TINBERGEN, NIKOLAAS (NIKO)

(b. The Hague, Netherlands, 15 April 1907;

d. Oxford, United Kingdom, 21 December 1988), ethology, animal behavior, natural history, popularization of science, filming.

Tinbergen was, with Konrad Lorenz, one of the two main founders of the science of ethology, or biological study of animal behavior, and for this he received the Nobel Prize. Tinbergen approached animal behavior studies experimentally in the field, and advocated a rigorous separation of causal, functional, developmental, and evolutionary analysis—known as “Tinbergen’s four why's.” He began as a bird-watcher and field-worker with insects in the Netherlands, was interned as a hostage during the German occupation during World War II, and after the war moved to Oxford in England. He established a group of students, many of whom became well known in their field, and brought the science of animal behavior to a wide public. He was a brilliant communicator as well as a natural field biologist, and gained international recognition with his photography and several behavior films. In later life he focused his studies on childhood autism.

Early Life in Holland. Born and bred in the Netherlands, Niko moved to England in later years, but he had the good fortune to grow up in a country with an extremely rich natural history, and in a family with strong academic interests and background. His father Dirk Cornelis Tinbergen was a grammar school teacher of Dutch language in The Hague, and a respected scholar of medieval Dutch. He had a PhD and was the author of several books, including a widely used Dutch grammar. He was also keen on drawing as a hobby, a passion that Niko acquired from him. Dirk Cornelis was a hardworking, very organized person, intellectually stimulating, full of humor and joie de vivre, as well as a devoted father and family man, often taking his family on country walks and holiday trips.

Niko’s mother Jeanette was also a schoolteacher. She was more mathematical than her husband, but keen on literature, and spoke French, English, and German as well as Dutch. She was the heart of the family, a warm and impulsive person. They lived in a rather bourgeois street of terraced houses (Bentinckstraat, The Hague): simple, thrifty and rather austere, and the work ethic dominated.

There were five children. The eldest was Jan, who in academic achievement stood out above the others from a very early age. He became a physicist, later turned to economics, and ended up with the 1969 Nobel Prize in Economics, twenty honorary doctorates, a knighthood, and many other honors. He was very hardworking, even as a young boy, and whereas Niko larked about as a boy and just scraped through his school exams, Jan worked. In later life, Jan was the man of meticulous quantification, whereas Niko watched birds and had broad ideas. The two brothers were never particularly close.

Of the other siblings, Niko’s older sister Jakomien and younger brother Dik did not follow prominent academic careers; she became a teacher and head of languages in a grammar school, and Dik an engineer and later director of Public Energy in The Hague. For Niko the closest sibling was his brother Luuk, eight years his junior. Niko was in awe of him; he thought Luuk the most intelligent in the family, a keen naturalist and artist. When Niko was in his twenties he did many natural history projects on birds jointly with Luuk. Luuk later became a professor in Groningen University and a prominent ecologist, but he suffered from depression and took his own life at the age of thirty-nine.

Although the example of his parents and the interests of his siblings must have facilitated Tinbergen’s career as a natural historian, scientist, and writer, his development was not instigated by his family. Niko mapped out his own course, encouraged but not guided by his parents. As a boy, his life was about roaming around in the Dutch countryside, watching little creatures, walking, and messing about with nature, identifying birds, and bringing oiled birds home from the beach and cleaning them.

At his government grammar school in The Hague he did not do particularly well, except at sports (he even briefly played on the Dutch national hockey team). But in his spare time he became a fanatic teenage naturalist, encouraged by his biology master Dr. Abraham Schierbeek. What probably largely predetermined Tinbergen’s career as a biologist was his involvement in a youth organization, the Nederlandse Jeugdbond voor Natuurstudie (NJN; Dutch Youth League for Nature Study).

Spurred on by his NJN friends, his brother, his parents, and his schoolteacher, by the age of sixteen Tinbergen had produced his first publications, in popular natural history magazines, and he had made a significant start with wildlife photography, which in the 1920s was a new development. Yet he was somewhat suspicious of academia, and at the end of his school years he could not see a career in biology, a subject he saw dominated by morphology and taxonomy. He had a flair for languages, with an excellent command of Dutch and good working knowledge of German, French, and English, but he disliked any of these subjects in the disciplined, formalized context of school. He was full of doubts.

At the end of school in 1925, his parents persuaded him to take a “working holiday” at a biological field station on the Baltic Sea, the bird migration station (Vogelwarte) directed by Johannes Thienemann in the Kurische Nehrung, in East Prussia (now in Kaliningrad, part of Russia). He spent most of his time there with photography rather than science, yet when he returned he immediately enrolled for a five-year degree course in biology at the University of Leiden, close to The Hague.

Later, Tinbergen wrote: “I started my studies in Leiden at the tail end of a period of the most narrow-minded, purely ‘homology-hunting’ phase of comparative anatomy, taught by old professors” (Tinbergen, 1989, p. 438). For him, biology in the university consisted of lists of facts and dry comparisons, contemplated in endless lectures in stuffy rooms. But outside with his friends, he could study birds in their nests and insects on bright flowers, along beaches and drifting skies. He spent a minimum of time on course work, was absent as often as he could be, just making sure he would pass, while all the time doing exciting extracurricular projects in order to keep sane. The undergraduate study itself had little impact on him, but his activities away from university all the more.

His extracurricular activities during his student years brought him in contact with several people who had a long-lasting influence on him. They included Gerard Tijmstra, a maverick who at the time was a math teacher as well as an ornithologist, and who induced Tinbergen to start serious observations on gulls. There was Jan Verwey, who lectured in zoology at Leiden, and who was very much a field man and bird-watcher; he produced some of the first analyses of bird behavior (herons), drawing attention to their “ritualized movements” and “behavior out of context.” He later became director of the Dutch marine institute. Verwey and Tinbergen clicked, and spent many hours on bird observations together. There was A. F. J. (Frits) Portielje, a supervisor at the Amsterdam zoo, a keen observer of animals in captivity, also of gulls (he wrote a seminal paper on their behavior), and in Holland a widely known popularizer of things natural.

Tinbergen, as an undergraduate, watched and wrote about the behavior of herring gulls, terns, several raptors and owls, about migration and bird territories, and about shells and birds on the beaches, all in Dutch natural history magazines (Levende Natuur, Ardea, Amoeba, Meidoorn, Wandelaar, and others). In 1930, jointly with three friends, he published his first book Vogeleiland (Bird island), a natural history description of an area, De Beer near Rotterdam; he was the main author, but the authors’ names came in alphabetical order. In 1929, through the NJN, he met his later wife, Elisabeth (Lies) Rutten, the sister of one of his coauthors of Vogeleiland. In the last of his undergraduate years, 1930, Tinbergen did his first small, serious scientific study (though still largely descriptive), on the courtship behavior of common terns, published the following year in German—without any involvement of his university teachers.

Despite further doubts about an academic career, after graduating Tinbergen accepted a job as research assistant in the Zoology Department in Leiden, under Cornelis Jakob van der Klaauw. In this department Hildebrand Boschma was one of Tinbergen’s main contacts, specializing in taxonomy and physiology of invertebrates, not himself interested in zoological fieldwork, but accepting that such studies provided additional strength to the department. He encouraged Tinbergen and the two would maintain a regular contact and correspondence for decades after. Tinbergen had decided on a PhD project on a species of digger wasp, the bee wolf Philanthus triangulum, to be supervised by Boschma. He had been intrigued by these insects in a dune area in the center of the Netherlands, Hulshorst, where his parents had a holiday cottage, and he had previously done a small undergraduate project on them.

The bee wolf is one of the larger solitary wasps, yellow and black. In sandy dune country the female digs a tunnel about half a meter deep, with a few chambers at the end that she supplies with dead honeybees, several per chamber. She lays one egg in each chamber, and the larva feeds on the bees. When the female returns to the nest with a dead bee, she somehow finds the inconspicuous entrance to her nest, and Tinbergen investigated, among other things, what recognition criteria she used to find the right place. He designed elegant field experiments to address this, experiments that would become classics in later years. For instance, he surrounded the nest entrance with a 30-centimeter circle of pine cones before the bee left for a hunting trip, then moved the circle over a short distance after the bee had left, and observed the effect when the insect returned (confusion). He did this also using other objects, some flat, some tall and farther away from the nest, and he investigated the role of scent (absent). Similarly, he investigated the hunting behavior of the wasps, how they catch their honeybees (and only honeybees), and the important role of scent in recognizing honeybees.

Tinbergen’s PhD thesis, “Über die Orientierung des Bienenwolfes” (On the orientation of the bee wolf), was twenty-nine pages long, in German, and published in 1932 in Zeitschrift für Vergleichende Physiologie (Journal of comparative physiology). It was one of the shortest theses ever in this field, generally judged at the time to be a good paper, but nothing outstanding. It was one of the beginnings of field experimental studies of animal behavior, but years later Tinbergen was still amazed that he got away with it. The reason for its perfunctory quality was that Tinbergen had been selected as one of the participants in a one-year expedition to Greenland, and had to shorten his project; he became doctor of philosophy on 12 April 1932, got married to Lies Rutten on 14 April, and left for Greenland a few weeks later.

One of the remarkable aspects of Tinbergen’s PhD work on bee wolves was the difference with contemporary experimental methods, which in the laboratory sought to study animals under conditions that were all controlled as much as possible. In contrast, Tinbergen studied animals in conditions that were “natural,” and in which he attempted to change just one single variable. This was to characterize his later field-experimental work.

Greenland. In the context of the International Polar Year 1932–1933, the Tinbergens spent a year in Angmagssalik (now Tassiusaq), East Greenland. They stayed with Inuit, studied birds and the social life of sledge dogs, and collected a large number of Inuit utensils and objects d’art for the anthropological museum in The Hague. The Tinbergens’ many field notes on sledge dogs never saw the light of day, but the behavior studies of the snow bunting Plectrophonax nivalis and of the red-necked phalarope Phalaropus lobatus produced interesting papers after their return to the Netherlands. For both studies Tinbergen focused on the defense of a territory around the nest; he included discussions of the concepts of territorial behavior and gave detailed descriptions of behavior during the breeding cycle. The phalarope, a small wading bird, was especially interesting to him, being unusual among birds in that the female is brightly colored and defends the area, whereas the male is drab, and incubates and takes care of the chicks. Niko’s interest in territorial behavior was to be a main aspect of his future behavior studies, and this developed in Greenland.

More important than the immediate results of these Greenland field studies, however, was the effect of this interlude on Tinbergen himself. He gained confidence in his ability to carry out significant science under trying conditions, and in Holland he became an international scientist. Also, as an initially somewhat sentimental naturalist-conservationist, he became immersed in the Inuit culture of exploitation of animals and wildlife, and, staying most of his time there with the family of a shaman, he absorbed an Inuit view of animals as organisms just as plants are. He lost the notion of animals having sentiments; later this was to facilitate the concepts of animal behavior that he developed.

The Tinbergens returned from Greenland to Leiden in the fall of 1933, to his job as a research assistant to Van der Klaauw. He published a book in Dutch on his experiences, Eskimoland (1935), illustrated with many of his photographs, and a series of articles in popular natural history journals.

Development of Ethology in Leiden in 1930s. The subject of territorial defense by birds, which emerged in Tinbergen’s studies in Greenland, made him aware of a need to underpin his fieldwork with a theoretical base and a clear phrasing of questions. As he wrote in a Dutch article, instead of what birds are fighting for, we should be asking “what drives birds to fight, what do they fight, and what is the effect of their fighting?” He began to be involved with contemporary theories of animal psychology.

In the Netherlands at the time the leading student of animal behavior was Johan Bierens de Haan, who was to become an important force in Tinbergen’s development. The two men exchanged a frequent, voluminous, and often personal and friendly correspondence over more than twenty years. Tinbergen’s senior by twenty-four years, the animal psychologist Bierens de Haan was a lecturer at the University of Amsterdam and author of several books and papers. He saw animal instinct as having a clear purpose: It was innate and species-specific; it involved first of all an “awareness” followed by a “feeling,” followed by a “striving.” That, in turn, produced overt behavior, and according to him, a good animal observer would be able to recognize this chain of events by intuition. In response, Tinbergen urged that physiological phenomena be separated from psychological ones, and that science should be interested only in the former, as “subjective phenomena cannot be observed objectively in animals, it is idle either to claim or to deny their existence” (Tinbergen, 1951, p. 4).

The views of Bierens de Haan were broadly in line with those of the British psychologist William McDougall. Almost diametrically opposed to this were the views of the American John Broadus Watson, the man behind behaviorism, an approach that was equally an anathema to Tinbergen. Watson considered that all behavior was acquired, none was innate; every animal was a trained response machine and any behavior could be taught. Tinbergen was repulsed by the preoccupation of the behaviorists with white rats and monkeys in cages, pressing levers; later he said that behaviorism had given him a mental allergy to white rats from which he never fully recovered.

Another, different approach to animal behavior concentrated on its “directiveness.” It was that of the English biologist Edward Stuart Russell, who assumed that “the objective aim or ‘purpose’ of an activity controls its detailed course” (e.g., animals mate in order to produce offspring). Tinbergen objected that this precluded any physiological explanation of behavior (1951, pp. 3–4).

What struck Tinbergen about all the animal behavior theorists was that none of them knew animals in their natural environment, none of them was a field biologist. But, at least initially, he had little alternative to offer. Later, he would refer to the

haphazard, kaleidoscopic attempts at understanding animal behavior done by the future ethologists … made difficult rather than facilitated by the many early brands of psychology to which we turned for enlightenment, but which had disappointed us so bitterly. (Tinbergen, 1989, p. 440)

In the Zoology Department at Leiden in the 1930s, Tinbergen was charged by Van der Klaauw with a lecture course in comparative anatomy, and with the organization of student courses in animal behavior. His lectures and field courses were highly popular, and resulted in several outstanding research projects by Tinbergen with his undergraduate students, also independent PhD projects, and publications that were important in the development of ethology.

At the site of his previous PhD work in central Holland, the dunes of Hulshorst, one set of joint projects employing Tinbergen’s inductive approach (see below) was done, mostly with insects and all including a strong experimental element. This included extensions of his studies on Philanthus, including the PhD project of his student Gerard Baerends on the caterpillar-killing wasp Ammophila (Baerends was later to become a highly influential figure in Dutch zoology, and in animal behavior sciences), and research on the courtship behavior of a butterfly, the grayling Eumenis semele. In all these projects, sequences of behavior patterns were dissected into separate components, and causal factors as well as subsequent effects of each component were determined through simple, but ingenious, experiments in the field. The students loved it, and the resulting clear publications, in both scientific journals and more popular magazines, made a large impact.

In a similar approach in the laboratory, Tinbergen and his students carried out projects on the breeding behavior of small fish in a more or less natural context in tanks, especially on the three-spined stickleback Gasterosteus aculeatus. These were to become critical in his assessment and development of the theoretical views then being promoted by Konrad Lorenz, the German scientist who was to be the main force in Tinbergen’s further career in animal behavior.

The paths of Tinbergen and Lorenz crossed for the first time during a symposium on instinct held in Leiden, in 1936. Their characters were poles apart: Tinbergen the naturalist, gentle and self-deprecating, worrying, analyzing, and experimenting; Lorenz keeping animals at home, ebullient and brilliant, a philosophical mind, bubbling with ideas without testing or following them through. Nevertheless, the two hit it off immediately and became lifelong friends, despite many later controversies between them, and despite World War II that saw them passionately committed to opposite sides.

Lorenz had attracted Tinbergen’s attention with his 1935 German-language paper “Der Kumpan in des Umwelt des Vogels,” (The Companion in the Bird’s World) which described how instincts function in the social life of birds. He treated behavior patterns as if they were organs that can be compared between species, arguing that each behavior pattern was “released” by a combination of species-specific stimuli in the environment, like a key that fitted a lock. He also referred to behavior as occurring in different functional contexts, such as breeding or fighting, though he did not propose an internal systematic arrangement of behaviors. Such ideas were music to Tinbergen’s ears, and fitted in well with his experiments at that time on stickleback fish and insects. From his side, Lorenz found in Tinbergen’s experiments the scientific testing of his own as yet unsubstantiated ideas.

Their single joint publication, much quoted afterward, came after Niko and family stayed with Konrad in his home Altenberg in Austria in the spring of 1937, and the two scientists experimented with Lorenz’s greylag geese on the behavior of these birds when rolling a stray egg back into the nest. They described an “instinct action” with its specific releaser, and a separate direction component also with its own stimuli, jointly resulting in the egg being steered into the nest. Lorenz provided elaborate terminology and theory in the paper, Tinbergen the sections dealing with the experiments themselves.

Other classic experiments from that time in Altenberg involved cardboard models that were pulled overhead over young goslings and turkeys (later published in Tinbergen’s Study of Instinct [1951]). The models resembled a bird of prey (short neck, long tail) when pulled in one direction, and a duck when pulled in the other. The goslings responded as the investigators expected, reinforcing the idea of a very simple set of stimuli that directs behavior. Later students have found it difficult, however, to repeat the results.

Much of Tinbergen’s prewar approach to animal behavior is summarized in his “An Objectivistic Study of the Innate Behaviour of Animals,” published in 1942. In this he argued that animal behavior has internal and external causes, and can be arranged in hierarchical fashion. For instance, environmental factors would cause a fish or a bird to be in “reproductive drive,” then other stimuli would cause “subdrives” such as nest-building, courting, or fighting. Such stepwise organization schemes would apply to all behavior: There would be a hierarchy and such a hierarchy could be analyzed physiologically. Different drives (e.g., reproduction, aggression, predator-defense, feeding) would be mutually exclusive. All such behavior patterns would be inherited and innate, and he referred to them as “stereotyped movements” (later “fixed action patterns”), each set off by a release mechanism that was triggered by a specific stimulus.

Tinbergen’s “Objectivistic Study” paper contained many definitions and categorizations, with long discussions of terms such as instinct reaction chains, reflexes, vacuum activities, intention movements, and substitute activities. His ideas of hierarchical organization went much further than those of Lorenz. The paper also singled out the importance of studying not merely the causal background of behavior, but also its function, especially in communication, that is, behavior that is designed to carry information for other animals. It contrasted sharply with the subjectivist approach of McDougall and Bierens de Haan, where the animals’ feelings were paramount.

Tinbergen’s contribution firmly showed ethology to be an exact science. By expressing the principles more clearly than Lorenz had ever done, Tinbergen became the mouthpiece of the new discipline in the English-speaking world. In the meantime, in 1939 he had been appointed as “lector” in Leiden (comparable to a present-day readership).

World War II. In May 1940 the Netherlands were overrun by the German forces. In 1941 most of the teaching staff of Leiden University resigned in protest against the treatment of their Jewish colleagues, and in 1942 many of them, including Tinbergen as well as many other of the most prominent figures in Dutch society (professors, cabinet ministers), were taken hostage by the Germans. They were to be the subject for reprisals after actions by the Dutch underground; in the end, some twenty of them were shot. The hostages were interned in a former training college for priests in Sint-Michielsgestel in the south of Holland.

Tinbergen was to spend two years in the hostage camp, in reasonable comfort but with the threat hanging over him. The inmates organized lectures, plays, and concerts, and there was considerable intellectual activity as well as political discussion. Tinbergen was able to write. One product of that time was a text in Dutch, Inleiding tot de diersociologre (Introduction to Animal sociology), published as a 184-page paperback after the war, in 1946. It was visually appealing, with many of his drawings, but it was never much of a success, having been written in a rather schoolmasterly style. There was a kaleidoscope of social behavior, with several arguments about behavior organization as in his previous publications. As an interesting kind of throwback, in places Tinbergen insisted that animals behave for the good of the species. Given the context in which it was written, it was not surprising that the author was not in top form. Another result of his hostage camp efforts was a series of handwritten and richly illustrated booklets about animals, for his children at home. Two of these were later published in English: Kleew (1947), about gulls, and The Tale of John Stickle(1954), about the behavior of sticklebacks.

After Tinbergen was finally released from the hostage camp in September 1944, he would spend seven more months under the German occupation, living with his family close to his field study sites near Hulshorst. Life in Leiden was too difficult in that period, known as the “hunger winter,” with serious shortages.

Leiden after the War. After the war was over, rebuilding a research establishment took time, against a background of the problems of day-to-day living for a family with four children, in a shattered country. There was nothing to work with, not even notepaper, not even a bicycle to get around, and this in the face of a huge burden of lecturing to the flood of new students that followed the five years of war (for instance, Tinbergen had to lecture on animal morphology to some seven hundred medical students). He threw himself into the new, difficult life, and with missionary zeal even started a new journal for behavior studies, Behaviour, specifically to address his own young science.

In January 1947 he was appointed to a full professorship at Leiden, in experimental zoology. In his inaugural lecture, titled “Nature Is Stronger Than Nurture,” and subtitled “In Praise of Fieldwork,” Tinbergen outlined the aims and methods of ethology, the biological study of animal behavior. In the years following, he maintained a program of field research with his students just as in the 1930s, based in the same site Hulshorst, and also near the colonies of herring gulls on the Friesian island of Terschelling.

One set of (now well-known) field experiments there was aimed at the analysis of the pecking response gull chicks directed at their parents’ bills (thus eliciting food regurgitation): which colors, bill shapes, and movements could make the chicks peck. It was published in Behaviour in 1950; it could be criticized in its methods, but the innovative approach opened new avenues in biology.

In 1946 Tinbergen had made a three-month lecture tour through the United States and Canada, organized by Ernst Mayr. In a set of six lectures at Columbia University he set out the approach to ethology that was to become the framework for his magnum opus, The Study of Instinct, which he wrote during 1947 and 1948, but published only in 1951. It was the major product of his postwar years in Leiden. It gave an outline of the entire structure of animal behavior, its internal and external mechanisms, its development, and its biological function and evolution. It provided order in the perceived chaos of behaving animals, with simple explanations and ideas about how to watch and study, with no jargon, and with easy-to-read graphs and pleasant natural drawings. Later much of the structure of behavior proposed here was dismantled again, but The Study of Instinct served its purpose. It was ethology’s first real text, and it was critical for establishing the field’s identity.

It begins by explaining how ethology relates to physiology, psychology, other biological sciences, behaviorism, and vitalism, outlining the questions that Tinbergen thought important. The following chapters describe the hierarchical organization of behavior and the role of external “releasing” stimuli, and of internal factors such as hormones and the central nervous system, in causing behavior. The last three chapters discuss the development of behavior in an individual’s lifetime, including learning and conditioning, and discuss adaptation and evolution.

At the same time he was writing The Study of Instinct, Tinbergen published many scientific and popular papers. By any standards, he was highly successful in Leiden: professor at an unusually young age, many admiring students, renowned internationally, his major book about to burst onto the world scene, editor of the main international journal in his field, and able to travel as much as he liked. Yet in 1949 he abandoned his chair in Leiden for a job as demonstrator, well below the level of lecturer, in Oxford, England.

Reasons for his move were partly a missionary zeal to spread his ethology message in the English-speaking world, and partly that he had enough of the Netherlands, with its judgmental provincialism, its crowds, its choking rules and regulations, its celebration of financial gain, and the university where he had to spend too much time on administrative matters.

Oxford. Tinbergen was recruited to Oxford by the then-head of the Department of Zoology, Alister Hardy. He arrived in September 1949 with his family, Lies expecting their fifth child. It was a big upheaval, much greater than it would have been in the early twenty-first century, starting life in a quite different kind of society, in a foreign language, with children between four and fifteen going to local schools. But they were happy, and also, by the time Tinbergen started in the Department of Zoology his job had been upgraded to that of lecturer. He soon attracted a group of outstanding research students; there was an atmosphere of high spirits and tremendous enthusiasm.

In England Tinbergen again started behavioral fieldwork in gull colonies, initially in Norfolk, later on the Farne Islands and in Ravenglass, on the Irish Sea. With his students, he addressed by comparative and experimental methods the founding ideas of ethology as developed in the Study of Instinct. Among the products of this period were scores of popular articles, as well as theoretical papers (e.g., on “derived” activities, 1952) and several books. His postdoctoral student Esther Cullen’s paradigmatic contribution (1957) showed the varied aspects of bird behavior that had evolved in response to environmental requirements, in her classical paper on the adaptiveness of the behavior of a cliff-nesting gull, the kittiwake. Although the research was largely Cullen’s own initiative and work, it was always seen as a product of the Tinbergen group.

The Study of Instinct appeared in 1951, to excellent reviews all round; it established Tinbergen alongside of Lorenz as the leading scientist in this field. Although many of the underlying ideas had come from Lorenz, Tinbergen was perceived as responsible for ethology’s scientific foundation. The Study of Instinct was soon followed by Social Behaviour in Animals (1953), which made little impact, and by The Herring Gull’s World (1953), a detailed description of the behavior of the herring gull, and the book that Tinbergen himself was always most pleased with. There was the more popular Bird Life (1954), and Curious Naturalists (1958), in which he wrote about his fieldwork for a naturalist readership.

Niko Tinbergen attained great authority with his work in Oxford, and jointly with Konrad Lorenz in Germany, he was surrounded by the success of ethology. Then, out of the blue in 1953, came a potentially devastating criticism from Daniel Lehrman, a comparative psychologist based at Rutgers University in Newark. It was directed especially at Lorenz, but also at Tinbergen. Lehrman argued that there was no such thing as simply innate behavior. There was no evidence for a single causal background of similar behavior patterns in different species. There was no evidence for any underlying neuro-physiological mechanisms, which in any case were likely to be different between species. Lehrman saw the simple behavioral models of Lorenz and Tinbergen as a danger to understanding.

Tinbergen invited Lehrman to Oxford, they argued, and later they were to become good friends. Tinbergen agreed with many of Lehrman’s points, especially with the criticism that ethology made a clear distinction between innate and learned behavior (nature/nurture), and Tinbergen agreed that there had been much oversimplification. But he also made Lehrman see that he had rejected some useful aspects and methodologies of ethology. Lorenz was much more offended, and, unlike Tinbergen, saw nothing of value in Lehrman’s critique. He later interacted with Lehrman at ethological meetings and other conferences, but he never fully appreciated Lehrman’s objection that Lorenz’s sharp distinction between innate and learned behavior stood in the way of a better understanding of how behavior develops in the individual.

In subsequent years, Tinbergen no longer focused on any of the causal relationships underlying animal behavior; he published one more long paper on the comparative behavior of gull species (1959), but after that his interest concentrated on what he saw as the functional and evolutionary significance of behavior, on the effects of behavior patterns on the animals’ survival. These were topics that suited his talents best: the study of how behavior patterns contribute to the animals’ survival in the world, in their natural habitat.

At Oxford University itself, Tinbergen remained somewhat of an outsider. His lectures were popular, but the social side of university life did not appeal to him, and his interests were academically rather narrow. He was a fellow of the ancient Merton College for some years, a college typical of Oxford’s dreaming spires and full of ritual, but he resigned from that and instead moved to the more modern, down-to-earth Wolfson College. He had few friends in the university, and in general the Tinbergens rather kept to themselves.

But the research with his dynamic group of PhD students and collaborators continued vigorously in the late 1950s and the 1960s. He maintained intensive fieldwork, in which he actively participated himself. Especially in the large gull colonies of Ravenglass, on the Irish Sea, several innovative behavior studies were launched. One was a study of a simple behavior pattern of black-headed gulls, the removal of eggshells from their nests after eggs had hatched; this became a classic (1962). In elegant field experiments, Tinbergen analyzed the stimuli that induce the behavior (especially color and texture), and simultaneously (also experimentally) its biological function, that is, how this behavior contributes to the maintenance of camouflage of the nest.

In 1963 he published the paper that is considered to be his most significant contribution to ethology, “On Aims and Methods of Ethology,” dedicated to Konrad Lorenz for his sixtieth birthday; its message became known as “Tinbergen’s four whys.” Tinbergen elaborated the approach that he took in The Study of Instinct: As a biological science, ethology deals with observable phenomena—the starting point is inductive. It is concerned with four different problems: that of causation, of effect (function, or survival value), of evolution, and of ontogeny (nature-nurture). Tinbergen reviewed these four aspects in detail, and the urgent need for experiments, and added a plea for what he saw as a continuing duty of ethology: the detailed observation and description of behavior.

During the 1960s Tinbergen’s active involvement in fieldwork declined to almost a full stop. He still supervised students, still contributed several papers; there were further popular books (including a Time/Life book Animal Behavior, 1965) and many popular articles. However, his enthusiasm for carrying out research himself disappeared. It was replaced by his developing interest in moviemaking (which he had been doing on a small scale since the 1940s), which culminated in the television movie on the behavior of the herring gull, Signals for Survival. He spent several years filming it himself, in the huge bird colony on Walney Island, and edited it jointly with the experienced Hugh Falkus. It won the coveted Italia Prize for documentaries (1969).

Last Projects. Internationally, numerous distinctions came his way in the 1960s, including Fellowship of the Royal Society in 1962, at the age of fifty-five, honorary memberships of many societies, and invitations to lecture from all over the world. At Oxford his status was recognized rather belatedly, with a full professorship in 1966. He still had many students; he was ably assisted in their supervision (since the mid-1950s) by Michael Cullen, who, with a wide-ranging but severely critical and quantitative mind, was a perfect complement to Tinbergen. In Oxford Tinbergen was one of the instigators of the new undergraduate course in human sciences. He became closely involved in setting up and maintaining the Serengeti Research Institute in Tanzania, which he visited annually. But coincident with his declining interest in field studies, he struggled with health problems, especially with profound depressions that incapacitated him.

The main change in Tinbergen’s interest was toward the application of ethology in studying the behavior of people, and toward human problems. Having suffered lifelong feelings of guilt about his lack of interest in the suffering of people, he followed Konrad Lorenz and his former student Desmond Morris in the use of knowledge acquired in the study of animals to understand ills of humanity such as aggression and warfare. His inaugural lecture in Oxford (1968), titled “On War and Peace in Animals and Man,” was published in Science and created much discussion about whether comparisons of human and animal behavior were permissible. Tinbergen compared animal group territories with those of people and pointed out the malfunction of our “innate” appeasement gestures when long-range weapons were being used. He urged scientists not blithely to apply animal results to people (and he criticized Lorenz for this), but merely to use the methodology of ethology in the human context.

Tinbergen’s wife Lies took an interest in the behavior of children, and she and Niko started an observational study on the unusual behavior of autistic children, which was to be his last project. It culminated in their 1983 book “Autistic” Children: New Hope for a Cure. The Tinbergens’ research, papers, and lectures on childhood autism were controversial, as they drew profound conclusions on an emotional subject, with only anecdotal evidence as support. Using an ethological analysis, studying approach of and avoidance by children, the researchers concluded that defective parental behavior is the main cause of autism. Criticism was to be expected, and it was especially severe because of Niko Tinbergen’s international standing as a behavioral scientist.

In 1973 Tinbergen, jointly with Lorenz and Karl von Frisch, was presented with the Nobel Prize for Physiology or Medicine “for their discoveries concerning organization and elicitation of individual and social behaviour patterns.” A Nobel Prize for such a wide theme is unusual; more often, it is awarded for a single discovery. Von Frisch had, indeed, made such a single discovery, in the communication system of honeybees, but Lorenz and Tinbergen were awarded for their new approach. In his acceptance lecture, Tinbergen concentrated almost entirely on his autism studies, and on what he saw as an evaluation of the Alexander technique, an alternative and nonscientific technique to improve human body posture and movement. It drew much criticism.

In his retirement Tinbergen withdrew entirely from academia and science, and from moviemaking. He was close to his family, he corresponded warmly with many people but saw few friends, and was often severely depressed. He died of a stroke at the age of 81, on 21 December 1988.

Evaluation. The contribution of the bird-watcher who received the Nobel Prize was that of an innovator. He suggested the questions one should ask of the behavior of animals, both in the field methods used to study them, and in the experiments that change just one or two factors in the animals’ environment rather than take them into totally controlled captivity. Tinbergen’s contributions were all the more effective because he was a talented communicator, in many different ways.

Among students of animal behavior, Tinbergen is known for his “four whys”: the why of causation, of ontogeny, of survival value, and of evolution; these were the questions he addressed by experiment and comparison. Where Lorenz had a wealth of ideas, Tinbergen analyzed and experimented, and sorted wheat from chaff. Tinbergen’s first model of the hierarchical organization of behavior has been overtaken by others, but it was Tinbergen himself who initiated this process. From Lorenz’s first vague suggestions, it was Tinbergen who articulated the system of a hierarchy of behavior patterns. In general, the ideas of both Lorenz and Tinbergen about causation of behavior have largely been discarded, but studies such as Tinbergen’s first ventures into problems of survival value, of biological function, have developed and amplified hugely into what is now called behavioral ecology, whereas Lorenz, not being a field naturalist himself, had little to offer. Tinbergen’s first, simple experiments caused others to formulate questions of optimal performance, and the concept of optimality has been a major foundation of behavioral ecology. In his study of the evolution of behavior of gulls, Tinbergen had a major stroke of luck in the work of his postdoctoral student Esther Cullen, by putting her to work on the kittiwake, which demonstrated so beautifully how adaptation to a particular niche had repercussions for a whole range of species-specific behaviors.

Tinbergen’s experimental methodology was greeted enthusiastically at the time, but on close inspection many of his studies had failings that would not have passed a present-day reviewer, and his lack of quantification was criticized later, even by his own students (although partly this was the state of the science at the time). Some of the celebrated simplicity of the experiments caused flaws, among others because in the absence of blind tests there often was a subjective influence of the observer. But Tinbergen encouraged such critical rejection; his arguments made sense, and what mattered most were the ideas that he presented. It was Tinbergen’s rational questioning approach to the behavior of animals in their natural environment for which he will be remembered.

Publications and Impact. Tinbergen published sixteen books, several translated into many languages, and some 360 scientific and popular papers. Of these, The Study ofInstinct was the best known, and continues to be widely quoted in the early twenty-first century. About two-thirds of his papers were popular articles, mostly in Dutch, about one-third in English. His single most important scientific paper, “On Aims and Methods of Ethology” (1963), is quoted even more often than The Study of Instinct. His most important movie, Signals for Survival, had a strong science content as well as beautiful imagery.

Apart from the Nobel Prize for Physiology or Medicine (1973), Tinbergen received many other rewards. He had chairs in Leiden and in Oxford, numerous visiting professorships in universities in many countries, and honorary doctorates in Edinburgh and Leicester. He was a Fellow of the Royal Society, a fellow first of Merton, then of Wolfson College in Oxford, a foreign member of the U.S. National Academy of Sciences and of the Royal Netherlands Academy of Sciences and Arts, a member of the German Academy of Natural Sciences, and an honorary member of the German Ornithological Society, an honorary fellow of the American Academy of Arts and Sciences and of the Royal College of Psychiatry. He also received distinguished awards from many other societies, especially the Swammerdam Medal of the Dutch Academy of Sciences and Arts and the Godman Salvin Medal of the British Ornithological Union, as well as others. He supervised some forty PhD students, several of whom became highly influential (Gerard Baerends, Desmond Morris, John Krebs, and Richard Dawkins, among others).

BIBLIOGRAPHY

Hans Kruuk's Niko’s Nature, cited below, includes a complete bibliography.

WORKS BY TINBERGEN

With G. van Beusekom, F. P. J. Kooymans, and M. G. Rutten. Het Vogeleiland [Bird island]. Laren, Netherlands: A. G. Schoonderbeek, 1930.

“Zur Paarungsbiologie der Flussseeschwalbe (Sterna h. hirundo L.)” [On the biology of reproduction in the common tern]. Ardea 20 (1931): 1–18.

“Über die Orientierung des Bienenwolfes (Philanthus triangulum Fabr)” [On the orientation of the bee wolf]. Zeitschrift für vergleichende Physiologie 16 (1932): 305–334.

Eskimoland. Rotterdam: D. van Sijn & Zonen, 1935.

“Field Observations of East Greenland Birds. I. The Behaviour of the Red-Necked Phalarope (Phalaropus lobatus L.) in Spring.” Ardea 24 (1935): 1–42.

“The Function of Sexual Fighting in Birds, and the Problem of the Origin of ‘Territory.’” Bird Banding 7 (1936): 1–8.

With Konrad Lorenz. “Taxis und Instinkthandlung in der Eirollbewegung der Graugans, I” [Taxis and instinctive movement in the egg-rolling of the greylag goose]. Zeitschrift für Tierpsychologie 2 (1938): 1–29.

The Behavior of the Snow Bunting in Spring. Transactions of the Linnaean Society of New York 5. New York, 1939.

“Die Übersprungbewegung” [Displacement activities]. Zeitschrift für Tierpsychologie 4 (1940): 1–40.

“Ethologische Beobachtungen am Samtfalter, Satyrus semele L.” [Ethological observations on the grayling butterfly]. Journal für Ornithologie 89 (1941): 132–144.

“An Objectivistic Study of the Innate Behaviour of Animals.” Bibliotheca Biotheoretica 1 (1942): 39–98.

Inleiding tot de diersociologie [Introduction to animal sociology]. Gorinchem, Netherlands: Noorduijn, 1946.

Kleew: The Story of a Gull. New York: Oxford University Press, 1947.

De Natuur is sterker dan de leer, of de lof van het veldwerk [Nature is stronger than nurture: In praise of fieldwork]. Leiden: Luctor et Emergo, 1947. Leiden University inaugural lecture, 25 April 1947.

“The Hierarchical Organisation of Nervous Mechanisms Underlying Instinctive Behaviour.” Symposia of the Society for Experimental Biology 4 (1950): 305–312.

With A. C. Perdeck. “On the Stimulus Situation Releasing the Begging Response in the Newly-Hatched Herring Gull Chick (Larus argentatus argentatus Pont).” Behaviour 3 (1950): 1–39.

The Study of Instinct. Oxford: Clarendon Press, 1951.

“The Curious Behavior of the Stickleback.” Scientific American 193 (December 1952): 22–26.

“‘Derived’ Activities: Their Causation, Biological Significance, Origin and Emancipation during Evolution.” Quarterly Review of Biology 27 (1952): 1–32.

“On the Significance of Territory in the Herring Gull.” Ibis 94 (1952): 158–159.

“A Note on the Origin and Evolution of Threat Display.” Ibis 94 (1952): 160–162.

The Herring Gull’s World. London: Collins, 1953.

Social Behaviour in Animals. London: Methuen, 1953.

Bird Life. London: Oxford University Press, 1954.

The Tale of John Stickle. London: Methuen, 1954.

“On the Functions of Territory in Gulls.” Ibis 98 (1956): 401–411.

Curious Naturalists. London: Country Life, 1958.

“Comparative Studies of the Behaviour of Gulls (Laridae): A Progress Report.” Behaviour 15 (1959): 1–70.

With G. J. Broekhuysen, F. Feekes, J. C. W. Houghton, et al. “Egg Shell Removal by the Black-Headed Gull, Larus ridibundus L.: A Behaviour Component of Camouflage.” Behaviour 19 (1962): 74–117.

“On Aims and Methods of Ethology.” Zeitschrift für Tierpsychologie 20 (1963): 410–433. Facsimile reprinted in Animal Biology 55, no. 4 (2005): 297–321.

Animal Behavior. Life Nature Library. New York: Time Incorporated, 1965.

With Eric A. Ennion. Tracks. Oxford: Clarendon Press, 1967.

“On War and Peace in Animals and Man.” Science 160 (1968): 1411–1418.

With Hugh Falkus and Eric A. Ennion. Signals for Survival. Oxford: Clarendon Press, 1970. Book based on the 1969 film of the same name. VHS version: McGraw-Hill, 1970.

The Animal in Its World: Explorations of an Ethologist, 1932–1972, vol. 1, Field Studies. London: George Allen & Unwin, 1972. A compilation of ten of Tinbergen’s scientific papers on experimental fieldwork, including studies on bee wolf, grayling butterfly, black-headed gull, and fox.

The Animal in Its World: Explorations of an Ethologist, 1932–1972, vol. 2. Laboratory Experiments and General Papers. London: George Allen & Unwin, 1972. A compilation of eight of Tinbergen’s scientific papers, on laboratory studies (of thrushes and sticklebacks) and ethological theory.

“The Croonian Lecture, 1972: Functional Ethology and the Human Sciences.” Proceedings of the Royal Society of London, Series B, Biological Sciences 182 (1972): 385–410.

With Elisabeth A. Tinbergen. Early Childhood Autism: An Ethological Approach. Advances in Ethology, supplements to Journal of Comparative Ethology (Zeitschrift für Tierpsychologie) 10. Berlin: P. Parey, 1972.

“Ethology and Stress Diseases.” In Les Prix Nobel en 1973. Stockholm: Norstedt, 1974, and Science 185 (1974): 20–27. Nobel Lecture. Available from http://nobelprize.org/nobel_prizes/medicine/laureates/1973/tinbergen-lecture.html

With Elisabeth A. Tinbergen. “Autistic” Children: New Hope for a Cure. London: Allen and Unwin, 1983.

“Watching and Wondering.” In Studying Animal Behavior: Autobiographies of the Founders, edited by Donald A. Dewsbury. Chicago: University of Chicago Press, 1989.

OTHER SOURCES

Burkhardt, Richard W. Patterns of Behavior. Chicago: University of Chicago Press, 2005.

Cullen, Esther. “Adaptations in the Kittiwake to Cliff-Nesting.” Ibis 99 (1957): 275–302.

Kruuk, Hans. Niko’s Nature. Oxford: Oxford University Press, 2003.

Lorenz, Konrad. “Der Kumpan in der Umwelt des Vogels: Der Artgenosse als auslösendes Moment sozialer Verhaltungsweisen” [The companion in the bird’s world: Fellow members of the species as releasers of social behavior]. Journal für Ornithologie 83 (1935): 137–215, 289–413.

Röell, D. René. The World of Instinct: Niko Tinbergen and the Rise of Ethology in the Netherlands (1920–1950). Assen, Netherlands: Van Gorcum, 2000.

Hans Kruuk

Nikolaas Tinbergen

Nikolaas Tinbergen (1907-1988) is known for his studies of stimulus-response processes in wasps, fishes, and gulls. He shared the Nobel Prize in medicine in 1973 for work on the organization and causes of social and individual patterns of behavior in animals.

Nikolaas Tinbergen, a zoologist, animal psychologist, and pioneer in the field of ethology (the study of the behavior of animals in relation to their habitat), is most well known for his studies of stimulus-response processes in wasps, fishes, and gulls. He shared the 1973 Nobel Prize in physiology or medicine with Austrian zoologists Karl von Frisch and Konrad Lorenz for his work on the organization and causes of social and individual patterns of behavior in animals.

The third of five children, Tinbergen was born April 15, 1907, in The Hague, Netherlands, to Dirk Cornelius Tinbergen, a school teacher, and Jeanette van Eek. His older brother Jan studied physics but later turned to economics, winning the first Nobel Prize awarded in that subject in 1969. The Tinbergens lived near the seashore, where Tinbergen often went to collect shells, camp, and watch animals, many of which he would later formally research.

After high school, Tinbergen worked at the Vogelwarte Rossitten bird observatory and later began studying biology at the State University of Leiden, Netherlands. For his dissertation, Tinbergen studied bee-killer wasps and was able to experimentally demonstrate that the wasps use landmarks to orientate themselves. Tinbergen first established the traditional routes of the wasps near their burrows, then altered the landscape to see how the wasps' behavior would be affected. Tinbergen was awarded his Ph.D. in 1932.

Tinbergen married Elisabeth Rutten in 1932 (they had five children together). Soon afterward, the Tinbergens embarked on an expedition to Greenland, where Tinbergen studied the role of evolution in the behavior of snow buntings, phalaropes, and Eskimo sled dogs. When he returned to the Netherlands in 1933, he became an instructor at the State University, where he organized an undergraduate course on animal behavior. Tinbergen's work had been recognized in the field of biology but it was not until after he met Lorenz—the acknowledged father of ethology—that his work began to form a directed body of research. Tinbergen took his family to Lorenz's home in Austria for a summer so the two men could work together. Although they published only one paper together, their collaboration lasted a number of years.

During 1936, Tinbergen and Lorenz began constructing a theoretical framework for the study of ethology, which was then a fledgling field. They hypothesized that instinct, as opposed to simply being a response to environmental factors, arises from an animal's impulses. This idea is expressed by the concept of a fixed-action pattern, a repeated, distinct set of movements or behaviors, which Tinbergen and Lorenz believed all animals have. A fixed-action pattern is triggered by something in the animal's environment. In some species of gull, for instance, hungry chicks will peck at a decoy with a red spot on its bill, a characteristic of the gull. Tinbergen showed that in some animals learned behavior is critical for survival. The oystercatcher, for instance, has to learn which objects to peck at for food by watching its mother. Tinbergen and Lorenz also demonstrated that animal behavior can be the result of contradictory impulses and that a conflict between drives may produce a reaction that is strangely unsuited to the stimuli. For example, an animal defending its territory against a formidable attacker, caught between the impulse to fight or flee, may begin grooming or eating.

Regarding his collaboration with Lorenz, Tinbergen is quoted in Nobel Prize Winners as saying: "We 'clicked' at once…. [Lorenz's] extraordinary vision and enthusiasm were supplemented and fertilized by my critical sense, my inclination to think his ideas through, and my irrepressible urge to check out 'hunches' by experimentation." Tinbergen and Lorenz's work was disrupted by World War II.

Tinbergen spent much of the war in a hostage camp because he had protested the State University of Leiden's decision to remove three Jewish faculty members from the staff. After the war ended, he became a professor of experimental biology at the University. In 1949, Tinbergen traveled to Oxford University in England to lecture. He stayed at Oxford, establishing the journal Behavior with W. H. Thorpe and working in the University's animal behavior division. His 1951 book The Study of Instinct is credited with bringing the study of ethology to many English readers. The book summarized some of the newest insights into the ways signaling behavior is created over the course of evolution. In 1955, Tinbergen became an English citizen, and in 1966 he was appointed a professor and fellow of Oxford's Wolfson College. When the work of Tinbergen, Lorenz, and von Frisch, who had demonstrated that honeybees communicate by dancing, received the Nobel Prize in 1973, it was the first time the Nobel Committee recognized work in sociobiology or ethology.

It was Tinbergen's own hope that the ethologists' body of work would help in understanding of human behavior. "With von Frisch and Lorenz, Tinbergen has expressed the view that ethological demonstrations of the extraordinarily intricate interdependence of the structure and behavior of organisms are relevant to understanding the psychology of our own species, " wrote P. Marler and D. R. Griffin in Science. "Indeed, [the Nobel Prize] might be taken … as an appreciation of the need to review the picture that we often seem to have of human behavior as something quite outside nature, hardly subject to the principles that mold the biology, adaptability, and survival of other organisms."

The ability of an organism to adapt to its environment is another element of Tinbergen's work. After he retired from Oxford in 1974, he and his wife attempted to explain autistic behavior in children to adaptability. The Tinbergens' assertion that autism may be caused by the behavior of a child's parents caused some consternation in the medical community. Tinbergen believed that much of the opposition to his work was caused by the unflattering view of human behavior it presented. "Our critics feel we degrade ourselves by the way we look at behavior, " he is quoted as saying in Contemporary Authors. "Because this is one of the implications of ethology, that our free will is not as free as we think. We are determinists, and this is what they hate…. They feel that our ideas gnaw at the dignity of man."

Tinbergen was wrote a number of books and made many nature films during his lifetime. Among his publications were several children's books, including Kleew and The Tale of John Stickle. Among the numerous awards he received are the 1969 Italia prize and the 1971 New York Film Festival's blue ribbon, both for writing, with Hugh Falkus, the documentary Signals for Survival, which was broadcast on English television. Tinbergen died December 21, 1988, after suffering a stroke at his home in Oxford, England.

Further Reading

Contemporary Authors, Volume 108, Gale, 1983, pp. 489-90.

Nobel Prize Winners, H. W. Wilson, 1987, pp. 1059-61.

"Learning from the Animals, " in Newsweek, October 22, 1973, p. 102.

Marler, P., and D. R. Griffin, "The 1973 Nobel Prize for Physiology or Medicine, " in Science, November 2, 1973, pp. 464-467. □

Tinbergen, Nikolaas (1907–88)A Dutch-born, but later British, zoologist who shared (with KonradLorenz and Karl vonFrisch) the 1973 Nobel Prize for Physiology or Medicine for studies of animal behaviour under natural conditions. Tinbergen was especially noted for his studies of social organization among gulls. He became professor of animal behaviour at the University of Oxford in 1966, and emeritus professor in 1974.

Tinbergen, Niko(laas) (1907–88) Dutch-born British zoologist and ethologist. Working first at Leiden University, he moved to Oxford in 1947, becoming professor of animal behaviour in 1966. With Konrad Lorenz he was a pioneer of ethology, working with animals in their natural setting. In later years Tinbergen attempted to apply ethological principles to human beings, especially autistic children. He shared the Nobel Prize for physiology or medicine with Lorenz and Karl von Frisch.

Tinbergen, Nikolaas (1907–88) A Dutch-born, but later British, zoologist who shared (with Konrad Lorenz and Karl von Frisch) the 1973 Nobel Prize for Physiology or Medicine for studies of animal behaviour under natural conditions. Tinbergen was especially noted for his studies of social organization among gulls. He became Professor of Animal Behaviour at the University of Oxford in 1966, and Professor Emeritus in 1974.