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The Nature and Nurture of LoveFrom Imprinting to Attachment in Cold War America$

Marga Vicedo

Print publication date: 2013

Print ISBN-13: 9780226020556

Published to Chicago Scholarship Online: September 2013

DOI: 10.7208/chicago/9780226020693.001.0001

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The Study of Instincts

The Study of Instincts

(p.43) Chapter Two The Study of Instincts
The Nature and Nurture of Love

Marga Vicedo

University of Chicago Press

Abstract and Keywords

This chapter analyzes Konrad Lorenz's views about instincts and imprinting in the context of his efforts to develop the new science of ethology with Niko Tinbergen. It first introduces Lorenz's and Tinbergen's project to explain the biological basis of animal and human behavior. It then examines more closely Lorenz's conception of instincts and imprinting. Next, it turns to Lorenz's role in discussions about maternal care and infants' needs. After psychiatrists like John Bowlby showed interest in his work, Lorenz increasingly extrapolated his ideas about imprinting in birds to the human realm.

Keywords:   Konrad Lorenz, instincts, imprinting, ethology, Niko Tinbergen, maternal care, infant needs, John Bowlby


In his 1953 book Child Care and the Growth of Love, John Bowlby asserted: “The theories put forward in this book, … are in strict agreement with what biological science has shown to be true of both bodily and mental growth.” He referred to the “European biologists” who had shown in birds and dogs that “emotional experiences at certain very early and special stages of mental life may have very vital and long-lasting effects.” The European biologists he was referring to were the Austrian Konrad Lorenz and the Dutchman Nikolaas Tinbergen, two students of animal behavior who aimed to institute the biological study of instincts as an independent branch of research: ethology.1

Several scholars have illuminated different aspects of the history of ethology. Richard Burkhardt, John Durant, Theodora Kalikow, Robert Richards, and others have examined central conceptual and institutional developments in the field. Klaus Taschwer and Benedikt Föger’s biography of Lorenz and Hans Kruuk’s biography of Tinbergen offer fascinating accounts of these two complex men. Together with D. R. Röell’s study of ethology in the Netherlands and Burkhardt’s comprehensive book about research on animal behavior, this literature provides a thorough account of the development and reception of ethology.2 Here I expand on those aspects of ethology that influenced ideas about infants’ needs and maternal (p.44) care and focus on Lorenz’s influence in child studies, a topic that has received less scholarly attention.

Ethology, and Lorenz’s work on imprinting in particular, became fundamental to the construction of mother love and love for mother as biological instincts. To show this, I first introduce Lorenz’s and Tinbergen’s project to explain the biological basis of animal and human behavior. They believed that human as well as animal behavior is a matter of instincts. I then examine more closely Lorenz’s conception of instincts and imprinting. Next I turn to Lorenz’s role in discussions about maternal care and infants’ needs. After psychiatrists like Bowlby showed interest in his work, Lorenz increasingly extrapolated his ideas about imprinting in birds to the human realm.

Ethology: Lorenz and Tinbergen Search for the Biological Basis of Behavior

Konrad Lorenz (1903–1989) was the son of a world-famous orthopedic surgeon and professor at the University of Vienna. Lorenz arrived late in his parents’ marriage, eighteen years after his only brother. He grew up in Altenberg, a small Austrian town near Vienna, in a mansion whose extensive grounds, and the surrounding forest near the Danube, served as his playground and, later, as his observational and experimental site.3 In this wealthy, educated household, his devoted parents indulged his passion for animals. Lorenz raised dogs, fish, and a variety of birds with the help of Margarethe (Gretl) Gebhardt, a neighbor’s daughter with whom he shared games and animals beginning in childhood. Following his father’s desires, Lorenz studied medicine at the University of Vienna, where he earned his doctorate in 1928. In 1933 he also obtained a doctorate in zoology there. A few years earlier, in 1927, he had married Gretl, who became a medical doctor and a lifelong supporter of her husband’s projects.

Lorenz was interested both in animal and human behavior from early on, as can be seen in his training in comparative anatomy, ornithology, and human psychology. In medical school, Lorenz studied with the renowned comparative anatomist Ferdinand Hochstetter. Under his tutelage, Lorenz learned to identify homologous anatomical structures and use them to trace common ancestries. Later, Lorenz used instinctive behaviors to reconstruct evolutionary phylogenies. In ornithology he received the approval and encouragement of the influential German ornithologist Erwin (p.45) Stresemann. In addition, the assistant director of the Berlin zoo Oskar Heinroth, who studied instinctive behavior patterns in birds, also mentored Lorenz. In human psychology, Lorenz was a student of Karl Bühler, director of the Psychological Institute at the University of Vienna and a world leader in several areas of psychology, including Gestalt perception and child development. Bühler had also written about instincts and, in a successful book that went through several editions, reviewed the various schools of psychology, including Gestalt psychology, behaviorism, and psychoanalysis. Lorenz did one of his auxiliary fields for his PhD exams with Bühler and regularly attended his seminars.4

From Bühler, Lorenz learned about the major discussions in human psychology, such as the debate between psychologists William McDougall and John Watson on the role of instincts in human conduct. Whereas McDougall defended the notion that humans were moved by several key instincts and emotions, Watson argued that they were born with very few innate predispositions. Lorenz soon concluded that much work remained to be done to prove the importance of instincts in human social behavior. Convinced of the scientific and social significance of his goals, Lorenz felt ready for that task. Observing birds, he wondered what his studies implied for human behavior. As he put it in a 1931 letter to Heinroth: “Who knows what will become of today’s human psychology if one can only know what is instinctive behavior and what is rational behavior in humans? Who knows how human morals with their drives and inhibitions would look if one could analyze them like the social drives and inhibitions of a jackdaw.”5

Only a few years later, Lorenz presented a unitary framework for explaining animal and human social behavior. Between 1927 and 1935, he published detailed observations of the behavior of half-tamed jackdaws, ravens, and night herons, as well as his views about instincts.6 Then in a 1935 paper, “Companions as Factors in the Bird’s Environment: The Conspecific as the Eliciting Factor for Social Behaviour Patterns,” he put forward a general framework for understanding social behavior that he would defend with only minor modifications for the rest of his career. In this massive essay, Lorenz included observations on almost thirty types of birds and described their social behavior as a set of instinctive responses that had been built by natural selection because of their survival value.7

Lorenz posited the existence of innate templates or schemas (later called innate releasing mechanisms) that, when activated by internal and external releasers, lead the bird to perform certain instinctive behaviors. (p.46) He elaborated on the role of various fellow members of the species, the “companions,” that act as releasers of a bird’s social behavior: the parental companion, the infant companion, the sexual companion, the social companion, and the sibling companion. According to Lorenz, the bird’s reactions to the specific stimuli from each of these companions form a functional system. For each of these relationships, there is an innate releasing mechanism, some type of conduct or image in the companion that functions as releaser, and an instinctive pattern of behavior that is “released” in an automatic and uniform manner.8

In this paper Lorenz asserted that “social behavior patterns in particular are to a large part governed by instincts even in the highest animals.” He concluded with a call to arms, urging researchers to “recognize that the instinct, governed by its own laws and fundamentally differing from other types of behavior, is also to be found in human beings” and encouraging them to investigate human instincts.9 At this point, however, Lorenz continued studying birds with renewed energy after meeting Niko Tinbergen, a young researcher with whom he had been corresponding.

Nikolaas Tinbergen was born in The Hague in 1907, the son of a grammar school teacher. He and his four siblings were raised in an austere household by parents who had a strong sense of public duty. Throughout his youth, Tinbergen was more interested in the outdoors and in field hockey than in intellectual matters. After a trip to an ornithological field station in Germany, he decided to turn his interest in wildlife into a career in field zoology.10

Tinbergen obtained his PhD in biology at Leiden University in 1932 with a dissertation on the homing behavior of the digger wasp Philan-thus, based on field experiments he had done in the sand dunes area of Hulshorst. After he married Elisabeth (Lies) Rutten, they spent one year in Greenland with a meteorological expedition. Tinbergen then returned to his post as assistant in the Department of Zoology at Leiden University. With his students, Tinbergen continued his experiments on the hunting and homing behavior of wasps, added observational work on the parental behavior of hobbies (birds), and carried out experiments on the territorial behavior of sticklebacks (fish). When he met Lorenz, he was ready for a theoretical framework that could help make sense of those diverse observations and experiments.

At first sight Lorenz and Tinbergen seemed an unlikely team because they had very different personalities, but they immediately found common ground in their shared interests. Flamboyant and self-assured, Lorenz reveled (p.47) in being the center of attention at scientific and social gatherings. He was an animated speaker who, much to the delight of his audiences, readily went down on all fours to imitate a variety of animal sounds and gestures. Convinced of the scientific and social importance of his work, he presented his views with messianic zeal. Tinbergen, on the other hand, battled with self-doubt throughout his life and often questioned the value of his contributions to science and society. But they both enjoyed being outdoors and were drawn to physical activities. Above all, they shared a deep love for nature. Both men had spent their childhood and youth in the open air, observing animals, and they continued this throughout their lives.

When Tinbergen spent three and a half months with Lorenz at his Altenberg home and research site in 1937, they carried out observations and experiments on a variety of birds. Among the best known are their experiments to test young turkeys’ reaction to simulated predators and to analyze the egg-retrieval movements of greylag geese. In the latter, they filmed a goose rolling a stray egg back to her nest with her beak. Once the goose started, she would automatically continue her motion toward the nest even when the investigators replaced the egg with objects like a huge egg or a cube, and even when they took the egg away altogether.11

Lorenz and Tinbergen believed that their individual research skills complemented each other. Lorenz was impressed by Tinbergen’s ability to design experiments, a good counterpart to his own interest in observation. Tinbergen admired Lorenz for his theoretical ideas, while Lorenz realized that his theories would benefit from the evidence provided by Tinbergen’s work. So joining forces would benefit them both. From then on, although they developed most of their research separately, they conceived of their work as part of a common enterprise.

The two men decided to build and promote a modern biological science of animal behavior—ethology—focused on the study of instincts. This was not an entirely new endeavor. As Richard Burkhardt has shown, many of the elements of their approach to studying instincts were already present in the work of researchers of the previous generation. What distinguished Lorenz and Tinbergen was their concerted effort to make the study of instinct a distinct and central branch of biology.12 Together they defended the notion that observation should be the key method for studying animal behavior, and they encouraged the use of photography in research.13

Tinbergen promoted their ethological program during a 1938 trip to (p.48) the United States, where he delivered a series of lectures at the American Museum of Natural History and the Linnaean Society of New York and participated in other scientific meetings. Meanwhile Lorenz remained in Austria, continuing his research, pursuing funding, and trying to secure a permanent position. The prospects for ethology looked promising.

But World War II rudely interfered, putting a stop to their efforts to institutionalize ethology as an independent branch of biology and separating them geographically as well as politically.

In 1938, soon after Germany annexed Austria, Lorenz joined the Nazi party and sought to get ahead in a regime that he thought would support his scientific interests, including human psychology. He tried to obtain the chair in human psychology at the University of Vienna left vacant by the Nazis’ dismissal of his former teacher Karl Bühler, “for political and world-view reasons.” His wife, the respected psychologist Charlotte Bühler, had one Jewish parent and was dismissed on racial grounds.14 They both emigrated and eventually found jobs in the United States, although Karl Bühler never again worked at the prominent level he had attained in Europe. Lorenz did not obtain his position, but two years later he was awarded a professorship of psychology at Albertus University in Königsberg, in East Prussia. Barely a year later, however, he was drafted into the German army.

After a brief stint as a motorcycle-riding instructor, Lorenz spent two years (from summer 1942 to spring 1944) as a psychiatrist in Poznan, Poland. From the scarce documents available, historians have been unable to determine whether his work as an assistant to race psychologist Rudolf Hippius contributed to Nazi activities. Later Lorenz denied his membership in the Nazi party and said he never suspected the Nazis had mass murder in mind.15

During this period, in addition to articles on bird behavior and the concept of instinct, Lorenz wrote on the consequences of interfering with instinctive behavior in animals and humans. Comparing wild and domesticated animals, Lorenz defended the superiority of the wild forms. He pointed to the short extremities, fat belly, and early sexual maturation and promiscuity of females in some domestic breeds as evidence of degeneration from their wild state. Lorenz believed that civilization in humans, like domestication in animals, led to degeneration owing to the deterioration of innate releasing mechanisms. In humans, he argued, civilization led to physical degeneration as well as a decline in innate moral and aesthetic capacities.16

(p.49) Presenting domestication in animals and civilization in humans as analogous helped Lorenz to justify extrapolating animal research to the human realm and also to foreground the social relevance of his own expertise. Note the elaboration of this chain of reasoning in the following text Lorenz published in a 1940 newspaper:

Fighting spirit and motherly love, the characteristics that are necessary for the preservation of the species, are being lost not only in animals, but in humans as well, through the process of civilization. This is why one may draw comparisons between the two realms without further ado. Race politics knows that the continuous ups and downs, the flowering and decline of cultures arise when the victorious people become complacent. Today, the biologist researches consciously and with scientific precision the causes of these phenomena.17

Here Lorenz established the social significance of his expertise by arguing that biologists could help to improve the race, or at least to avoid its further degeneration. In turn, the urgent need for biological knowledge justified the extrapolation from animals to humans. By focusing on a major Nazi concern, race purity, Lorenz defended his views about the perils of civilization for the degeneration of the human race while advancing the importance of his scientific work.

During the Nazi period in Germany and Austria, Lorenz took advantage of a political climate favorable to these ideas by presenting his views in the rhetoric prevalent at the time. In several writings dating from the early 1940s, he couched his concerns about social decadence in the language of racial degeneration and used medical metaphors to cast the alleged deterioration of human instincts as a cancer of the social body. Although he had not done any research on the behavior of humans, in 1943, eight years after his call to investigate instincts in humans in the “Companions” piece, he published another massive paper, “The Innate Forms of Possible Experience,” in which he summarized his views on the major areas of his thinking: animal behavior, evolutionary epistemology, the instinctual nature of human behavior, and the negative impact of civilization on human instincts. He postulated the existence of innate releasing schemas of behavioral systems that shape human experiences, including those related to aesthetic and ethical appreciation.18

This essay, written while he was a military psychologist in occupied Poland, was Lorenz’s last publication before the end of the war. In April 1944 he was sent to the front as a physician and almost immediately went (p.50) missing. The Russians sent him to a prisoner-of-war camp in Yerevan, Armenia. By February 1946, his wife heard about his situation. In February 1948, he returned to Austria.

Back home and in need of income, Lorenz published a book for the general public about his experiences raising and living with animals. In German, the title was Er redete mit dem Vieh, den Vögeln und den Fischen (He spoke with the beasts, the birds and the fish). It was translated into English as King Solomon’s Ring: New Light on Animal Ways. This was not a scientific book on animal behavior, but a collection of anecdotes that showcased Lorenz’s knowledge of birds and other animals as well as his enormous talent as a storyteller. He described animal behavior in a dramatic narrative consisting of major episodes of human interest such as falling in love, marriage, and fighting. This was embellished with tales of how his family life was deeply intertwined with the animals he was studying and illustrated with his own charming drawings. The book became a best seller and made Lorenz a household name internationally.19

Reunited with his fellow ethologists in Austria and Germany, Lorenz resumed his research on animal behavior. He obtained some funds from the Austrian Academy of Sciences to continue his work in Altenberg. In 1950 support from a German foundation, the Max Planck Society, enabled Lorenz, his family, some collaborators, and many animals to move from his deteriorating mansion to the castle of a German aristocrat in Buldern, in northern Germany, where he continued his research. In 1958 the Max Planck Society provided him with a permanent research setting. Lorenz and the physiologist Erich von Holst were made directors of the Max Planck Institute in Behavioral Physiology in Seewiesen, in southern Germany.

During the war, Tinbergen also went through dramatic experiences that affected him profoundly. Together with 80 percent of the faculty of his university, Tinbergen protested the dismissal of Jews and the Nazification of Dutch universities. He was detained and spent the war in a prison camp. With the paper he was allowed to use, he wrote a weekly letter to his children, telling them animal stories that he illustrated and would later publish in books for children. After his liberation, Tinbergen returned to teach at Leiden University in difficult postwar circumstances. When conditions improved, the university created a chair of experimental zoology for him. But Tinbergen, who was interested in expanding ethology outside the Continent, sent out feelers to colleagues in England and the United States. In late 1948 he accepted an offer to be a lecturer (p.51) in animal behavior at Oxford University, where he later became a professor of zoology.

Lorenz and Tinbergen reconnected at a 1949 symposium, “Physiological Mechanisms in Animal Behavior,” organized by the British Society for Experimental Biology in Cambridge. Although still bitter about the war and disappointed by Lorenz’s support for the Nazis, Tinbergen believed that reestablishing international connections with German scientists was necessary to heal the wounds of the past. Seeing Lorenz as the one who had provided the main conceptual framework for ethology, he decided to renew their relationship for the sake of the field.

Energized by the opportunity to launch his program again at the Cambridge symposium, Lorenz staked his claim to the independent status of ethology as a science. First he advanced the object of study: “The distinct and particulate physiological process whose discovery may be identified with the origin of comparative ethology as an independent branch of science is represented by a certain type of innate, genetically determined behaviour patterns.” As for method, Lorenz argued that “unprejudiced observation” was the appropriate way to study genetically determined behavior patterns. In his view, ethologists should start by developing “a morphology of behaviour” by “a thorough observation and description of all the behaviour patterns at the disposal of the species.” The systems of behavioral patterns uncovered by ethology would then reveal natural laws of universal character.20

Drawing extensively on the conceptual apparatus he had developed in the 1930s, Lorenz again proposed a unitary framework for animal and human behavior. This implied considering human psychology from the standpoint of the central object of ethology, the instincts. He claimed that “the laws we have found in animal behaviour find an enormously important application to the special phenomena of human psychology.” However, he did not specify which laws or which applications he was referring to.21

Tinbergen presented the ethological program in his 1951 book The Study of Instinct, which became the foundational text of ethology. As the “scientific study of behaviour,” ethology first selected “a specific object, a special group of phenomena: innate behaviour.”22 Tinbergen also presented Lorenz’s views on human instincts, arguing that Lorenz’s writings in this area had already provided substantive evidence for the instinctual nature of human social behavior.

In sum, Lorenz and Tinbergen aimed to create an independent science (p.52) of ethology that would explain social behavior by focusing on the study of instincts in animals and humans. But what, exactly, were the main objects of this science, the instincts? In the next section I focus on Lorenz’s views about instincts, for two reasons: During the early years of ethology, Lorenz was the main theorist of the notion of instinct. And Bowlby and other child analysts borrowed the concept from his writings.

The Nature of Instincts

Lorenz and Tinbergen portrayed social behavior as guided or even determined by instincts. But what were instincts? And how could an investigator identify instinctive behavior? These were key questions for their ethological program.

Lorenz originally thought of instincts as reflexes, or chains of reflexes, set off by external stimuli, but he abandoned this view under the influence of the German physiologist Erich von Holst and the American biologist and animal researcher Wallace Craig. In 1937 Lorenz met Holst, who had shown that automatic behaviors generally cataloged as reflexes, such as creeping movements in earthworms, were caused by internal stimuli. Richard Burkhardt has shown that Lorenz was also heavily influenced by his correspondence with Craig.23 Noting that animals are restless before performing an instinctive act but usually relaxed afterward, Craig argued that animals desire to perform their innate behaviors. In addition, the animal sometimes performs the action in the absence of the relevant environment or stimulus (“vacuum response”).24

In Lorenz’s new vision of animal behavior, the organism’s central nervous system produces a series of internal stimuli that lead to an appetitive behavior, expressed in restlessness and searching. Impulses build up inside the organism. Then they are unleashed by an innate releasing mechanism (IRM) and by social releasers (SR), which “open the doors” to the organism’s appetitive actions. Usually the IRM discharges in response to an external stimulus, but it can also explode “in vacuo.” On some occasions “displacement activities” substitute for the organism’s normal, natural, or instinctive actions.25 For Lorenz every fixed-action pattern, a stereotyped action performed by all members of a species, has its own independent drive system. If a fixed-action pattern is not carried out, its “action-specific energy” builds up until it is eventually released. The intensity of a behavioral response depends on the amount of accumulated action-specific energy and the strength of the external stimulus.26

(p.53) Lorenz devised a psychohydraulic model to explain how instincts work. Although he introduced a new vocabulary to refer to the components in the model, he relied on concepts of drive, energy, and release prevalent at the time in physiology, psychology, and psychoanalysis.27 Like many of those models, Lorenz’s motivational model is analogous to a hydraulic reservoir. He presented the following diagram: The stream flowing from the tap represents the internal accumulation of action-specific energy. The spring represents the central inhibition that must be overcome. The valve is the internal releasing mechanism. The liquid spouting from the lower tap represents the instinctive movement (see fig. 2.1).

The Study of Instincts

Figure 2.1. Konrad Lorenz’s psychohydraulic model of instinctive action. From Konrad Lorenz, “The Comparative Method in Studying Innate Behaviour Patterns,” Symposia of the Society for Experimental Biology 4 (1950):256.

The objects of study in ethology—instincts—sometimes went by different names: drive activities, instinctive movements, instinctual behavior patterns, fixed-action patterns, fixed motor patterns, innate behavior, innate behavior patterns, “genetically determined behavior patterns,” or simply instincts. At the 1949 meeting in Cambridge, Lorenz had proposed “endogenous movements” to unify the diversity of concepts, but this term (p.54) never acquired widespread acceptance, and Lorenz himself continued to use a variety of names.28

Many students of animal behavior also used these terms interchangeably, but it is important to note some important differences among them. A fixed-action pattern, one of the most common names in the ethological literature, refers to a pattern of behavior that is a stereotyped action performed by all members of a species. Whether animals perform an action in a stereotyped way can be ascertained by observation. But the other names all refer to something that cannot be determined by observing a behavior, namely, its innateness or instinctual nature. Tinbergen and Lorenz were interested in innate fixed-action patterns. But how could innate or instinctive behavior be identified?

According to Lorenz, instinctive actions share a series of features: they are species-specific, carried out by all members of a species in a stereotyped way; they always follow a given stimulus; and they continue until their consummation. That is, they are “machinelike” in character, an automatic sequence that, once set in motion, continues inexorably until its programmed completion. These behaviors are also satiable. Consequently, they are less likely to take place after several repetitions. But the action is sometimes performed “in vacuo” when the appropriate stimulus is not present.29

Further, the central characteristic of these innate behavior patterns is that they cannot be modified by training. Contrary to other theorists of instinct such as Darwin and William James, Lorenz posited a radical separation between instinctive and learned behaviors. In a given behavior, there may be a conglomerate of both types, what he called an “intercalation” of the innate and the learned (Instinkt-Dressurverschränkung), but the innate and learned components can be separated. In addition, he thought that an instinct could be treated “as an organ, whose individual range of variation can be neglected in the general biological description of a species.” In Lorenz’s account, the instinctive is uniform across the species.30

Although many of the characteristics of instinctive behavior can be observed, Lorenz argued that to be sure a behavior is innate, one needs to perform deprivation, or “Kaspar Hauser,” experiments.

One day in May 1828, a disoriented young man was found in a town square in Nuremberg, Germany. He could not talk, walked with difficulty, and was afraid of many objects and animals. The authorities concluded that Kaspar Hauser, as he later called himself, had been locked in a cellar from infancy and had lived in isolation from the external world. His case, (p.55) like those of other severely deprived children and “wolf children,” or wild children, fascinated scientists because humans raised in isolation constitute a sort of experiment that can illuminate the nature-nurture controversy. The reasoning goes like this: if a behavior or trait that cannot have been influenced by training appears in an individual, then it must be inborn or instinctive.

The justification for using isolation experiments to find animals’ instincts is based on a similar intuition. Lorenz believed a behavior that appears in an animal raised in isolation has to be innate, since it is not the result of imitation, practice, exercise, conditioning, or any other type of learning. Lorenz’s mentor Oskar Heinroth and his wife, Magdalena, had engaged in an extensive program of such experiments in the early decades of the century to ascertain what was innate and what was learned in various species of birds from central Europe.31 Following their insight, Lorenz believed that through observations and isolation experiments, researchers could identify an animal’s instincts.32 This was a key point, since he claimed that most social behavior is instinctual or innate.

Although Lorenz said that many social patterns of behavior are innate, he also claimed that the object that will release those patterns is not innate, at least in some species. Instead, it is acquired through imprinting.


As I mentioned before, in his 1935 paper “Companions as Factors in the Bird’s Environment,” Lorenz introduced the bird’s companions as factors eliciting its instinctive behavior. In this paper, Lorenz also examined imprinting, which his mentor Heinroth and others before him had already observed in some types of birds and which was well known to farmers and breeders.

Imprinting is the process whereby some species of birds attach to the first moving object they see after hatching. Usually that object is their mother, and through that process the bird comes to recognize its conspecifics and develops the social responses of its species. The process of imprinting is very important for these species because, although their social behavior patterns are innate, the birds do not recognize the members of their species until they are imprinted. Lorenz had observed this phenomenon in jackdaws, geese, and many of his hand-reared birds. Lorenz elaborated on the significance of imprinting, and described its main characteristics:

(p.56) I have described this behavior of the Greylag gosling because it provides a virtually classic example of the manner in which a single experience imprints the relevant object of the infantile instinctive behavior patterns in a young bird which does not recognize this object instinctively. This object can only be imprinted during a quite definite period in the bird’s life. A further important feature is the fact that the Greylag gosling obviously “expects” this experience during a receptive period, i.e., there is an innate drive to fill this gap in the instinctive framework.33

The Study of Instincts

Figure 2.2. Konrad Lorenz followed by birds imprinted on him. Courtesy of Konrad Lorenz Archive, Altenberg, Austria.

For Lorenz, imprinting has clear characteristics. First, it takes place during an early critical period. Afterward the brain, like hardened wax, cannot be molded. Second, the bird has an innate urge to acquire the imprint of its species and will do so through a single impression, without conditioning, trial and error, or any learning period. As Lorenz put it, imprinting “has nothing to do with learning.”34 Third, it is irreversible.

By fixing the object of the bird’s reaction in infancy, imprinting also determines future social and emotional reactions. If the infant bird is not imprinted on a member of its own species, it will still develop the instinctive social and sexual behavior patterns characteristic of its species, but it will direct them toward the wrong object. For example, some of the birds imprinted (p.57) on Lorenz courted him later. Imprinting thus has irreversible consequences for the animal’s behavioral development.35

In normal circumstances, the mother provides the image of the right species, filling the “gap” in her infant’s instinctual framework. The mother thus enables the baby bird to release its instinctive social behavior toward the appropriate objects. In that sense the mother-infant relationship is essential for the bird’s adequate development. One could say the baby bird has a built-in, instinctive need for the mother. Furthermore, the mother has also been designed by evolution to provide the appropriate responses to her infant.

The mother’s behavior toward her offspring is also innate, according to Lorenz. In his view, one should not talk about the parental instinct in the singular because parental care includes many small innate components. In birds, it includes building a nest, then feeding and protecting the chicks. Lorenz considered these behaviors innate, especially those involved in maternal care. Discussing the innate schema of the infant companion, Lorenz noted that in most cases parents recognize their progeny instinctively. In addition, the characteristics that release parental conduct cannot be acquired through learning, since “the adult bird’s own offspring are of course the first freshly-hatched conspecifics which it sees, and yet it must react to this first encounter with the entire repertoire of parental behavior operating to preserve the species.”36

In his writings on humans and other animals, Lorenz usually talked about parental behavior or parental care or parental love, but most of his examples of those behaviors and emotions came from the female of the species. He argued that the chick’s innate responses form a functional whole with the responses of the mother. Invoking his major test for innateness, Lorenz noted that the fact that the mother engages in maternal care toward her first infant without previous training proves its instinctive character. Furthermore, Lorenz claimed that maternal behavior is automatic and independent of the offspring’s behavior; this too confirmed its instinctive nature. For example, he reported that a Cairina mother would “rescue” a mallard duckling from the experimenter’s hands, even though minutes later she would bite and kill it when it tried to mix with her own chicks. In a sense, the mother cannot refrain from helping the infant in the first place. Her instincts impel her to act. For Lorenz, the “automatic nature of these parental care responses” proves that “the unitary treatment of the offspring is thus determined within the instinctive framework of the adult bird and not in the rôle that the infant plays in its environment.”37 (p.58) That is, the actions involved in caring for the offspring are predetermined in the instinctive framework of the parents.

In sum, Lorenz postulated the existence in some species of birds of an innate need for imprinting—a process that in normal circumstances leads an infant to attach to its mother. He also postulated an innate mechanism that leads a mother to look after her infant. In the case of maternal care, the mother’s actions are predetermined within her instinctive framework. When Lorenz talked about innate behavior and its releasers, he often employed the metaphor of a lock and key. For any particular lock, “the form of the key-bit is predetermined.”38 The same holds for the behavior within the mother-infant dyad. The preservation of the mother-infant behavioral system, understood as an interlocking system of instincts, is essential for adequate infant development. But could these findings be generalized to humans?

For humans, Lorenz postulated the existence of an innate releasing schema toward babies, one that is especially strong in women. He highlighted the significance of the “inborn schemata of the infant” or, as he called it later, an innate releasing mechanism. According to Lorenz, the existence of this innate system of behavior toward infants could be deduced from the identification of innate feelings associated with particular objects described as herzig, a term that combines the connota-tions of “sweet,” “neat,” and “cute” in English. An encounter with such an object releases the instinctive movement of “taking in the arms,” as he said he had witnessed in a striking episode. When she was an infant, Lorenz’s daughter saw a doll and, in seconds, ran to take it in her arms with a “motherly” expression. The automatic character of the response and the determination shown in a behavior she performed for the first time seemed comparable to the instinctive reactions of animals.39 Lorenz described his daughter’s response to a doll as equivalent to the Cairina mother’s response to a chick from another species.

Lorenz presented a diagram illustrating the features that release parental behavior. On the left are faces possessing the features that release parental behavior in humans: small heads, round features, big eyes (herzig features). They are contrasted on the right with faces that lack those characteristics (see fig. 2.3).40

The Study of Instincts

Figure 2.3. According to Lorenz, small, round heads with big eyes, bulging craniums, and retreating chins elicit human parental responses; organisms without those features (in the right column) do not. From Konrad Lorenz, “Die angeborenen Formen möglicher Erfahrung,” Zeitschrift für Tierpsychologie 5 (1943):276. Courtesy of Konrad Lorenz Archive, Altenberg, Austria.

Before World War II, Lorenz had consistently maintained that his views about the role of instincts in social behavior applied to humans as well as to animals. Elaborating on those views, his paper at the 1949 Cambridge symposium claimed that the existence of human instincts was (p.59) proved by the existence of human emotions. Here he was following scholars as diverse as Charles Darwin, William James, William McDougall, and John Watson, who had maintained that an emotion always correlates with an instinct, as I discussed in chapter 1. Except for McDougall, Lorenz did not refer to these authors. In his “Companions” paper, Lorenz had written that McDougall had “demonstrated that particular instinctive behaviour patterns are dependent upon specific emotions as subjective correlates.” Now Lorenz said he agreed with McDougall’s view that “man has just as many ‘instincts’ as he has qualitatively distinguishable emotions.” He then used this correlation as a tool to infer instincts from emotions.41 This correlation between emotions and instincts explains why we often find Lorenz, as well as other authors, using the terms maternal care and maternal love interchangeably and assuming that if the first is instinctive, so is the second.

Following this line of reasoning, Lorenz argued that parental behavior (p.60) in humans is innate. Incorporating the material used in his earlier writings, he contended that this was “proven” by the existence of the emotions associated with looking at and interacting with babies. In Lorenz’s words:

It is a distinct and indubitably sensuous pleasure to fondle a nice plump, appetizing human baby…. In this case, the existence of a true innate releasing mechanism in man has been clearly proven…. Also, the objective and subjective reactions activated by the mechanism are clearly distinguishable. A normal man—let alone a woman—will find it exceedingly difficult to leave to its fate even a puppy, after he or she has enjoyed fondling and petting it. A very distinct “mood,” a readiness to take care of the object in a specific manner, is brought about with the predictability of an unconditioned response.42

As with other instincts, the behavior’s emotional quality, the fixity of the response, its universality, and its machinelike character proved the instinctual basis of parental, and especially maternal, behavior.

In 1953, ten years after his long paper on innate forms of possible experience, Lorenz published a condensed but otherwise almost verbatim account of his views about human instincts. In this paper too, Lorenz devoted much attention to parental care, repeated his views about the innate releasing mechanism toward the cute—especially strong in women—and asserted the instinctive nature of mother love as well as monogamous love.43

After the war, building on the success of his popular book King Solomon’s Ring, Lorenz devoted himself to promoting his ideas and reaching large audiences. He published more often in English, had many of his earlier essays and books translated, and welcomed foreign visitors and researchers to his research station. In addition, he traveled extensively in the United States, where in 1954–55 he delivered talks at prestigious universities such as Harvard, Cornell, and Clark University, as well as public lectures in cities like Boston and New York City. He also attended numerous international meetings throughout Europe. In some of those meetings, encouraged by psychiatrists’ interest in his work, Lorenz’s assertions about instinctive behavior in humans became increasingly bold.

The WHO Meetings: Imprinting from Birds to Infants

Lorenz found an eager audience for his views on social behavior in an international Study Group on the Psychobiological Development of the Child, organized by the World Health Organization (WHO). Chaired by (p.61) one of the pioneers in studies about children and human emotions, Frank Fremont-Smith of the Josiah Macy Jr. Foundation, this group met in Geneva in 1953, in London in 1954, and in Geneva again in 1955 and 1956. Besides Lorenz, members included the British psychoanalyst and psychiatrist John Bowlby, the American anthropologist Margaret Mead, and the Swiss psychologist Jean Piaget. Guest speakers included biologist Julian Huxley and psychoanalyst Erik Erikson. The meetings focused on developments in ethology and their implications for child psychology.44

Bowlby made sure the mother-child relationship would be a focal point of the discussions. In his introduction during the first meeting, he highlighted his interest in ethology and noted that his investigations into the effects of separation from the mother had led him to Lorenz’s work. He was interested in finding out if imprinting operated in humans and could help explain infants’ negative reaction to separation from their mothers. He also noted that ethological studies of the mother-child relationship should be of interest to psychoanalysts, who had already placed the mother-child dyad at the center of social development. As Bowlby put it, “The phenomenon of imprinting at once struck me as possibly important to my work. Whether it really has anything to do with the effects of separation we shall see. The other thing that fascinated me in [Lorenz’s] work was the mother-child relationship of animals. The mother-child relationship is manifestly an example of instinct, in the ethological meaning of the word, and it is also at the centre of psychoanalysis.”45

As early as 1934, Lorenz had been aware that some psychologists and psychiatrists showed interest in imprinting. He wrote to Stresemann: “I have finally gotten in contact with the psychologists, that is, they now know about me. The psychiatrists also start to be interested in the phenomenon of imprinting. They believe there could be something similar in humans, which I do not think is impossible. But to see it clearly, I would need a series of 6 male and 6 female Kasper Hausers, who will hardly be granted to me!”46

Now, speaking before an audience with few experts in biology, Lorenz made daring pronouncements about human behavior. In a memorandum he sent to the WHO regional office a few weeks before the first meeting, Lorenz said he would focus on two processes of interest to students of child development: innate releasing mechanisms (IRMs) and imprinting. He would talk about the existence of IRMs in the human species and deal “with the extreme probability of imprinting in human children.” Finally, he would treat the pathological disintegration of IRMs and the pathology of imprinting. Lorenz’s presentation on imprinting elaborated on the significance (p.62) of this phenomenon, which until that point had been observed in only a few species: “Though imprinting has been found in its typical form in birds and insects rather than in mammals, I really do believe it to be fundamentally akin to those very lasting object-fixations of human beings, chiefly because these fixations also seem to be dependent on early childhood impressions and seem also to be largely irreversible. Some psychiatrists and psychoanalysts here I believe share this opinion, at least as a working hypothesis.”47

During the discussions, Lorenz acknowledged that little could be said with certainty about humans or any mammal: “We don’t know a thing about them…. Maybe in about five years I can just tell you something about small monkeys, or lemurs, with which we intend to start.” Further, he pleaded: “As to experiments, I must ask you not to expect too much knowledge about imprinting in man from ethologists.”48

Yet Lorenz had already stated his belief in the instinctual nature of human social behavior in no uncertain terms. Here, and in much the same words as in his previous writings, Lorenz asserted that human maternal behavior was a clear instance of innate behavior, as proved not by biology but by social interactions. This is how he put it:

But now let me proceed to what interests us most, the mother-child relationship. One of the best instances of the I.R.M., except for the snake, is our reaction to the quality of cute…. Now, let’s look at the properties which produce the impression of a thing being cute. The head must have a large neurocranium and a considerable recession of the viscerocranium, it must have an eye which is below the middle of the whole profile. Beneath the eye there must be a fat cheek. The extremities must be short and broad. The consistency of the body ought to be that of a half-inflated football, elastic; movements that are rather clumsy elicit the reaction very strongly; and finally the whole thing must be small, and must be the miniature of something.49

In short, “cute” equals “baby”—as he had already asserted in his 1943 and 1953 papers.

But how can we know whether parental care toward babies is instinctual? To answer the question, Lorenz appealed to the results of social “experiments”:

Now, in order to see whether many people have got that I.R.M., we ought to do a mass experiment with thousands or millions of experimental persons. Just (p.63) this experiment has already been done: it has been done by the doll industry, which, of course, sells the supranormal object best. The exaggeration of key-stimuli can be very nicely shown in the “cupie” [kewpie] doll, and the “Käthe Kruse Puppe” in German, and if you want facts on what I say, then go to Walt Disney’s films and see how Walt Disney represents cute animals.50

In the supranormal or supernormal object the characteristics that stimulate the release of an innate behavior are exaggerated. For example, Tinbergen had shown that the oystercatcher (a bird) prefers a giant egg, even an artificial one, to a normal egg. In other cases it is not size but some other characteristic of the object that triggers an animal’s instinctual reaction.51

Lorenz portrayed the appeal of a doll as support for the existence of a female maternal instinct. Yet Lorenz had not conducted research on these industries, nor did he present data on these issues from other sources. In addition, he did not consider any alternative explanations for the behaviors he was discussing. For example, he did not examine the possible role of environmental influences on maternal behavior or attitudes. By this point several researchers, including the American psychologist Leta Hollingworth and the sociologist Ruth Reed, had done extensive research showing that society’s emphasis on women being nurturing, their roles as caretakers, and social expectations that they should be “maternal” all influenced their interest in babies.52

Lorenz, however, asserted not only that maternal behavior is instinctive but also that the value societies place on such behavior is innate:

We must keep in mind that mother-love is not more necessary to the survival of the species than the drive to copulation. Why, then, are those drives to copulation “brutish” and why is “maternal love” sublime? This is simply our emotional valuation of instinctive behaviour in man—and it is largely dependent on supply and demand. I am convinced that we have something very deep, innate, in our behaviour, which tends to devalue sex and eating and to value very highly mother-love, social behaviour, defence of the family, and so on.53

Anthropologist Margaret Mead, who became a close friend of Lorenz but was also one of the few people in these meetings who sometimes criticized his views, pointed out that no such universal valuation exists, since one could find “societies which put a high value on sex and eating, and a low value on maternity.”54 Lorenz did not respond to her objection.

(p.64) When he turned to parental behavior during the third meeting, held in 1955, Lorenz maintained that there was only a quantitative difference between men’s and women’s reactions to babies. Only cultural mores prevented “giving utterance to these, certainly instinctive, urges” in males. However, he also proposed that the urge to develop different gender roles is innate:

Well, I had better come out and be honest about what I am aiming at. I do believe that there is a certain unlearned element—something like an IRM— which makes the little boy actually seek for somebody to take over the father role. Sylvia Klimpfinger has evidence for that in a hospital—a hostel—where all the children are reared by the female staff alone, and all these children—the boys more significantly than the girls—go for the gardener who is the only male accessible to them. This led me to suspect that there might be an unlearned preference for what to imitate—boys to imitate Pa and for girls to imitate Ma.55

In sum, after asserting that biologists knew nothing about the biological basis of behavior in mammals, let alone humans, Lorenz argued that parental and sexual roles are innate in humans, as are their ethical valuations of those roles. According to him, humans possess an internal releasing mechanism for parental behavior toward babies, as shown by a universal tendency to consider baby features cute. In addition, people instinctively give a high moral value to mother love. Finally, there is an innate preference for boys to imitate their fathers and for girls to imitate their mothers. The combination of those points amounted to an argument for the instinctual basis of traditional gender roles and, specifically, gendered parental roles.

Since Lorenz did not devote specific books or articles exclusively to this topic, one might be tempted to conclude that his views about maternal care were not central to his career. On the contrary, I believe they played an important part in his success, especially in the United States. His role as an expert on maternal care transcended the confines of academic meetings and was key to his prominent visibility in different disciplines and among the general public. As historian Gregg Mitman has shown, Lorenz’s and also Tinbergen’s focus on the family life of animals was the center of their films and media work.56 In the United States, many of Lorenz’s public appearances emphasized his expertise on “motherhood” in ducks and humans. Magazines and television programs also focused on his ability to substitute for a bird’s mother. His “mothering” abilities made him appealing to large audiences. When Life magazine run a story about (p.65) (p.66) Lorenz in 1955, for instance, the title presented him in his most popular role, as “An Adopted Mother Goose” (see fig. 2.4).57

The Study of Instincts

Figure 2.4. Lorenz as “An Adopted Mother Goose.” The caption of the picture in Life Magazine reads: “Standing chest-deep in water on his Westphalia preserve, Dr. Lorenz assembles his family of goslings by making noises like mother goose.” Life Magazine 39 (July/August 1955):73. Photo by Thomas D. McAvoy. TIME & LIFE Images. Courtesy of Getty Images.

As an adopted mother, Lorenz enjoyed a privileged position from which to observe the social life of animals. In his best-selling King Solomon’s Ring, Lorenz detailed his role as a devoted “foster mother” of jackdaws, ducklings, and goslings. His success in this role conferred authority on his views concerning maternal care and even infant development. Thanks to his ability to “talk to the beasts, the birds, and the fish,” Lorenz, like King Solomon, could then share the animals’ wisdom about mother love. His lessons seemed to come straight from nature, right from the goose’s mouth.

At the same time, Lorenz used his intimate knowledge of animal family life to extrapolate from birds to humans. This extrapolation was especially successful in the case of his work with geese. As Klaus Taschwer has put it, geese were Lorenz’s emblematic animal.58 Lorenz’s best-known images show him swimming with or being followed by geese. Observers could readily see him as part of their family life. In a fascinating examination of his changing representations of his “goose child Martina,” Tania Munz has shown how Lorenz used geese, and specifically Martina, to instantiate different—and sometimes contradictory—views over the span of his career.59 Lorenz literally attributed to Martina those characteristics and pathologies that he believed were typical of animal and human behavior. In this sense, when Lorenz talked about animals, he was often talking about humans as well.

Lorenz used several strategies to achieve this identification of humans with animals, especially geese. First, he established the equivalency of humans and animals at the emotional level. He was fond of citing his mentor Heinroth’s words: “I regard animals as very emotional people with very little intelligence!”60 He also often emphasized that greylag geese exhibited social and especially family behaviors similar to those of humans. The strong pair bond leading to monogamy, the healthy family, the coy female that did not engage in indiscriminate mating: these were all features of the healthy family that Lorenz saw as the fundamental pillar of a stable society, in geese and in humans.

In addition, Lorenz used numerous rhetorical strategies to promote the identification of humans with his animals. He gave the animals he raised human names, like Martina. Sometimes he told the life stories of individual animals, like the greylag goose Martina and the jackdaw Tshock. These biographies were built around the moments of significance in a human life—birth, attachment to the mother, falling in love—and included (p.67) all the dramatic elements of a human life: jealousy, overcoming obstacles, faithfulness, courage, and so forth. The animals played humanlike roles: parent, groom, bride, sibling, or child. They lived humanlike lives, and a human playing the role of an animal (Lorenz) narrated their ad-ventures. For Lorenz, the human-animal boundary was highly permeable.

All those elements helped readers empathize with the geese, and they also provided evidence for Lorenz’s contention that, emotionally, humans are like animals. If we are like animals, for him that also meant that much of our emotional equipment is inborn and much of our social behavior is determined by instincts. As I said earlier, Lorenz had already stated in his early writings that social behavior in higher animals is to a large extent determined by instincts. This bold assertion was made explicit and more appealing by illustrating how those instincts work in the lives of animals.

Lost in translation between the scientist and the foster mother, between the geese and the humans, was an important aspect of imprinting that remained hidden in many subsequent discussions. Birds do not imprint on an individual, but on the species through an individual. Lorenz’s mentor Heinroth, who first highlighted the phenomenon, always talked about how geese imprint on humans and then see humans as their parents. Lorenz also wrote that imprinting provides the image of the species to the animal imprinted. However, he always emphasized his personal relationship with a bird imprinted specifically on himself. When Lorenz wrote about those birds that followed him, he always noted that they considered him their mother. For example, as Munz has noted, he presented his most famous goose, Martina, as “my goose child.”61

Lorenz’s emphasis on his particular relationship with a bird imprinted on him helped to make imprinting in animals parallel to the human mother-infant dyad. In this sense, it facilitated the extrapolation of imprinting from animals to humans. To a great extent, I believe, this displacement from the species to the individual made it easier for psychoanalysts like Bowlby to see imprinting as analogous to the relationship of human children to their mothers.


Konrad Lorenz and Niko Tinbergen presented ethology as a new science that would focus on the biological study of instincts. They argued that human social behavior, like the social behavior of other animals, is guided by instincts. After World War II, at a time of heightened scientific and (p.68) public interest in the development of emotions and the mother-child relationship, Lorenz’s studies of imprinting and animal social life received much publicity. For many psychoanalysts and psychologists, including John Bowlby, several psychopathologies and sociopathologies could be traced back to the disruption of the mother-infant bond. To them, Lorenz’s work on imprinting seemed relevant to understanding the nature of early relationships.

Here I have shown how Lorenz extrapolated his views about animals to humans and, specifically, endorsed the relevance of imprinting for understanding child development. Encouraged by Bowlby and other child researchers’ eager reception of his views, Lorenz increasingly presented himself as an expert on the mother-child dyad. At least in the United States, Lorenz’s success was closely tied to his role as “an adopted mother goose.”

During the 1950s many child analysts, in turn, would use Lorenz’s ideas to support their own views. One of the most prominent was Bowlby, who—with Lorenz’s endorsement—gradually turned to animal research to back his view that the mother is the child’s psychic organizer and in 1958 proposed a synthesis of ethology and psychoanalysis to explain children’s attachment to their mothers.


(1) . John Bowlby, Child Care and the Growth of Love, ed. Margery Fry (Harmondsworth, UK: Pelican, 1953), 15. Bowlby included “Lorenz, K.” in the “List of Authorities referred to but not named” on p. 183.

(2) . The most complete history of ethology is Richard W. Burkhardt, Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology (Chicago: University of Chicago Press, 2005); see also John R. Durant, “Innate Character in Animals and Man: A Perspective on the Origins of Ethology,” in Biology, Medicine and Society, 1840–1940, ed. Charles Webster (Cambridge: Cambridge University Press, 1981), 157–92; Donald A. Dewsbury, Comparative Psychology in the Twentieth Century (Stroudsburg, PA: Hutchinson Ross, 1984); Richard W. Burkhardt Jr., “The Founders of Ethology and the Problem of Human Aggression: A Study in Ethologists’ Ecologies,” in The Animal/Human Boundary: Historical Perspectives, ed. Angela N. H. Creager and William Chester Jordan (Rochester, NY: University of Rochester Press, 2002), 265–304. For the other authors, see further notes in this chapter.

(3) . On Lorenz’s life and work, see Klaus Taschwer and Benedikt Föger, Konrad (p.256) Lorenz: Biographie (Vienna: Paul Zsolnay, 2003). See also Burkhardt, Patterns of Behavior.

(4) . On Heinroth, see Burkhardt, Patterns of Behavior, 135ff., and Karl Bühler, Die Krise der Psychologie (Jena: Gustav Fischer, 1927). On Bühler, see James F. T. Bugental, ed., “Symposium on Karl Bühler’s Contributions to Psychology,” Journal of General Psychology 75, no. 2 (1966):181–219. On his influence on Lorenz, see Veronika Hofer, “Konrad Lorenz als Schüler von Karl Bühler: Diskussion der neu entdeckten Quellen zu den persönlichen und inhaltlichen Positionen zwischen Karl Bühler, Konrad Lorenz und Egon Brunswik,” Die Zeitgeschichte 28 (2001):135–59.

(5) . Lorenz to Heinroth, February 22, 1931; cited in Burkhardt, Patterns of Behavior, 141.

(6) . See Konrad Lorenz, “Beobachtungen an Dohlen,” Journal für Ornithologie 75 (1927):511–19; Lorenz, “Beiträge zur Ethologie sozialer Corviden,” Journal für Ornithologie 79 (1931):67–127; Lorenz, “Betrachtungen über das Erkennen der arteigenen Triebhandlungen der Vögel,” Journal für Ornithologie 80 (1932):50–98; Lorenz, “Beobachtungen an freifliegenden zahmgehaltenen Nachtreihern,” Journal für Ornithologie 82 (1934):160–61; and Lorenz, “Über die Bildung des Instink-tbegriffes,” Die Naturwissenschaften 25 (1937):289–300, 307–18, 324–31.

(7) . Konrad Lorenz, “Der Kumpan in der Umwelt des Vogels, der Artgenosse als auslösendes Moment sozialer Verhaltensweisen,” Journal für Ornithologie 83 (1935):137–215, 289–413. This article was published in three English translations, but only the last one is complete: Lorenz, “The Companion in the Bird’s World,” Auk 54 (1937):245–73; Lorenz, “Companionship in Bird Life: Fellow Members of the Species as Releasers of Social Behavior,” in Instinctive Behavior: The Development of a Modern Concept, ed. Claire H. Schiller (New York: International Universities Press, 1957), 83–128; and Lorenz, “Companions as Factors in the Bird’s Environment: The Conspecific as the Eliciting Factor for Social Behaviour Patterns,” in Lorenz, Studies in Animal and Human Behavior, vol. 1 (Cambridge, MA: Harvard University Press, 1970), 101–258 (this version will be cited here).

(8) . Lorenz borrowed the concepts of “companion” and “releaser” from the work of Jakob von Uexküll, who had incorporated some of Lorenz’s observations on the social behavior of jackdaws into his own theory of the perceptual worlds of animals. See Jacob von Uexküll, “A Stroll through the Worlds of Animals and Men: A Picture Book of Invisible Worlds,” in Schiller, Instinctive Behavior, 5–80. On Uexküll see Anne Harrington, Reenchanted Science: Holism in German Culture from Wilhelm II to Hitler (Princeton, NJ: Princeton University Press, 1996), and see Burkhardt, Patterns of Behavior, on early influences on Lorenz’s work.

(9) . Lorenz, “Companions as Factors,” 254, 258.

(10) . On Tinbergen, see Hans Kruuk, Niko’s Nature: A Life of Niko Tinbergen and His Science of Animal Behavior (Oxford: Oxford University Press, 2003); Burkhardt, Patterns of Behavior; D. R. Röell, The World of Instinct: Niko Tinbergen (p.257) and the Rise of Ethology in the Netherlands (1920–1950) (Assen, Netherlands: Van Gorcum, 2000). For Tinbergen’s reflections on his career, see Niko Tinbergen, “Watching and Wondering,” in Studying Animal Behavior: Autobiographies of the Founders, ed. Donald A. Dewsbury (Chicago: University of Chicago Press, 1989), 431–63. On the early collaborative work between Lorenz and Tinbergen in 1936–1937, see Burkhardt, Patterns of Behavior, 187–88, 205–13.

(11) . Konrad Lorenz and Niko Tinbergen, “Taxis and Instinctive Behaviour Pattern in Egg-Rolling by the Greylag Goose,” in Lorenz, Studies in Animal and Human Behaviour, 1:316–50; originally published as “Taxis und Instinkthandlung in der Eirollbewegung der Graugans,” Zeitschrift für Tierpsychologie 2 (1938):1–29.

(12) . Burkhardt, Patterns of Behavior, passim.

(13) . On the role of film in ethology, see Gregg Mitman, Reel Nature: America’s Romance with Wildlife on Film (Cambridge, MA: Harvard University Press, 1999), chap. 3; Tania Munz, “Die Ethologie des wissenschaftlichen Cineasten: Karl von Frisch, Konrad Lorenz und das Verhalten der Tiere im Film,” montage AV 14, no. 4 (2005):52–68.

(14) . Mitchell G. Ash, “Psychology and Politics in Interwar Vienna: The Vienna Psychological Institute, 1922–1942,” in Psychology in Twentieth-Century Thought and Society, ed. M. G. Ash and W. R. Woodward (Cambridge: Cambridge University Press, 1987), 157.

(15) . On Lorenz and National Socialism, see Burkhardt, Patterns of Behavior, chap. 5; Ute Deichmann, Biologists under Hitler (Cambridge, MA: Harvard University Press, 1996), 179–205; Benedikt Föger and Klaus Taschwer, Die andere Seite des Spiegels: Konrad Lorenz und der Nationalsozialismus (Vienna: Czernin, 2001); and Theodora Kalikow, “Konrad Lorenz’s Ethological Theory, 1939–1943: ‘Explanations’ of Human Thinking, Feeling, and Behaviour,” Philosophy of the Social Sciences 6 (1976):15–34.

(16) . Konrad Lorenz, “Über Ausfallserscheinungen im Instinktverhalten von Haustieren und ihre sozialpsychologische Bedeutung,” in Charakter und Erziehung: Bericht über den 16. Kongress der deutschen Gesellschaft für Psychologie in Bayreuth, ed. Otto Klemm (Leipzig: Johann Ambrosius Barth, 1939), 139–47; Konrad Lorenz, “Durch Domestikation verursachte Störungen arteigenen Verhaltens,” Zeitschrift für angewandte Psychologie und Charakterkunde 59, nos. 1–2 (1940):2–81.

(17) . Konrad Lorenz in Königsberger Allgemeine Zeitung, November 2, 1940. Cited in Föger und Taschwer, Die andere Seite, 134–35.

(18) . Konrad Lorenz, “Die angeborenen Formen möglicher Erfahrung,” Zeitschrift für Tierpsychologie 5 (1943):235–409.

(19) . Konrad Lorenz, King Solomon’s Ring: New Light on Animal Ways (New York: Crowell, 1952); originally published as Er redete mit dem Vieh, den Vögeln und den Fischen (Vienna: Borotha-Schoeler, 1949). In some editions the English (p.258) translation included a subtitle: He Spoke with the Beasts, the Birds, and the Fish. According to Tania Munz, the book went through some forty printings and was translated into twelve languages. See Tania Munz, “‘My Goose Child Martina’: The Multiple Uses of Geese in Konrad Lorenz’s Writings on Animals, 1935–1988,” Historical Studies in the Natural Sciences 41 (2011):431.

(20) . Konrad Lorenz, “The Comparative Method in Studying Innate Behaviour Patterns,” Symposia of the Society for Experimental Biology 4 (1950):221, 222, 234.

(21) . Ibid.., 265.

(22) . Niko Tinbergen, The Study of Instinct (Oxford: Clarendon Press, 1951), 1, 2.

(23) . On Craig’s work and on his influence on Lorenz, see Burkhardt, Patterns of Behavior, chaps. 1 and 3, respectively.

(24) . Lorenz, “Companions as Factors,” 250–51. On 250–52 Lorenz presents a short history of the concept of instinct, noting the sources of some of the elements he used for his own conception. See also Wallace Craig, “Appetites and Aversions as Constituents of Instincts,” Biological Bulletin 34 (1918):91–107. On Lorenz’s views about instincts, see Burkhardt, Patterns of Behavior; Theodora J. Kalikow, “History of Konrad Lorenz’s Ethological Theory, 1927–1939: The Role of Meta-theory, Theory, Anomaly and New Discoveries in a Scientific ‘Evolution,’” Studies in the History and Philosophy of Science 6 (1975):331–41; Robert Richards, “The Innate and the Learned: The Evolution of Konrad Lorenz’s Theory of Instinct,” Philosophy of the Social Sciences 4 (1974):111–33; Colin G. Beer, “Darwin, Instinct, and Ethology,” Journal of the History of the Behavioral Sciences 19 (1983):68–80; and Ingo Brigandt, “The Instinct Concept of the Early Konrad Lorenz,” Journal of the History of Biology 38 (2005):571–608.

(25) . Konrad Lorenz, “The Establishment of the Instinct Concept” (1937), in Lorenz, Studies in Animal and Human Behavior, 1:259–315.

(26) . Lorenz, “Comparative Method,” 255. Tinbergen’s model of motivation was different from Lorenz’s model. Tinbergen postulated a hierarchical system of centers. See Tinbergen, Study of Instinct, 125. Also see Lorenz, “The Past Twelve Years in the Comparative Study of Behavior” (1952), in Schiller, Instinctive Behavior, 288–310.

(27) . J. B. S. Haldane, “The Sources of Some Ethological Notions,” British Journal of Animal Behaviour 4 (1956):162–64. For a comparison of Lorenz’s and Mac-Dougall’s models, see Paul E. Griffiths, “Instinct in the ‘50s: The British Reception of Konrad Lorenz’s Theory of Instinctive Behavior,” Biology and Philosophy 19 (2004):609–31.

(28) . Lorenz, “Comparative Method,” 221.

(29) . Lorenz, “A Consideration of Methods of Identification of Species-Specific Instinctive Behaviour Patterns in Birds” (1932), in Lorenz, Studies in Animal and Human Behavior, 1:57–100; Lorenz, “Establishment of the Instinct Concept.”

(30) . Lorenz, “Consideration of Methods,” 65; Lorenz, “Companions as Factors,” 248.

(p.259) (31) . See Burkhardt, Patterns of Behavior, chap. 3.

(32) . Lorenz’s most extended discussion of deprivation experiments is found in “The Value and the Limitations of the Deprivation Experiment,” chap. 7 of his Evolution and Modification of Behavior (Chicago: University of Chicago Press, 1965), 83–100.

(33) . Lorenz, “Companions,” 126; emphasis in original.

(34) . Ibid., 245.

(35) . Ibid., 124ff.

(36) . Ibid., 168.

(37) . Ibid., 185.

(38) . Ibid., 244.

(39) . Lorenz, “Die angeborenen Formen,” 274; my translation.

(40) . Ibid., 276.

(41) . Konrad Lorenz, “Companions,” 251; Lorenz, “Comparative Method,” 263; William James, Principles of Psychology (1890; reprint, New York: Holt, 1900), 442; John B. Watson, Psychology from the Standpoint of a Behaviorist (Philadelphia: Lippincott, 1919), 231; and William McDougall, An Introduction to Social Psychology (1908; reprint, Boston: Luce, 1916), 29.

(42) . Lorenz, “Comparative Method,” 265.

(43) . Konrad Lorenz, “Über angeborene Instinktformeln beim Menschen,” Deutsche medizinische Wochenschrift 78 (1953):1566–69, 1600–1604.

(44) . The verbatim transcriptions of the proceedings of the study group meetings were published by the Tavistock Institute in 1956 (both the 1953 and the 1954 meetings), 1958, and 1960. They were later collected in a single volume: J. M. Tanner and Barbel Inhelder, eds., Discussions on Child Development (London: Tavistock, 1971), but with separate pagination for each original volume. All citations here refer to this edition.

(45) . Bowlby, in Tanner and Inhelder, Discussions on Child Development, 1:27.

(46) . Konrad Lorenz to Erwin Stresemann, May 14, 1934; quoted by Föger and Taschwer, Andere Seite, 55–56.

(47) . Konrad Lorenz, “Memorandum on Ethology,” January 7, 1953, PP/BOW/H.132, John Bowlby Papers, Western Manuscripts and Archives, Wellcome Library, London; Lorenz, in Tanner and Inhelder, Discussions on Child Development, 1:117.

(48) . Lorenz, in Tanner and Inhelder, Discussions on Child Development, 1:215–16, quotation on 211.

(49) . Ibid., 222.

(50) . Ibid., 223.

(51) . See Tinbergen, Study of Instinct, 45. Tinbergen describes other examples of “supernormal” stimuli on 44–45.

(52) . Leta S. Hollingsworth, “Social Devices for Impelling Women to Bear and Rear Children,” American Journal of Sociology 22 (1916):19–29; Ruth Reed, (p.260) “Changing Conceptions of the Maternal Instinct,” Journal of Abnormal Psychology and Social Psychology 18 (1923):78–87. On the maternal instinct, see Marga Vicedo, “Mother Love and Human Nature: A History of the Maternal Instinct” (PhD diss., Harvard University, 2005).

(53) . Lorenz, in Tanner and Inhelder, Discussions on Child Development, 1:227–28.

(54) . Mead, in Ibid., 1:228.

(55) . Lorenz in Tanner and Inhelder, Discussions on Child Development, 3:36, 69; see also 45.

(56) . Mitman, Reel Nature.

(57) . “An Adopted Mother Goose: Filling a Parent’s Role, a Scientist Studies Goslings’ Behavior,” Life Magazine 39 (July/August 1955):73–74, 77–78. For an analysis of Lorenz’s use of his role as “mother” to study animals, see Vicedo, “Outside or Inside the Animal?”

(58) . For an analysis of Lorenz’s connection with greylag geese, see Klaus Taschwer, “Von Gänsen und Menschen: Über die Geschichte der Ethologie in Österreich und über ihren Protagonisten, den Forscher, Popularisator und Ökopolitiker Konrad Lorenz,” in Wissenschaft, Politik und Öffentlichkeit, von der Wiener Moderne bis zur Gegenwart, ed. Mitchell G. Ash and Christian H. Stifter (Vienna: WUV, 2002), 331–51.

(59) . Munz, “‘My Goose Child Martina.’“

(60) . Konrad Lorenz, On Agression (1966; reprint, New York: Bantam, 1969), 203.

(61) . Munz, “‘My Goose Child Martina,’” 432.