Sensory Physiology and the Return of the Animal Mind in the Career of Donald Redfield Griffin, 1934-1986
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SENSORY PHYSIOLOGY AND THE RETURN OF THE ANIMAL MIND IN THE CAREER OF DONALD REDFIELD GRIFFIN, 1934-1986 by Richard Stephen Nash A dissertation submitted to Johns Hopkins University in conformity with the requirements for the degree of Doctor of Philosophy Baltimore, Maryland January, 2016 © 2016 Richard Stephen Nash All Rights Reserved ABSTRACT Donald Redfield Griffin (1915-2003) was an American zoologist best known for his discovery of echolocation and for his later work on animal consciousness. He was a central figure in behavioral biology and sensory physiology in the United States, and he made important contributions to the disciplinary and intellectual development of animal behavior research in the second half of the twentieth century. During his early career, he focused on the sensory physiology of animal navigation. Along with fellow Harvard graduate student Robert Galambos (1914-2010), in the late-1930s Griffin discovered the ultrasonic method of orientation in bats; in 1944 he coined the term “echolocation” to describe this phenomenon as a general method of perception. In addition to his discovery of echolocation, Griffin also made several contributions to understanding the physiological basis of bird migration and navigation, and he popularized zoologist Karl von Frisch’s (1886-1982) dance language theory of the honeybee in the United States. In 1976, Griffin surprised the scientific world by raising the question of animal consciousness, a taboo in professional science for most of the twentieth century. Although the animal mind was of central importance in post-Darwinian biology, the onset of behaviorism and mechanistic conceptions of behavior in the twentieth century relegated such inquiry to the dustbin of pseudoscience and amateurism. Beginning with his provocative book, The Question of Animal Awareness (1976), Griffin devoted the second phase of his career to making animal consciousness a scientifically respectable topic once again. Here again he made significant contributions to the study of animal behavior by establishing a new field of science, cognitive ethology, which is centered on ii the evolutionary and comparative analysis of consciousness and cognition in animal behavior. Doctoral Advisor: Sharon Kingsland, PhD Thesis Committee: Daniel Todes, PhD Robert Kargon, PhD Angus Burgin, PhD Nathaniel Comfort, PhD iii ACKNOWLEDGMENTS First and foremost, I am happy to thank my advisor Sharon Kingsland for her patience, criticism, and guidance of my project. Without her keen insight and close reading, I could never have completed this dissertation. Dan Todes has similarly been a magic well of intellectual support as he has challenged and inspired me to make this work my own. I also thank Angus Burgin, a tremendous influence in my approach to intellectual history and historical scholarship in general. Finally, I thank Bob Kargon and Nathaniel Comfort for their unique insights, and for their close reading and criticism of my dissertation. My experience in the Johns Hopkins’ History of Science, Technology, and Medicine program has been incredibly fulfilling and edifying. The wide-ranging intellect of the faculty and of my fellow graduate students has shown me the true meaning of scholarship and collegiality. I would like to thank in particular Graham Mooney, Larry Principe, Maria Portuondo, Bill Leslie, Yulia Frumer, and Michael Dennis for their various contributions to my intellectual development. In addition I am happy to thank Kathy Olesko for first inspiring me to study the history of science when I was a naïve undergraduate at Georgetown University. Portions of my research were made possible by a generous grant from the Rockefeller Foundation’s Archive Center. In addition, my research has been crucially assisted by several excellent archivists and librarians, including Bethany J. Antos (Rockefeller Archive Center), Christine Ruggere (JHU), Kelly Spring (JHU), Jim Stimpert (JHU), Sandra Jackson and ILL Staff (JHU), and Lawrence Marcus (Library of Congress). I am forever grateful for your knowledge and assistance. iv Several graduate students and colleagues have supported me in countless ways over the past several years. It has been a great pleasure to share ideas, questions, doubts, and drinks with Ada Link, Jean-Olivier Richard, Layne Karafantis, Yixian Li, Pen Hardy, Julia Cummiskey, Eli Anders, Emily Margolis, Emilie Raymer, Matt Franco, Kirsten Moore, Simon Thode, Todd Christopher, Rex, Tom Berez, Tulley Long, Nick Radburn, and Kat Smoak. I owe a further debt of gratitude to my friends and colleagues in the JAS- Bio community. I also thank my great friends for their significant roles in my life, especially Blake and John Lennon, Tom Williams, and Kartik Venguswamy. I apologize to the many other colleagues and friends who deserve special recognition; it is unfortunate that acknowledgments are written toward the end of our intellectual endeavors, when our minds gurgle and lurch toward clarity. Finally, I thank my family. Aimee Hess, my beautiful wife, has been a most incredibly supportive and devoted advocate. I thank her especially for generously reading several drafts, offering keen criticism and editorial advice over the duration of this long project. That Aimee has recently brought my lovely daughter Ella into this world is merely icing atop an otherwise scrumptious cake. I also thank my parents, Rick Nash and Kelly Covey, my in-laws the Hess family, and my siblings, Paige, Coleman, and Kenneth, for their encouragement and understanding as I have wandered into the bizarre world of academia. v TABLE OF CONTENTS Abstract…………………….…………………………………………………………...…ii Acknowledgments……………………….……………………………………………….iv CHAPTER 1: Donald Griffin and the Mechanistic Character of American Biology……..1 CHAPTER 2: Obstacle Avoidance and the Mechanistic Conception of Bats…………...21 CHAPTER 3: The Wartime Discovery of Echolocation………………………………...65 CHAPTER 4: The Trouble with Bird Migration……………………………………….120 CHAPTER 5: Birds, Bats, and Bees: The Complexity of Animal Behavior...................175 CHAPTER 6: Cognitive Dissidence and the Return of the Animal Mind……………..218 BIBLIOGRAPHY……………....…………………………………………………..…..256 CURRICULUM VITAE………….……………………………………………….……272 vi CHAPTER ONE Donald Griffin and the Mechanistic Character of Twentieth-Century American Biology Donald Redfield Griffin (1915-2003) was an American biologist and leader in advancing the study of animal behavior, sensory physiology, and physiological ecology in the mid-to-late-twentieth century. He received his Ph.D. from Harvard University in 1942 and went on to a distinguished career at Cornell University, Harvard University, and finally Rockefeller University, from which he retired in 1986. Retirement did not end his scientific research, and he continued to study the behavior of birds and mammals throughout his life. The scientific work for which he is best known was the discovery that bats navigated their environments by emitting ultrasonic sounds and analyzing their echoes, a method that Griffin dubbed “echolocation” in 1944. This work, initially conducted in collaboration with Robert Galambos (1914-2010) at Harvard, solved the centuries-old problem of how bats sensed their environments without relying on vision. Griffin also conducted important research on bird migration and homing behavior, another longstanding biological problem that had drawn the attention of many researchers. In this work he developed innovative methods to study these problems and clarified many of the key issues involved. In tandem with other ornithologists, he advanced the celestial theory of navigation by showing that many migratory species used the motion of the sun and stars to orient themselves homeward from unfamiliar territory. Griffin’s colleagues admired his research for its rigor, and for the care he took to base conclusions about animal behavior and perception on scientific evidence derived from laboratory experiments and field observations. It came as a surprise therefore when his colleagues heard him raise a formerly taboo subject – the thorny question of animal consciousness – at a scientific conference in 1975, following it up with a book in 1976, The Question of Animal Awareness.1 The idea that animals possessed anything like human awareness or consciousness, although important in Darwinian biology and psychology in the nineteenth century, had long since been relegated to the dustbin of pseudoscience and amateurism. Griffin himself had been trained at Harvard to eschew anything but a mechanistic approach to behavior that avoided any reference to the animal mind. Yet thirty-five years later he was emboldened to raise this problem and to challenge his colleagues to take animal consciousness seriously as an object of scientific inquiry. This dissertation explores how this change occurred in Griffin’s thought. I will argue that, contrary to his own statements that the intellectual transition to animal consciousness was relatively abrupt, it was in fact a gradual but logical development of his increasingly sophisticated understanding of how animals navigated and perceived their environments. In reopening the problem of animal consciousness, Griffin was not departing from his earlier habit of thought but was carrying his life’s work to a logical endpoint, to the seemingly strange idea that scientists needed to think harder about what it was really like to be a bird or a bat. By pursuing such questions, Griffin concluded that the subject of animal consciousness had to be raised