Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology

by Richard W. Burkhardt Jr.

Overview

It is hard to imagine, by their very name, the life sciences not involving the study of living things, but until the twentieth century much of what was known in the field was based primarily on specimens that had long before taken their last breaths. Only in the last century has ethology--the study of animal behavior--emerged as a major field of the life sciences.

In Patterns of Behavior, Richard W. Burkhardt Jr. traces the scientific theories, practices, subjects, and settings integral to the construction of a discipline pivotal to our understanding of the diversity of life. Central to this tale are Konrad Lorenz and Niko Tinbergen, 1973 Nobel laureates whose research helped legitimize the field of ethology and bring international attention to the culture of behavioral research. Demonstrating how matters of practice, politics, and place all shaped "ethology's ecologies," Burkhardt's book offers a sensitive reading of the complex interplay of the field's celebrated pioneers and a richly textured reconstruction of ethology's transformation from a quiet backwater of natural history to the forefront of the biological sciences. Winner of the 2006 Pfizer Awad from the History of Science Society

Product Details

ISBN-13:	9780226080901
Publisher:	University of Chicago Press
Publication date:	03/15/2005
Edition description:	1
Pages:	648
Product dimensions:	6.00(w) x 9.00(h) x 1.60(d)

About the Author

Richard W. Burkhardt Jr. is professor of history at the University of Illinois at Urbana-Champaign. He is the author of The Spirit of System: Lamarck and Evolutionary Biology.

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PATTERNS OF BEHAVIOR
KONRAD LORENZ, NIKO TINBERGEN, AND THE FOUNDING OF ETHOLOGY

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By Richard W. Burkhardt, Jr.
The University of Chicago Press
Copyright © 2005 The University of Chicago
All right reserved.
ISBN: 978-0-226-08090-1

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Chapter One
CHARLES OTIS WHITMAN, WALLACE CRAIG, AND THE BIOLOGICAL STUDY OF ANIMAL BEHAVIOR IN AMERICA

Instinct and structure are to be studied from the common standpoint of phyletic descent. C. O. WHITMAN, "ANIMAL BEHAVIOR," 1898

I have about decided that comparative psychology is the line for me. Whether there will be bread and butter in it or not, I don't know-that's a subordinate question. WALLACE CRAIG TO C. C. ADAMS, 1898

In the summer of 1898, at the Marine Biological Laboratory in Woods Hole, Massachusetts, the American zoologist Charles Otis Whitman delivered a lecture containing the insight that Konrad Lorenz would later call the "Archimedean point" of comparative behavior studies. As Whitman put it: "Instinct and structure are to be studied from the common standpoint of phyletic descent." In other words, instinctive behavior patterns could be used just like body parts in studying how animals evolved.

One could write a history of ethology beginning with other investigators in other settings, but Whitman in the United States is as useful a starting point as any other would be. His case opens up a wide range of questions involving theories and practices, actors and objects, allies and competitors, boundaries and bridges, local circumstances and broader horizons. All of these relate in turn to the issues of who could speak authoritatively about animal life and how to study it.

The actors in animal behavior studies in Whitman's day constituted a stunningly diverse fauna, stirring in marvelous ways. Single-celled organisms moved silently toward or away from the experimenter's chosen stimuli. So-called higher animals performed more complicated motions, often accompanied by buzzing, cooing, clucking, or other ways of signaling their presence. Then too there were the human observers of the animals' behavior-watching, wondering, poking, probing, controlling, conversing, writing, and in general constructing systems of relations connecting themselves with their animal subjects and with the additional material, institutional, and social necessities pertaining to their scientific lives. The roster of investigators of animal behavior in the United States included Whitman and his behaviorally oriented students, William Morton Wheeler, Samuel J. Holmes, Oscar Riddle, and Wallace Craig; Jacques Loeb and Herbert Spencer Jennings, paired in history by their famous debate over tropisms and the behavior of lower organisms; Charles H. Turner, the African-American biologist whose experimental investigations of color sense and form sense in bees predated Karl von Frisch's earliest work on these topics; Francis H. Herrick, a pioneer in the study of the domestic life of birds; and C. C. Adams and Victor Shelford, both animal ecologists. Then too there were Margaret F. Washburn, Robert M. Yerkes, and John Broadus Watson, psychologists whose work, at least in the early part of their careers, focused primarily on animal behavior. Beginning in 1911, the United States could also claim the Journal of Animal Behavior, the first major scientific journal devoted exclusively to the study of animal behavior.

These various individuals did not constitute a close-knit community of investigators, sharing a common view of the aims and methods of animal behavior study. The biologists and psychologists, not surprisingly, tended to differ from each other in their ideas and their practices. Yet the lines that separated them were by no means hard and fast. Certain biologists and psychologists, at particular moments of their careers, felt more affinity with each other than they did with members of their own disciplines. Instructively enough, though, none of them sought to establish the study of animal behavior as an independent discipline. Each viewed the study of animal behavior as a means by which his or her own parent discipline could be reformed and developed. Each wanted to make animal behavior studies an integral part of the broader, ongoing enterprise that was either biology or psychology.

This was particularly true of Whitman. Whitman was an eloquent proponent of the view that specialized studies should be pursued only as parts of the organized whole, which in this case meant biology. He put this view into practice in his organization of the Division of Biology at the University of Chicago. He likewise articulated this view in his presidential address to the Society of American Naturalists in 1897, urging: "We need to get more deeply saturated with the meaning of the word 'biological,' and to keep renewing our faith in it as a governing conception. Our centrifugal specialties have no justification except in the ensemble." He encapsulated the same vision in his 1902 call for a special experiment station-a "bio logical farm"-where "the study of life-histories, habits, instincts and intelligence" would be conducted alongside "the experimental investigation of heredity, variation, and evolution."

To Whitman it was apparent that the success of biology as a governing conception would depend not simply on the idea's intellectual appeal but also on the establishment of the attitudes, practices, and institutions necessary to sustain it. He called the approach he championed "experimental natural history." He specifically distinguished this from the narrower, physiological, lab-oriented approach of his colleague Jacques Loeb. In Loeb's approach, Whitman complained, "instinct reduces itself in the last analysis to heliotropism, stereotropism, and the like" and "the whole course of evolution drops out of sight altogether."

In the new, university zoology laboratories that symbolized modern biology across the country, investigators took up Loeb's tropism studies with enthusiasm. Whitman himself, however, did not believe the laboratory to be a sufficiently ample setting for promoting biology as properly conceived. Nor did he believe the problem could be solved simply be supplementing lab studies with fieldwork. He envisioned his "biological farm" as the special setting where experimental natural history would be pursued and the "governing concept" of biology would be ratified.

This chapter focuses primarily on the work and careers of Charles Otis Whitman and his student Wallace Craig. Their stories illuminate the problems of pursuing naturalistic, experimental, and evolutionary studies of behavior at a time when the institutional support for such studies was yet to be created and when other forms of research were proving more successful in commandeering the resources and approbation of the American scientific community. They were not the only investigators in the United States in the early twentieth century who pursued animal behavior studies. They were, however, the scientists who contributed most directly to the particular approach to animal behavior studies that came to be known as ethology.

CHARLES OTIS WHITMAN AND AMERICAN BIOLOGY

When Charles Otis Whitman delivered his lecture on "animal behavior" at the Woods Hole Marine Biological Laboratory in 1898, he was arguably the most influential biologist in America. The MBL was the Mecca of American biology, the seaside magnet that in the summer months drew America's finest biologists to a single, vital center, and Whitman was its first director. He also headed the Division of Biology and the Department of Zoology at the University of Chicago, where he had assembled as distinguished a biological research faculty as could be found anywhere in the country. Before going to Chicago in 1890 he had served for three years as the first chair of the zoology department at Clark University. In addition, he had founded the Journal of Morphology (in 1887) and been the key figure in establishing in 1890 the American Morphological Society (later to be renamed the American Society of Zoologists). The range of his interests and expertise was unmatched among his zoologist or biologist colleagues in America. How he came to his interest in animal behavior and how that interest fitted with the rest of his concerns and career deserve careful attention.

Ethologists have routinely reported that their fascination with animal life began in childhood, long before they had any inkling of what it might mean to be a scientist. Whitman's own case fits the model. Born in 1842, Whitman as a youth was an ardent bird collector. He distinguished himself by his skill in mounting the specimens he shot, but he also kept birds-and amphibians and mammals too-as pets. Among his live birds were pigeons. As he later reported to his student Wallace Craig, his pigeons fascinated him, and he "sat and watched them by the hour, intensely interested in their feeding, their young, and in everything that they did."

The historian Philip J. Pauly has described how Whitman, the son of Adventist parents in rural, western Maine, broke with his parents' extreme religious views and found his way first to Universalism and then to the natural sciences. Pauly argues that the intensity of the religious conflict of Whitman's youth was reflected in the strength of his later attachment to the idea of a law-governed, progressively developing universe, where supernatural events had no place. Whereas Whitman's parents believed that the Second Coming was imminent, Whitman put his own faith in modern science. He embraced a worldview in which gradual, progressive change and the evolution of life itself were the central processes and were to be explained in material rather than miraculous or mystical terms.

After working his way through Bowdoin College, Whitman taught high school in Massachusetts. In 1873 he was attracted to the natural history summer school instituted by the famed Harvard zoologist Louis Agassiz on Penikese Island in Buzzards' Bay. There, at the sea's edge, with living marine creatures directly at hand, Whitman got his first real taste of zoology. This led him to join the Boston Society of Natural History, to spend another summer at Penikese, and then to follow the example of a fellow Penikese student by going to Leipzig and studying zoology under the German parasitologist Rudolf Leuckart. It was under Leuckart at Leipzig that he was molded into a serious researcher.

In Leuckart's laboratory, Whitman mastered the latest techniques of microscopic analysis, particularly as adapted to the study of fertilization and development. For his doctoral thesis, he investigated the early embryonic development of the fish leech, Clepsine. His dissertation, published in 1878, reflected his newly acquired technical expertise, his command of contemporary science, and his commitment to a special vision of organic development. "In the fecundated egg," he wrote, "slumbers potentially the future embryo. While we cannot say that the embryo is predelineated, we can say that it is predetermined. The 'Histogenetic sundering' of embryonic elements begins with the cleavage, and every step in the process bears a definite and invariable relation to antecedent and subsequent steps." This recognition of the importance of tracing cells, structures, and even behavior patterns back to their very earliest origins became a hallmark of Whitman's work.

Over the course of the next two decades, Whitman moved often. He worked at Harvard, the Imperial University of Tokyo, the Zoological Station at Naples, the Allis Lake Laboratory (near Milwaukee, Wisconsin), Clark University, and then finally the University of Chicago and the MBL. Over the same period, his investigations of leeches grew to encompass these organisms in all aspects of their existence from their development to their systematics, phylogeny, ecology, and behavior. This came to be his model of proper biological research: the study of a single species or a group of closely related species exhaustively pursued in every aspect of its existence. In 1899 he defined biology as "the life-histories of animals, from the primordial germ-cell to the end of the life-cycle; their daily, periodical, and seasonal routines; their habits, instincts, intelligence, and peculiarities of behavior under varying conditions; their geographical distribution, genetic relations and oecological interrelations; their physiological activities, individually and collectively; their variations, adaptations, breeding and crossing." Whitman's focus on the living animal in every feature of its existence is epitomized in the claim that his students greeted each other not with the question "What is your special field?" but rather "What is your beast?"

In Whitman's animal behavior paper of 1898, his own special "beasts" featured prominently. Clepsine, the fish leech, was there. So too was Necturus, the freshwater salamander he began studying when he directed the Allis Lake Laboratory from 1886 to 1889. There also were his pigeons, the birds that had fascinated him as a boy and that he began studying in earnest again around 1895. They would be the almost constant focus of his research for the last decade and a half of his life.

Whitman presented his animal behavior paper in the form of two evening lectures in the Marine Biological Laboratory's summer lecture series of 1898. Here he set forth his views on the proper methods of studying behavior, the nature of instinct, the importance of studying instinct from a phylogenetic standpoint, the means by which behavior has evolved, and the relations-both ontogenetic and phylogenetic-between instinct and intelligence. Of this paper his student Frank R. Lillie later wrote: "No other of [Whitman's] papers illustrates better the qualities of his genius: the selection of a fundamental problem; painstaking study; publication only after years of observation and reflection; skill in laying bare the simple basis of an apparently complex group of phenomena; a grasp of the subject in all its bearings; and the use of the comparative or phyletic method of attack."

Whitman began his lecture with a few remarks on the subject of animal behavior generally. He then turned abruptly to the special topic of "modes of keeping quiet" among animals. Clepsine allowed him to make the point that if one wanted to understand the behavior of any animal species, one needed to have a thorough knowledge of the animal's entire behavioral repertoire. The "deceptive quiet" of Clepsine was such that an observer unfamiliar with Clepsine's habits would "almost certainly" draw the wrong conclusions about its sensitivity to stimuli. He drew the same lesson from his work with Necturus. It had taken him considerable experience with Necturus adults, he said-and two whole seasons rearing Necturus young-before he appreciated "the extreme timidity of these animals." This timidity, he explained, is "so deep-seated and persistent that one can form only a poor idea of it without considerable actual contact with it." Whitman underscored the importance of studying animal behavior under natural conditions. It was essential, he said, that one "observe and experiment under conditions that ensure free behavior."

Whitman paid particular attention in his lecture to instinctive behavior and to how instinct and intelligence were related. For him, the "first criterion of instinct" was that it could "be performed by the animal without learning by experience, instruction, or imitation." The fact that an animal's acts were adapted to purposeful ends, he explained, was not in itself proof of intelligence on the animal's part. Necturus always sneaked up on its prey, regardless of whether the prey was living or was simply a piece of meat. The creature, Whitman was convinced, had not the slightest appreciation of the importance of stealth. It was "quite blind" to the significance of its actions. Its movements, in Whitman's words, were "those characteristic of the species, not because they are measured and adapted to a definite end by intelligent experience, but because they are organically determined; in other words, depend essentially upon a specific organization." If Necturus, as a species, had to depend on its intelligence, he said, it was "difficult to see how it could escape immediate extinction." Its continued existence was assured by its instincts.

(Continues...)

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All rights reserved. No part of this excerpt may be reproduced or reprinted without permission in writing from the publisher.
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Table of Contents

Acknowledgments
Introduction
Theory, Practice, and Place in the Study of Animal Behavior
1. Charles Otis Whitman, Wallace Craig, and the Biological Study of Animal Behavior in America
2. British Field Studies of Behavior: Selous, Howard, Kirkman, and Huxley
3. Konrad Lorenz and the Conceptual Foundations of Ethology
4. Niko Tinbergen and the Lorenzian Program
5. Lorenz and National Socialism
6. The Postwar Reconstruction of Ethology
7. Ethology's New Settings
8. Attracting Attention
9. Tinbergen's Vision for Ethology
10. Conclusion: Ethology's Ecologies
Notes
Bibliography
Index

Editorial Reviews

"Burkhardt has written the definitive history of ethology. He has blended science and situation into a coherent and instructuve tale."

BJHS - Donald A. Dewsbury

"This monumental volume has been 26 years in the making and represents a mountainous effort of scholarship. . . . Burkhardt, a historian, makes a massive contribution to our knowledge of the development of animal behavior science. . . . A fascinating read, and a massive monument to the science of ethology."

Journal of the History of Behavioral Sciences - Hans Kruuk

"A fascinating and often entertaining account of the life and work of some of ethology’s key figures. Burkhardt has done a tremendous job, meticulously analysing and describing the rise of ethology. . . . In order to study the genomic or neural mechanisms of bevaviour, we need to know how behaviour works, and for that an ethological analysis is crucial. This wonderful book shows very clearly how early ethologists made such analysis possible."

Nature - Johan J. Bolhuis

"Patterns of Behavior offers more than a nuanced history of the origins of ethology. The juxtaposition of the stories of Tinbergen
and Lorenz provides an intriguing case study of how the advancement of scientific fields depends as much on the management of the
social relations among scientists as on intellectual brilliance and institution-building."

Science - Sharon Kingsland

"At last, a really excellent and complete account of ethology's checkered history. What emerges in Patterns of Behavior is a vivid demonstration that science is a human activity whose progress is influenced both by personality and by cultural context. This is a most distinguished contribution to the history of science."

Aubrey Manning

"This long, detailed, massively documented history of the origins of ethology . . . is a labor of love by a master historian of science. . . . Essential reading for anyone interested in the history of science in the twentieth century."

Alfred Kellyn Historical Review

"This excellent book reads like a novel. . . . The author reconstructs the tortuous history of the struggle for the creation of the discipline of ethology via intriguing biographies and against the backdrop of global history. . . . This work will remain as the reference for complex relationships between theoretical development and fieldwork during the 20th century, which is to say that the book clarifies our understanding as well of the current situation of the sister science of anthropology."

History of the Philosophy of Life Science - Marcel Otte

"This is a masterpiece. Twenty years in the making, this is the definitive account of the development of ethology. Beautifully written and immensely rich in detail, Patterns of Behavior is a wonderful reconstruction of the lives of the two founders of ethology, Niko Tinbergen and Konrad Lorenz . . . Patterns of Behavior is essential reading for anyone who teaches or conducts research in animal behaviour, anyone interested in the history of science, or those who simply want to discover how two very different personalities
created an entirely new way of looking at animals."

Animal Behaviour - T. R. Birkhead

"Not knowing one's roots is a common mistake in the early development of scientific disciplines, a problem that Burkhardt's thoughtful, comprehensive and readable book should help to resolve. It should be read by all animal behaviour researchers to help put their own efforts into perspective."

TREE - Michael Taborsky

"I thoroughly enjoyed reading this book. . . . I am convinced that  knowing the history of one's field makes for better instructors and so recommend that all students of aniumal behaviour read this book and most notably all those who teach it."

Ecoscience - Luc-Alain Giraldeau

"The most authoritative history of the field of ethology... A remarkable achievement. It brings together an immense amount of material in a scholarly, yet very readable, format. It also catches the enthusiasm of the participants in putting forward new ways of thinking about old problems, as well as some of the more acrimonious debates about particular issues."

Ethology - Jerry A. Hogan

"Burkhardt beautifully describes both the human story and the science that has come out of it. An excellent book."

Biologist - Colin Tudge

"This long, detailed, massively documented history of the origins of ethology . . . is a labor of love by a master historian of science. . . . Essential reading for anyone interested in the history of science in the twentieth century."

American Historical Review - Alfred Kelly