Niko Tinbergen is one of the founders and grandmasters of ethology, and the papers published here are among its most important documents. They are a source book for students of animal behavior and will give the historian of ideas an insight into one of the most influential movements in modern science.

Anybody who thinks ethology consists of passively imbibing the information offered by nature about animals has much to learn about both science and ethology. The first stage in behavioral analysis, of course, is to observe and record what is going on. This will involve intent and prolonged observation until what an untrained observer might dismiss as a sequence of unrelated behavioral performances is seen to fall into well-defined and functionally connected sequences or “behavior structures.”

These structures do not declare themselves in any obvious way. Their identification depends upon an imaginative conjecture on the part of the observer, which further observation may or may not uphold. As in other branches of science, this is a creative process in which the imagination must take the initiative. A very important element in the ethological approach is the comparison of behavior structures as they occur in different but related animals, which in turn opens up the possibility of identifying homologies of behavior. The word “homology” is not in itself easy to define, but in ethology it may be exemplified by saying that the behavior associated with mother love and suckling is obviously homologous or genetically cognate in man and apes. Obviously this complex behavioral repertoire did not spring into being with the inception of the species Homo sapiens.

Because it was felt to be too descriptive and not sufficiently experimental in character, the ethological method was slow to gain ground, for in Tinbergen’s early days as a research worker “experimental” was the boss word much as “molecular” is today. Everything had to be experimental: embryology, pathology, physiology, and if not theology, then certainly the study of animal behavior. Plus c’est la même chose, for today, after the triumphs of DNA, anybody who studies “molecular” something—it might be molecular taxidermy—instead of humdrum old everyday something feels an inexplicable increase of stature. The idea that there is something essentially meritorious about experimentation has been carried over from Bacon’s original use of the term1 and the advocacy that went with it. It just so happens however that some of Tinbergen’s earliest work was indeed experimental.2 Much of the success of his work is due to his adoption of a judicious blend of observation and critical experimentation.

Except in those laboratories, all too common in England, where behavioral analysis centered upon applying bracing electric shocks to sea anemones in order to lay bare their behavioral repertoire, ethology was soon recognized as one of the really important developments of modern biology and became the subject of excited discussions in zoological departments. Many bright young zoologists—among them Desmond Morris—cherished an ambition to study under Tinbergen in Oxford. At the time when many of these papers were written Tinbergen was professor of zoology at Leyden, but by juggling with half-vacancies Professor A.C. Hardy made a place for him at Oxford, and to our great good fortune Tinbergen came to England.

When Tinbergen approached our leading private research foundation for funds to support his research, they consulted both a very eminent neurophysiologist and myself. Fortunately they accepted my advice and not his, so that ethology miraculously became solvent for several years. The eminent neurophysiologist said of ethology, “Why, that’s just bird watching, isn’t it?”

Among the classical investigations recounted by Tinbergen in this volume are his analysis of the purpose (survival value) and ethology of the habit found in many nesting birds of removing from the nest the empty shells left after the chicks have hatched. A complex situation is revealed in which there is a nice balance between the hazard involved in leaving the telltale shell where it is and in abandoning the nest even for a fraction of a minute, thus exposing the fledglings to danger.

It is interesting that in spite of the rectitude of Tinbergen’s writing he is human enough to lapse occasionally into the kind of language appropriate to describing human behavior, as when he says (p. 288) that when robber gulls are around, gull parents are “loath” to leave the nest. No fewer than three lengthy and detailed papers are devoted here to the matter of egg-shell removal by black-headed gulls. The effect of his patient and scrupulous attention to detail has been to increase the stature of ethology to that of a major subdivision of biology. Among the gifts that people like Tinbergen acquire is a kind of feral wisdom that is almost unintelligible to townspeople, exemplified here by the use he makes of the indentification and interpretation of animal tracks (a subject on which he has written extensively).


The paper on childhood autism written by Tinbergen and his wife is a study in the application of ethological methods to human infants. Notice here, for example, the identification of certain elements of “autistic” behavior in the behavioral repertoire of normal children, and the implied hypotheses (a) that autism may often be the consequence by no means of neglect but rather of too much intrusive attention, and (b) that autism is behavioral in origin and not, as some influential psychiatrists maintain, the result of a genetic defect or of organic brain damage. And the sense of Tinbergen’s enthusiasm, kindness, lack of pretension, and acute observation one gets from reading this paper seems to me entirely authentic.

Tinbergen’s own childhood was spent amid a happy and cheerful family and he showed an early predilection for a naturalist’s pursuits. One of his school reports, with almost uncanny lack of prescience, said that his powers of application were not such as to equip him for a career as a natural historian. In reality, the power of sustained and sharply focused attention that is so necessary for the successful pursuit of ethology is supremely well developed in Tinbergen. He spent much of the war in a hostage camp and was able to reflect, not for the first time, that wild animals are less to be feared than malevolent human beings.

From time to time people in public life are invited to sign manifestoes prepared by bodies whose titles sound like “World Federation for Peace and Friendship”—manifestoes which always seem to be couched in terms such as these:

The nations of the world must henceforward live together in amity and mutual understanding and must learn not to have recourse to armed conflict to settle their political differences.

The question that always rises to my lips on these occasions is, “What large and influential body of men holds a contrary opinion?” Who still contends that the nations of the world should live together in enmity and chronic discord, and should go to war at the drop of a hat to settle the most trivial political disputes? Thoughts of the same degree of impatience have occurred to me in reading Sir Macfarlane Burnet’s new book. Burnet is in favor of stabilizing the growth of populations and the human ecosystem generally, and so are many others. The trouble is that these are formidably difficult tasks which it may be impossible to accomplish within societies that respect the rights of individuals. It wouldn’t be all that easy even if we were able to boss everybody about without too much regard to human inclinations and liberty of choice.

In my Reith Lectures, The Future of Man (1960), I tried to explain why the task of attempting to stabilize the human population is such an especially difficult one. To suppose that it can be accomplished (not that Burnet does think so) by so simple a means as limiting every family to two children is the merest naïveté. I am rather inclined to think that for many years to come human populations will alternate between scares of overpopulation and scares (as in the 1930s) of dying out altogether through underfertility.

Burnet describes himself as an elderly medical scientist with a considerable record of achievement and recognition. He does not add, though there is no reason why I should not, that in his chosen field of research he is something of a genius, if by a genius we mean someone whose intuitions are (a) original and highly important, and (b) quite often right. Burnet has revolutionized our entire conception of the immunological response, showing how it is to be thought of in terms of the population dynamics of lymphoid cells. Thus Burnet has a deep understanding of immunology from the inside. Unfortunately this deep understanding does not extend to philosophic matters. In the course of some unduly lengthy apologia about the circumstances leading up to the writing of the present book he says:

The idea of Man as the dominant mammal of the Earth whose whole behavior tends to be dominated by his own desire for dominance gripped me. It seemed to explain almost everything and I applied it to everything.

And Burnet is as good as his word, for “everything” includes industrial conflict and communism, law and justice, the “pleasant sins,” craftsmanship, science and art, affection and cruelty, values and religion and ruling class behavior.

I think that a man more deeply experienced in philosophy or the history of ideas would have realized that no one formula or known combination of formulas of this kind ever explains everything. Although Burnet’s approach is infinitely more sophisticated, the attempted solution of ontological problems or philosophic problems generally by means of single formulas such as he advocates is hardly different in kind from the solution profferred by the enthusiast who solemnly assures us that “Everything is Vibration.” Matter and mind and all human interactions are to yield to the formula. This kind of thing may be all right for psychotherapy but it doesn’t measure up to real life.


Tinbergen does write on human behavior but hardly ever tries to establish that significant homologies between animal and human behavior exist. Tinbergen therefore lies comfortably outside the fields of inquiry that Alexander Alland tries to demolish in The Human Imperative. Alland would like to reinstate the dignity of man after his sufferings at the hands of latter day evolutionists and ethologists. He singles out such books as African Genesis, Territorial Imperative, On Aggression, The Naked Ape for special censure, and he would certainly have added Dominant Mammal if he had known of it. The authors of these books, he tells us, “have been singularly unable to offer insights into the reasons for behavioral differences between groups or to explain the complexity of human social patterns.” Although this criticism is not a cruelly unjust one, I think it misses the point about the true scope and competence of ethology and physiology when applied to human behavior.

Biology and ethology are especially qualified to identify and expound the history of the behavioral characteristics which human beings have in common. The study and interpretation of the differences between individuals is essentially the subject matter of psychology. This subdivision of labor makes ethology a less revealing and interesting subject for human beings than psychology. It is a rather dull truth, though true nevertheless, that human beings need food and take energetic steps to procure it, but what is interesting about this biological inheritance of man is why some people eat this and not that, here and not there, now and not then3—and these are matters for psychology.

Again, human ambition may represent some very deep-seated biological heritage. But what is interesting about ambition is what the psychologists may be able to tell us about it—why ambition takes different forms in different men or after different upbringings. It is simply no good seeking ethological explanations of why X is a scientist, Y a musician, and Z a layabout, supposing each of these states of life to represent the fulfillment of an ambition. Alland’s great message is that human beings are free from the tyranny of their biological inheritance by reason of the fact that the principal modality of evolution in human beings is not now genetical and is not dependent upon genetic variation but is, as Lotka would have said, and as Waddington, Huxley, and I have repeatedly said, “exosomatic,” “exogenetic,” or psychosocial.

This still leaves us with the problem of what significance we are to attach to attempts to establish significant homologies of behavior between human beings and lower animals. The ethological approach can be a genuine eye opener to those who had not realized that much of the human behavioral repertoire is of great evolutionary depth.

Tinbergen himself once wittily observed that when human beings see a mouse nursing and cherishing her young they are inclined to say, “How like human beings they are!” but pointed out that what they ought to say is, “Aren’t human beings exactly like mice?” Not even the most passionate advocate of the rights of man could seriously suppose that the enormously complex behavioral and physiological repertoire associated with childbirth and childbearing sprang into being for the first time with the evolution of man. Nor does the admission of the existence of this evolutionary background derogate in any way from the dignity of human beings, who deserve extra marks for being so much more patient of the noisiness, greed, and untidyness of their offspring than animals are likely to be.

It is simply no good striking pious attitudes about these matters. Certain human behavioral traits are as much a part of our biological heritage as our livers are and our endocrine glands. Darwin put it all much better than anyone since:

We are not here concerned with hopes or fears, only with the truth as far as our reason permits us to discover it; and I have given evidence to the best of my ability. We must however acknowledge, as it seems to me, that man with all his noble qualities, with sympathy which he feels for the most debased, with benevolence that extends not only to other men but to the humblest living creature, with his godlike intellect which has penetrated into the movements and constitution of the solar system—with all these exalted powers—man still bears in his bodily frame the indelible stamp of his lowly origin.4

In this passage we may substitute “behavioral repertoire” for “bodily frame” and we may also—thank God (I am absolutely with Alland here)—delete the word “indelible.” We don’t have to put up with hemolytic disease of the newborn, which is a deeply programmed consequence of our biological constitutions, and I’ll be damned if I see why we should have to bash our neighbors on the head or be bashed by them ourselves.

By far the most important characteristic of human beings is that we have and exercise moral judgment and are not at the mercy of our hormones and genes. The ethological approach, however, does in some cases give us some idea of the magnitude of the task we have to undertake in eradicating the undesirable elements of our biological heritage. Thus it is a truth which all advocates of family planning come to learn by experience that the human propensity toward love-making and reproduction which is so deeply engraved in the human biological program cannot be eradicated merely by prudential appeals to the social desirability of having fewer children or by calling attention gloomily to the likely plight of the human race in the year 2000 if reproduction is allowed to go unchecked.

This Issue

March 8, 1973