holism we mean that there are emergent
properties arising at each hierarchical level and that these
properties cannot be understood in terms of those of a lower level.
The holist believes the living organism has properties that would
not be predictable on the basis of what we know of chemical
substances and the characteristics of human society cannot be
interpreted in terms of lower level biological properties. The old
adage of holism is that the whole is more than the sum of its parts.
Biology has undergone, in its most recent flowering, a period of
extraordinarily successful reductionism. It is hard to know exactly
when this began, but let me give a few milestones: the
interpretation of heredity in terms of unit characters by Gregor
Mendel; the demonstration by T. Boveri, E. B. Wilson, W. S.
Sutton, T. H. Morgan, and others that those characters resided in
the chromosomes of the nucleus; the discovery of the structure of
DNA by J. Watson and F. H. C. Crick; and finally the cracking of
the genetic code by M. W. Nirenberg and others who showed
which triplets of nucleotides in the DNA specified particular amino
acids in the proteins. By any measure these and other advances in
molecular biology have been staggering and at this very moment
the rapid progress continues unchecked.
If we turn to evolutionary biology, there has been a similar trend,
although it is less spectacular in its progress. Its origins of course
can be traced to Charles Darwin. The next step forward was the
rise of population genetics in the 1920s and 30s, especially the
work of R. A. Fisher, J. B. S. Haldane, and Sewall Wright. This
era of neo-Darwinism was truly reductionist because its concern
was for the rates of change of individual genes in a population over
time. In the 1940s and 50s the field was criticized because it was
oversimplified; it did not seem to reflect the real world, and
therefore unable to cope with what were perceived as the new and
more significant problems.
The next surge forward came in the 1960s when Robert
MacArthur and his colleagues and followers saw that one could
make simple theoretical models that applied not only to the more
complex aspects of evolution, but in particular to the morass
of problems in ecology. Their method of simplification and
6 · Philosophy and Less Grand Matters

approximation dramatically illuminated possible order out of

chaos. There was great resistance from the traditional evolutionists
and ecologists because it seemed that the very complexity of the
problems, which had been cherished as their most important
characteristic, was consciously ignored. MacArthur (1968)
countered these arguments with: “Think how physics would be
without its frictionless pulleys, conservative fields, ideal gases”
(1968:162). Theoretical ecology and evolution have already
proven, in a very short time span, to be enormously powerful as
analytic tools.
Nevertheless its proponents have not rejected holism, but they
believe that progress can most effectively be achieved by a balance
between holism and reductionism. Again to quote MacArthur
(1972): “Most scientists believe that the properties of the whole are
a consequence of the behavior and interaction of the components.
This is not to say that the way to understand the whole is always to
begin with the parts. We may reveal patterns in the whole that are
not evident at all in its parts. Species diversity, for example, is a
community property and is not a property of the individual
component species. It can be understood as a consequence of the
interaction of these species, but its patterns were discovered and
explained by people aware of communities; ecologists primarily
interested in the separate species have never made any progress in
unravelling community patterns” (1972:154, 155). In his own
research MacArthur showed that, using an overall holistic view as
his guide, he could generate simple hypotheses that were effective
in summing up the parts of a lower level of organization thereby
illuminating the properties of the higher level.
That there is a trend toward reductionism in modern population
biology is beyond doubt, and one example clearly illustrates this
fact. One of the important contributions of W. D. Hamilton (1964)
was the notion of inclusive fitness, the idea that fitness should
include the survival and reproduction of kin. This means, as was
explicit in the early ideas of the population geneticists, that the
genes are the object of selection. R. Dawkins (1976) has stated the
matter most elegantly in nonmathematical terms in his book The
Selfish Gene, whose title itself tells the main part of the story. He
talks of genes as being “replicators” and the organisms the genes
produce through development as being the “survival machines”
that are devices for keeping the replicators intact and functioning.
We shall discuss this whole matter in detail further on. Here the
 Philosophy and Less Grand Matters · 7
point I wish to make is that this is evolutionary reductionism in its
extreme form, and, as we shall see, these basic ideas have already
contributed to significant advances; they are the gateway to a new
understanding of evolution and the social organization of animals.
If one looks at the criticisms of sociobiology by anthropologists
and social scientists, they are almost entirely related to the idea that
a reductionist approach will not be useful in the social sciences.
The hierarchical study of human societies occupies a separate level
and must be considered in its own terms and not in terms of the
biological level lying below. In their view, human societies are too
complex, too special, too different from anything found in the
animal world to be interpreted in any meaningful way by
biological analysis. Their position is largely or entirely holistic.
For instance, their notion of culture is that it is an emergent
property unique to man. According to M. Sahlins (1976), culture
was developed in the hominid line about three million years ago.
It is a new condition that came into being as a result of the
complexity of the mind of man. To that extent the cultural
anthropologist would consider it biological, but once it came into
being, it took on a life of its own, and its new properties cannot be
understood in terms of the level below. It is, so to speak, self-
propelled and, like a soul, has become detached from its body.
More of this later; here I want only to stress that this is indeed a
holistic bias. (The very same argument can be made by those, such
as J. Jaynes [1977], who consider consciousness also to have arisen
suddenly in the early history of man, although consciousness,
according to Jaynes, arrived full-blown long after culture.)1
It has certainly been true for biology, as I previously illustrated,
that when a field is able to make advances by a reductionist
approach, the progress is most exciting and rapid. Furthermore it
is obvious that the more complex the field, the slower it achieves a
stage where it can make fast advances. This statement is one of
simple fact and applies not only to biology, but also to physics and
chemistry. And from this I would suggest it is not inconceivable
that the same process might occur in the social sciences at some
time in the future. If it does, clearly the lower level will be
biology. The sociobiologists have already claimed they have
1
The contrary view is admirably set forth in a book by D. R. Griffin (1976), who with
Darwin (1874) provides evidence for the idea that there might be a continuum between
what we call consciousness and various manifestations of behavior in animals.
8 · Philosophy and Less Grand Matters

found a bridge connecting the two levels, but this has been stoutly
resisted by many anthropologists and social scientists. Again, this
is obviously a matter to which we shall return.
One final word on holism. I do not mean to imply holism is bad
and reductionism is good, for they are both important. Often in a
particular field at a particular stage in its development it is
impossible to do anything other than examine the problems
holistically. Furthermore a holistic approach has, in many cases,
produced significant progress. It is probably true that it is a
necessary stage without which the reductionist progress could not
be made. Initially, it is the only way of describing the problems
and grouping the facts. Were this not done the chaos would be
complete. However, despite the strengths of a holistic approach,
one should not fear reductionism as an evil. When it comes to a
field, it should be greeted with caution, but also with pleasure. The
caution is needed because there is a degree of oversimplification
where the exceptions may accumulate to such an extent that clearly
they no longer prove the rule, but prove the need for a more refined
theoretical insight. The more traditional holism keeps the
perspective in the field, even when reductionism is rushing forward
at a dizzy pace.
It has always seemed strange to me that holism and reductionism
should elicit such strong passions among scholars. They are, after
all, only the philosophical methods characterizing different kinds
of scientific progress. The reductionists tend to be contemptuous
of all holists, for they feel they alone have the key to the universe.
Holists know they have a broad perspective, a large insight,
whereby they can see all the riches missed by the single-minded
reductionist. In principle it would appear so easy to be both at
once, but human nature is such that it enjoys taking positions on
philosophical or political dichotomies, ignoring totally the
possibility that some of these dichotomies are not genuine
antitheses of the either-or category, but are complementary. In
fact, I would go so far as to say that it is the holist who sees and
understands the dimensions of the problem and it is the
reductionist who in the long run will produce the most satisfying
type of explanation. The one cannot do without the other.

A DEFINITION OF CULTURE

There are probably few words that have as many definitions as

culture. I can remember when I was a student of Professor William
Weston at Harvard, there was a large room across the hall from his
 Philosophy and Less Grand Matters · 9
office where we made nutrient media to grow fungi and slime
molds. It was there that I learned the now lost art of how to make
potato agar from real potatoes. On the outside of the glass door to
this communal room was one word inscribed in large gold letters:
CULTURE.
At the other extreme there are those who use the word in a sense
I associate with Matthew Arnold and the Oxford English
Dictionary: culture is a refinement of tastes and artistic judgments;
it is the ultimate in the publication and ratification of the intellect.
Fortunately, definitions in science are arbitrary, and I shall define
the word in a sense somewhere in the middle of the great chasm
between the two uses of the word mentioned above. By culture I
mean the transfer of information by behavioral means, most
particularly by the process of teaching and learning. It is used in a
sense that contrasts with the transmission of genetic information
passed by the direct inheritance of genes from one generation to
the next. The information passed in a cultural fashion accumulates
in the form of knowledge and tradition, but the stress of this
definition is on the mode of transmission of the information, rather
than its result.
In this simple definition I have taken care not to limit it to man,
for, as so defined, there are among other animals, especially among
those that cooperate extensively such as primates. It would be easy
to alter the definition and say arbitrarily that it applied only to man,
and since any definition is fair game, there is nothing improper in
such a procedure. But I want to emphasize that this is not the
course I have taken.
There is a tendency to oppose the words biological and cultural,
but Marion Levy has pointed out to me why this is unfortunate.
Culture, as I have defined it, is a property achieved by living
organisms. Therefore in this sense it is as biological as any other
function of an organism, for instance, respiration or locomotion.
Since I am stressing the way information is transmitted, we could
call one cultural evolution and the other genetical evolution with
the understanding that they are both biological in the sense they
both involve living organisms.

ANTHROPOMORPHISMS

The existence of anthropomorphisms is a problem to which there is

no solution. Those interested in the similarities between man and
animal have no fear of anthropomorphisms, while those who see
10 · Philosophy and Less Grand Matters
man as special in some major way feel that our whole man-
oriented language is dangerous and misleading when applied to
animals. Here is a clear instance where human culture interferes
with our science.
Let us look at the prejudices of both sides of the argument. An
anthropologist might find the use of words such as slaves or castes
for ant colonies most undesirable. There are a number of reasons
he finds this usage unfortunate. For instance, it implies that the
most repugnant human morals are ascribed to the members of
some species of ant who are clearly too stupid to be immoral.
Much worse, it could imply that if ants have slavery, it is a natural
thing to do and therefore quite justified in a human society. These
arguments are not quite rational and can only be advanced under
extreme fervor of one sort or another. A more reasoned objection
would be that the motivations of ants and men might differ
radically, but by using the same words this distinction is lost.
A biologist, on the other hand, feels that the points made above
are too obvious to interfere with the dual use of the words. He
does not see any problem: in both ant and human slavery
individuals forcibly capture members of their own species or
related species and cause their captives to do work for the benefit
of the captors. It is unnecessary to drag in all the possible political,
psychological, or strictly human nuances; a very simple definition
of the word is sufficient. There is no need to be tyrannized by
words. If a biologist may not use the common words, he will be
forced to invent a whole set of jargon terms for nonhuman
societies, an unfortunate direction since there are too many jargon
words in any science as it is. I hope it will be sufficient if I make it
clear in the beginning that words either invented or frequently used
for human societies will also be used for animal societies with the
understanding that I am not implying anything human in their
meaning; they are to be considered simple descriptions of
conditions.
There nevertheless is a difficulty. It can be argued that no matter
how excellent and pure our stated intentions might be, the words
will unconsciously tend to make us interpret animal behavior in
human terms. But surely this danger exists no matter what terms
we use. It comes down to the very core of the problem of
objectivity: we see the world only through our own eyes, our own
minds. One might suppose it is easier to separate Newtonian
mechanics from our psyche than courtship and altruism in the
behavior of birds, but in fact they are both seen through our minds.
 Philosophy and Less Grand Matters · 11
If anything, in the behavior of birds it is possible to see the pitfalls
simply because they are more obvious. The difficulty of
attributing human motives (correctly or incorrectly) exists and will
continue to do so, no matter how cumbersome a vocabulary one
invents. When the reader finds words of common usage in the
pages that follow, he is urged to interpret them in a straightforward
way. Even if he subconsciously fails in this task, no great harm is
done for the question of what motivates other animals, as
compared to ourselves, is not the central subject of this book.
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