Second Nature: Domestication As Experiment and Metaphor in 20Th Century American Psychobiology

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AN ABSTRACT OF THE DISSERTATION OF

Nicholas Blanchard for the degree of Doctor of Philosophy in History of Science presented on August 7, 2014.

Title: Second Nature: Domestication as Experiment and Metaphor in 20th Century American Psychobiology.

Abstract approved: ______Paul L. Farber

By 1900 domestication was a promising, if somewhat vexed, subject in biology. Volumes had been written about domestication, but little serious scientific inquiry was directed toward the phenomenon. Expertise lay with practical men, primarily breeders and fanciers. The bulk of scientific commentary on domestication came from anthropologists who derived theories about man’s evolutionary past and future prospects based on an analogy with domesticated creatures. To an experimental ethos emerging near the turn of the 20th century, one increasingly dependent upon animals kept and bred in the laboratory, the available knowledge of domestication seemed inadequate, with its practical orientation and use of metaphor, analogy, and speculation. A small number of researchers working at various points along the fluid border between biology and psychology sought to reestablish the scientific understanding of domestication on the basis of experimental results.

I examine these latter efforts to determine how these investigators constructed new experimental understandings of domestication from the point of planning the experiments to interpreting the results and how these conceptions coincided with the widespread cultural resonances of domestication. Historians of science frequently correlate the experimental turn in biology and psychology not only with new standards of evidence, but also with new claims about disciplinary identity, expertise,

and objectivity. Domestication researchers, however, failed to produce a substantially new, clear, objective, and widely accepted explanation of the phenomenon by midcentury.

I argue that these efforts did not achieve the purported goals of experimental research, generally, not for any failure in the design of the experiments themselves, but for the continued cultural relevance of domestication, expressed in analogies, metaphors, and the wisdom of experience with domesticated animals, that corresponded with the values, social preoccupations, and professional circumstances of individual investigators. I argue, further, that the experience of domestication researchers demands a reevaluation of the impact of the experimental turn in biobehavioral research in the early years of the 20th century. This extends a recent historiogaphic tradition that recognizes continuities between pre-experimental and experimental work to include the relations of experimental scientists and non- scientific experts, the value of experience, and the use of analogy within the laboratory and without.

Second Nature: Domestication as Experiment and Metaphor in 20th Century American Psychobiology

by Nicholas Blanchard

A DISSERTATION

submitted to

Oregon State University

in partial fulfillment of the requirements for the degree of

Doctor of Philosophy

Presented August 7, 2014 Commencement June 2015

Doctor of Philosophy dissertation of Nicholas Blanchard presented on August 7, 2014

APPROVED:

Major Professor, representing History of Science

Director of the School of History, Philosophy, and Religion

Dean of the Graduate School

I understand that my dissertation will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my dissertation to any reader upon request.

Nicholas Blanchard, Author

ACKNOWLEDGEMENTS

The experience of writing this dissertation has been the most humbling, gratifying, enlightening, and remarkable of my life. I have encountered so many fascinating personalities in my researches: the credulous Clyde Keeler, “crushed by the gods”; Helen Spurway and Willard Small whose acknowledgement of assumptions attendant to domestication research were important precedents to my own; the tragic Barbara Burks; the controlling Milton Greenman. These figures and many others became minor fixtures in my life for a time. Their struggles and triumphs seemed, too often, to parallel my own. I thank them for sharing their stories with me, however difficult it was to tease them out.

I owe a tremendous debt to all those who have given me support and encouragement throughout my research and writing. The faculty and staff of the School of History, Philosophy, and Religion at Oregon State University has been gracious with their time and support since I arrived on campus absolutely green in the work of history. Ron Doel and Mary Jo Nye gave me an excellent introduction to the history of science and provided advice and encouragement always when I needed them most. Linda Richards, Tina Schweickert, Craig Beigel, Ron Gray, Mindy Gormley, Terry Christiansen, and Mason Tattersall each played the role of sounding board, colleague, mentor, and comic relief. I cherished my time with them. My most consistent source of support at OSU has unquestionably been Paul Farber. I was somewhat shocked when he agreed to serve as my major advisor, then newly in his retirement. However, his commitment to seeing this project through and to my growth as an historian has never wavered. He always seemed able to decipher what I was clumsily attempting and to gently, effortlessly set me on a clearer path. I will always be grateful for his genuine interest in my development and work.

This work would not have been possible without the support of the American Philosophical Society and the Philadelphia Area Center for the History of Science, which provided not only funding, but also access to critical resources. I would like to thank Roy Goodman at the APS Library and Babak Ashrafi and Bonnie Clause at

PACHS, especially, for their assistance and advice. Nina Long, librarian at the Wistar Institute Library in Philadelphia, also deserves special mention for her extraordinary efforts to provide me with resources pertaining to one of the most substantial domestication research programs ever undertaken. I thank the receptionists, librarians, curators, and archivists at the following institutions as well: the Library of Congress, Washington, D.C.; the Alan Mason Chesney Medical Archives of the Johns Hopkins Medical Institutions, Baltimore, Maryland; the National Library of Medicine, Bethesda, Maryland; the Rockefeller Archive Center, Tarrytown, New York; the Sterling Library of Yale University; the Firestone Library of Princeton University; the Proust Library of the University of Miami, Coral Gables, Florida; the Ina Dillard Russell Library at Georgia State College and University, Milledgeville, Georgia; the Valley Library, Oregon State University, Corvallis, Oregon. I am also grateful to Garland Allen for providing copies of his correspondence with Clyde Keeler and to Barbara Kimmelman for her support and helpful advice on this project.

Because no one has sacrificed more or been more unfailing with encouragement, I would like to thank my family. My parents have given me not only support, but the finest examples by which to live. My brother, Drew, remains my favorite confidant and a constant source of inspiration. I thank my children, Evan and Madelyn, for always reminding me of the world outside of work. Finally, I can only hope to someday return the love and support my wife, Theresa, has provided me throughout my work on this dissertation.

TABLE OF CONTENTS

Page

Introduction……………………………………………………………………………1

Section I: Dynamic Development in the Science of Man: The Culture of the Wistar Institute and the Origins of the Domestication Program………………………………………………………………………………18

From Synthetic Museum to Research Institute: The Founding Philosophy of the Wistar Institute………………………………………………………………………20

Uniting Philosophy and Experiment: Henry H. Donaldson and the Domestication Program at the Wistar Institute………………..……………..36

Analogy in Practice: The Wistar Institute Domestication Program and its Social Applications……………………..………………………………...58

Feral Rats, Captive Rats: A First Look at Domestication…………………....62

Experiments on Exercise and the Tone of Modern Life…………..………....72

Inbreeding, Hybridization, and Notions of Purity in Rats and Man…..……..77

Captive Breeding and the Elusiveness of Control……..………………….....94

Institutional Extension, Social Application……………………..………….109

Conclusion……………………………………………………..………...... 115

Section II: The Color of Temperament: Cultural Exchange Between Breeders and Geneticists…………………………..…119

Coat-color in History and Culture..………………………..………………..121

William E. Castle and the Bussey Insitute: The Culture of Coat-color Genetics…………………….…………….…….133

Domestication and the Genesis of Pigment Gene Pleiotropy………..……..145

Clyde E. Keeler- Education and Early Career……..……………………….146

TABLE OF CONTENTS (Continued)

Page

Life Beyond the Bussey……………………………………………..……...152

The Wistar Institute: Site of Discovery and Controversy…………….….....156

Research in Exile…………………………………………….……………..178

Applying Pigment Gene Pleiotropy to the Human Species………….……..202

Conclusion……………………………….………………………………....214

Section III: Beasts and the Burden of Metaphor: Domestication in American Psychobiology to Midcentury……………………………………………………....226

The Domestication Analogy and Notions of Progress in Pre-20th Century Social and Scientific Thought………………………………………..……..232

Domestication and Psychobiology: Identity and Approach………………...247

Psychobiology, Behaviorism, and the Fate of the Norway Rat………….....249

Freedom from the Tyranny of Instinct: Charles Otis Whitman, Progressivism, and Domestication Research at the Turn of the Century…..262

Charles R, Stockard, the Rockefeller Foundation, and the Preservation of the Constitutional View of Body and Mind at Midcentury……………...272

Domestication and Decline: The Degeneration Thesis of Curt P. Richter....307

Domestication and the Pychobiological critique of Behaviorism at Midcentury…………………………………………………………….....340

Conclusion……………………………………………………………………….....362

Bibliography………………………………………………………………………..378

Introduction

No doubt man selects varying individuals, sows their seeds, and again selects their varying offspring. But the initial variation on which man works, and without which he can do nothing, is caused by slight changes in the conditions of life, which must often have occurred under nature. Man, therefore, may be said to have been trying an experiment on a gigantic scale; and it is an experiment which nature during the long lapse of time has incessantly tried. Hence it follows the principles of domestication are important to us.

-C. Darwin, 1868.1

Charles Darwin leant domestication a grandeur it had not often held in natural history. By likening it to a great experiment, he suggested the sincerity and seriousness with which he and other naturalists should treat the phenomenon. By equating this experiment with the timeless activities of nature, he gave justification for this view.

There were, however, problems with this equation: to wit, the “experiment” existed in analogy, not reality. While Darwin had proposed concrete experiments to examine the origin and heritability of variations under domestication and voiced his “conviction of the high value of such studies,” he was forced to admit that “they have been very commonly neglected by naturalists.”2 It was axiomatic for many naturalists that domesticated creatures were “monstrosities” and “artifacts,” so far removed from the state of nature that they held little hope of illustrating even its most abstract and general principles.

Darwin’s trope of domestication as experiment seemed oxymoronic to many of his contemporaries, not only for their notions about domesticated animals, but also for the

1 Darwin, C.R. The Variation of Plants and Animals under Domestication. Vol.1. New York: Orange Judd, 1868. p.2-3. 2 Darwin, C.R. On the Origin of Species by Means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life. London: John Murray, 1859, facsimile of the first edition, Joseph Carroll (ed.), Peterborough, Ontario, Canada: Broadview Press Ltd., 2003. p.97.

2 social and epistemic barriers distinguishing the traditional craft of breeders and the activities of naturalists. To a vision of biology’s future that emerged near the turn of the

20th century- sequestered in the laboratory, under the most rigorously controlled conditions- domestication seemed even more contradictory to experimental ideals then being negotiated. Domestication had not been absent from scientific discourse, however.

Throughout the 19th century naturalists and budding anthropologists moved seamlessly between the examples of domesticated animals and man to discuss subjects ranging from human racial origins and slavery to the extent and future course of human civilization.

These associations, combined with the trove of popular notions and superstitions about domestication, may also have given pause to early experimentalists weighing Darwin’s call to investigate the phenomenon.

As experimentalists began to keep and to breed animals in the laboratory, however, they often looked to domesticated creatures, if not for the animals themselves, then for clues about how captivity and selective breeding would impact their subjects.

Domestication spoke to many of the questions that arose as new demands for objectivity and control confronted the desire to represent nature faithfully. If domestication was consistent with experiment, as Darwin’s analogy suggested, it would have bearing on issues of control and “naturalness” in modern biological research. Beginning around the turn of the 20th century, a handful of investigators finally heeded Darwin’s call and subjected domestication to experimental scrutiny.

In August of 1899, with “the dawn of the 20th century [and a new experimental era]… already in the sky,” embryologist Simon Henry Gage stressed the “transcendent importance” of domestication research at the American Association for the Advancement

3 of Science meeting in Columbus. He also acknowledged the ambiguous uses of domestication in earlier years, in which “sordid interests [played] too prominent a part.”

He continued, “Judging from the past, the study of domestic animals in any other way than in a scientific spirit and by the scientific method will prove barren, but studied in that spirit and by that method the result has always justified the effort, and has thrown as much, if not more, light upon biological problems than an equally exact study of a wild form.”3

For Gage, piety in the scientific spirit was enough to distinguish the scientific and

“sordid” interests in domestication. However, domestication was as much a cultural as a biological phenomenon, with its own practices, traditions, institutions, and myths.

Domestication was inescapably a human activity, and domestic animals were frequently made symbols of human virtue and vice. How did these prior notions coincide and conflict with the new agenda of experimental biology? Did the experimental era produce a new understanding of domestication loosed from its cultural moorings, or were they integral to its identity?

Ultimately, domestication was refashioned as an experimental subject in 20th century American biology and psychology. The primary goal of this work is to examine what this new identity meant for the understanding of domestication: its standing in the scientific domain, its social implications, and its previous understanding as a cultural entity. This objective demands a close look at the local contexts in which domestication was subjected to experiment and the individuals who attached such profound significance

3 Gage, S.H. The Importance and the Promise in the Study of Domestic Animals. Science. 1899. Vol10(245):305, 315.

4 to domestication not only as a proxy for organic evolution, but perhaps more importantly as a window into human nature and social change.

Domestication is notoriously difficult to define. It is a term as plastic as the animals and plants under its influence and as treacherous for the historian as for the biologist. Biological definitions suffer from imprecision and ambiguity. No universal thresholds separate domestication from captivity, taming, or acclimatization.

Maintenance in captivity over generations, control of breeding, dependence on human provision, morphological and/or behavioral change have been used to distinguish wild from domesticated varieties, but these criteria may be interpreted as matters of degree rather than qualitative distinctions. Further complicating any analysis of the problem, domestication has profound and varied cultural meanings, which, reciprocally, have influenced and been influenced by the term’s social and scientific usage. Domestication seems to reside almost exclusively in conceptual borderlands: between nature and culture, the wild and the civilized, human and animal, body and mind. It stands, like Janus, between worlds and yet, by facilitating reciprocal understandings, threatens to bring down the edifice between them.

In the literature of anthropology and philosophy, the definition and usage of

“domestication” have been significantly expanded and further complicated in recent decades. More recently, there has been an effort to rein in the excesses of this trend and offer a more precise definition of domestication for these fields. Rebecca Cassidy illustrates the dangers of the expanded meaning of domestication in contemporary anthropology: “What does it mean to domesticate desire (Brenner 1998)? Or Europe

(Hodder 1990)? Or death (Douglas 1978)? Or mobile phones (Ling 2001)?...

Domestication seems an attractive but ultimately unsatisfying euphemism, with a catch- all property that makes critique impossible.” She continues, “[T]he concept of domestication, like culture, is slippery and imprecise, those who invoke it often slide between distinct meanings. Others rely on a common sense interpretation of the term that leaves the assumptions it contains completely untheorized, although no less powerful.”4

It is one goal of this work to avoid the morass Cassidy describes, i.e. the proliferation of euphemistic meanings and the assumption of a common understanding of domestication. In an historical analysis of the meanings and implications of domestication, however, we must embrace a degree of ambiguity as our goal is not a general understanding of domestication but the understandings of particular historical actors- biologists and psychologists- in specific social and intellectual contexts. As anthropologist Nerissa Russell explains, “…the way domestication is defined is related to the definers’ view of the relationship between nature and culture and the place of humans with respect to nature.”5 Thus, another essential goal of this analysis is to uncover the assumptions that underlay these investigators’ definitions of domestication and informed the interpretation of their experimental results.

In one common view, however, experiments were meant to inform, not to be informed. Many pioneering experimentalists billed themselves as the vanguard of a new era of scientific biology, obliged to distinguish themselves from the older naturalists, who, lacking the appropriate rigor, were ostensibly inclined to prior assumptions and cultural biases. A longstanding historiographic tradition bolsters this impression, as it

4 Cassidy, R. Introduction: Domestication Reconsidered, in Where the Wild Things are Now: Domestication Reconsidered. Rebecca Cassidy and Molly Mullin (eds.) Oxford: Berg Publishers, 2007. p.3-4. 5 Russell, Nerissa The Wild Side of Animal Domestication. Society and Animals. 2002. Vol.10(3):294.

6 emphasizes the emergence of an experimental era of biology at the turn of the 20th century, characterized by the rejection of older, observational natural history and the rise of a more technical, objective, and epistemically pure experimental biology.6 A more recent (although by now established) historiography has challenged this standard story.

This newer interpretation acknowledges a significant shift toward experimental methodologies at this time, but rejects the notion that this transition represented a radical, and wholesale break with earlier goals and practices.7 Historians in this latter tradition have demonstrated the continuation (or perhaps appropriation) of various pre- experimental practices into the experimental era. The history of experiments on domestication reveals substantial continuities, not only in philosophical approaches, theoretical concerns, and practices, but also in the metaphors, norms, and other cultural resonances of domestication that survived the demand for greater objectivity. This look at early 20th century domestication research, however, addresses more basic concerns than whether the experimental turn was transformational or transitional; it highlights fundamental issues of identity, legitimacy, and social relevance in American biology and psychology in those years.

6 The most forceful late assertion of this position is Garland Allen’s Life Sciences in the Twentieth Century. Cambridge: Cambridge University Press, 1975. Slight amendments to the position appear in: Idem. Naturalists and Experimentalists: The Genotype and the Phenotype. Studies in History of Biology. 1978. Vol.3:179-209 and in Idem. Morphology and Twentieth Century Biology: A Response. Journal of the History of Biology. 1981. Vol.14(1):159-176. 7 See the Special Section on American Morphology at the Turn of the Twentieth Century in The Journal of the History of Biology 1981. Vol.14(1):83-191, especially Maienschein, J., Rainger, R. and Benson, K.R. Introduction: Were American Morphologists in Revolt? same volume p.83-87. See also the related edited volume The American Development of Biology. Ronald Rainger, Keith R. Benson, and Jane Maienschein (eds.) Philadelphia: University of Pennsylvania Press, 1988. For a more recent expression, see Nyhart, L.K. Natural History and the “New’ Biology in Cultures of Natural History. Nicholas Jardine, James Secord, and Emma Spary (eds.) Cambridge: Cambridge University Press, 1996. p.426-443.

These issues feature prominently in recent historical work on early 20th century

American biology and psychology. One line of investigation has looked at how the reconstruction of domesticated or semi-domesticated creatures into experimental organisms in these years was crucial in establishing identity and authority for American biologists and psyhologists. Robert Kohler situates a discussion about the epistemic authority of the laboratory space circa the 1910s within a tale of the rise of Drosophila from a semi-domesticated consumer of human refuse to the preeminent model organism of experimental genetics.8 Bonnie Clause explains that the development of the Norway rat as a “standard” laboratory animal at the Wistar Institute was inspired by contemporary efforts to standardize the production of manufactured goods, work practices, and other aspects of industry and social life in the Progressive Era.9 Karen Rader, in her aptly titled

Making Mice, describes how C. C. Little, alumnus of one pioneering center of mammalian genetics (Harvard’s Bussey Institute) and founder of another (Jackson

Memorial Laboratory), was inspired by Wilhelm Johannsen’s concept of “pure lines” to develop inbred strains of mice suited to the needs and whims of genetic and biomedical experimentalists. The semi-domesticated state of mice, as “hangers-on” to human civilization, Rader explains, allowed geneticists to avoid “trickier questions about how their genetics worked in the ‘wild.’”10 The apparent certainty of genetic background and behavior in standard laboratory animals was a crucial assumption that allowed

8 Kohler, R.E. Drosophila: A Life in the Laboratory. Journal of the History of Biology. 1993. Vol.26(2):281-310. 9 Clause, B.T. The Wistar Rat as a Right Choice: Establishing Mammalian Standards and the Idea of a Standardized Mammal. Journal of the History of Biology. 1993. Vol.26(2): 329-349. 10 Rader, K.A. Making Mice: Standardizing Animals for American Biomedical Research, 1900-1955. Princeton, New Jersey: Princeton University Press, 2004. p.36.

8 researchers to begin work on basic biological questions, and the results they achieved by adopting standard animals seemed only to justify the initial assumption.

Each of the above authors, and others still, recognize to some extent that the assumptions that attended the use standardized experimental animals were sometimes problematic. Rachel Ankeny and Sabina Leonelli, for instance, describe the “hybrid status” of experimental organisms, adapted to specialized purposes in the laboratory, but with a rich evolutionary history that is largely unacknowledged by the investigators using them.11 This troubling duality is true of domesticated animals generally, not only those adapted to life in the laboratory. Even with animals that were considered recently domesticated, there were often troubling gaps between what was known about the animal in captivity and its progenitor in the wild. In some cases, it was precisely this ahistoricity, this rootlessness, that made domesticated animals so appealing to laboratory experimenters concerned with theoretical questions rather than the complicated origins of their chosen research material. Ambiguity, both in the animal subject and in the laboratory space, conferred credibility, a sense that the results achieved in one instance could be applied universally without concern for situational anomalies.12 Although this perception was crucial in establishing the authority of laboratories and the standard animals within them, it is illusory. Laboratories and their denizens have histories, and these remain meaningful for the work they produce.

Of course, not all researchers in these years were willfully ignorant of the historical particularities of their research animals. Cheryl Logan notes that the two

11 Ankeny, R. and Leonelli, S. What’s So Special about Model Organisms? Studies in History and Philosophy of Science. 2011. Vol.42:313-323. 12 Kohler, R.E. Landscapes & Labscapes: Exploring the Lab-Field Border in Biology. Chicago: University of Chicago Press, 2002. p.6-11.

9 individuals most responsible for standardizing the Norway rat for experimental research,

Henry H. Donaldson and Adolf Meyer, were committed to a biological perspective that emphasized, at least initially, the ontogenetic and phylogenetic histories of their animal subjects. Their goal in standardizing the rat was making generalizations, but not in the sense of the rat being a “one-size-fits-all” model of generalized mammalian growth and behavior, as it later became. Rather, it was predicated on the observation that rats had developmental patterns similar to humans, and thus, that the Norway rat would not be a general mammalian model, but a specific one for human development and behavior.13

Logan’s focus on the perspective behind the choice of a standard animal is valuable as it not only considers whether the animal would bear appropriate experimental results, but reveals the presuppositions that researchers brought to the laboratory table as well as just how unsettled questions of objectivity, generality, and the place of experiment were in the early years of experimental psychology and biology.

The investigators featured in this work were keenly interested in the pre-domestic lives of the species in their laboratories. Many of them maintained a holistic biological perspective that emphasized the integrated, adapted nature of the organism, developed in deep, historical associations with its particular conditions of life. Just as Christopher

Lawrence and George Weisz describe for the development of medical holism in the first half of the 20th century, the biological holism associated with domestication research drew almost equally from strains of cognitive and cultural holism (involving “integrative

13 Logan, C.A. The Altered Rationale for the Choice of a Standard Animal in Experimental Psychology: Henry H. Donaldson, Adolf Meyer, and ‘the’ Albino Rat. History of Psychology. 1999. Vol.2(1):3-24; also Idem. “[A]re Norway Rats… Things?”: Diversity Versus Generality in the Use of Albino Rats in Experiments on Development and Sexuality. Journal of the History of Biology 2001. Vol.34(2):287- 314.

10 and comprehensive intellectual approaches to phenomena” and a “style of cultural and political critique aimed at various crises of modern Western society,” respectively).14

Domestication, as it described a range of seemingly interrelated morphological, physiological, and behavioral phenomena, appealed to familiar holistic notions of constitution, habitus (the amalgamated body and mind), and the general unity of knowledge. Culturally, domestication had already been employed in countless analogies with human civilization by anthropologists and social critics to complain about the fragmentation of modern life.

A related, historical approach has focused on the setting of research and its role in establishing disciplinary boundaries as well as epistemic and social authority. Kohler’s work on Drosophila, for instance, spawned a larger study of the laboratory space and its implications for the relationship between lab and field-work in modern biology. He describes how the authority of the laboratory as a “placeless place,” increasingly influenced the methods of biology in the field, a setting characterized by variation and unpredictable contingencies.15 The laboratory not only raised boundaries between lab work and field work, but also between ways of knowing, bearing on the standing of

“experts” as the laboratory increasingly became the home of trustworthy knowledge in the first decades of the 20th century. However, Kohler’s critics have pointed out that this

14 Lawrence, C. and Weisz, G. Medical Holism: The Context in Greater Than the Parts: Holism in Biomedicine, 1920-1950. C. Lawrence and G. Weisz (eds.). Oxford: Oxford University Press, 1998. p.6- 7. 15 Ibid. p.9.

11 interpretation is largely one-sided, perpetuating “the history of twentieth century biology from the perspective of the Roman wandering in the barbarian hinterland.”16

Domestication, especially in its cultural heritage, is impossible to imagine without consideration of its traditional custodians, the breeders. Although relations between breeders and naturalists in the 19th century had always been tenuous, scientific experts who dared to discuss domestication or artificial selection routinely cited evidence and even interpretations from practical breeders. In 1878 agricultural scientist Manly Miles declared with more admiration than lament that knowledge of animal heredity remained

“almost exclusively based on the experience of practical men.”17 A minor cottage industry has emerged in the history of biology to explore the relations of practical and scientific knowledge around the turn of the 20th century, finding generally that the exchange has been more complicated than hegemonic.18 Several of these analyses have shown how the fluid interactions of practical and scientific experts influenced the interpretation of research as well as its perceived social relevance. Most notably, Barbara

16 Mitman, G. Review: Landscapes and Labscapes by Robert E. Kohler. Journal of the History of Biology. 2003. Vol.36:599-601. 17 Quoted from Rosenberg, C.E. No Other Gods: On Science and American Social Thought. Baltimore: Johns Hopkins University Press, 1976. p.198. Originally from Miles, M. Stock Breeding: A Practical Treatise on the Applications of the Laws of Heredity to the Improvement and Breeding of Domestic Animals. New York: D. Appleton and Company, 1878. p.iii. 18 See Rosenberg, C.E. No Other Gods. p.167, 185-195; Kimmelman, B.A. A Progressive Era Discipline: Genetics at American Agricultural Colleges and Experiment Stations, 1900-1920. Ph.D. Dissertation, university of Pennsylvania, 1987; Rader, K.A. Making Mice. p.32-35; Paul, D.B. and Kimmelman, B.A. Mendel in America: Theory and Practice, 1900-1919, in The American Development of Biology Ronald Rainger, Keith R. Benson, and Jane Maienschein (eds.) p.281-310; Cooke, K.J. From Science to Practice or Practice to Science? Chickens and Eggs in Raymond Pearl’s Agricultural Breeding Research, 1907- 1916. Isis. 1997. Vol.88(1):62-86; Rossiter, M. The Organization of the Agricultural Sciences, in The Organization of Knowledge In Modern America, 1880-1920 Alexandra Oleson and John Voss (eds.), Baltimore: Johns Hopkins University Press, 1979. p.211-248; Sapp, J. The Struggle for Authority in the Field of Heredity, 1900-1932: New Perspectives on the Rise of Genetics. Journal of the History of Biology. 1983. Vol.16(3):311-342; Danbom, D.B. Publicly Sponsored Agricultural Research in the United States from an Historical Perspective, in New Directions for Agriculture and Agricultural Research: Neglective Dimensions and Emerging Alternatives Kenneth A. Dahlberg (ed.), Totowa, New Jersey: Rowman & Allanheld, Publishers, 1986. p.106-131.

Kimmelman and Kathy J. Cooke argue independently for the importance of the ideal of rural vigor and breeder’s notions of environmental influence in shaping American eugenics in the first decades of the 20th century.19

Nerissa Russell writes, “From the scholarly perspective, I believe that one of the difficulties of studying domestication, its simultaneously biological and social character, also is one of its virtues. Domestication is a concept that can bridge disciplines as well as mediating or even negating the nature/culture dichotomy.”20 A similar sentiment has driven my obstinate pursuit of so complicated and elusive a subject. The biologists and psychologists featured in this work, I endeavor to show, were no less enchanted by this dual character. A look at their work, in the context of early 20th century American experimental science, integrates several themes seen in the histories of experimental animals, research settings, and social and epistemic boundaries. Perhaps the most important precedent to this work is Richard W. Burkhardt, Jr.’s analysis of the role domestication played in Konrad Lorenz’s tailoring of his scientific interpretation to fit the political exigencies of the moment during the rise and after the fall of the Third Reich.21

Domestication, in Burkhardt’s account, played a significant, albeit supporting role behind his main objective to trace the development of ethology as a modern scientific discipline.

Here I have chosen to highlight domestication in large part to explore its uniquely rich cultural relations and how these have influenced its understanding in modern American

19 Kimmelman, B.A. The American Breeders’ Association: Genetics and Eugenics in an Agricultural Context, 1903-13. Social Studies of Science. 1983. Vol.13(12):183-189; Cooke, K.J. The Limits of Heredity: Nature and Nurture in American Eugenics Before 1915. Journal of the History of Biology. 1998. Vol.31:269-274. 20 Russell, N. The Wild Side of Animal Domestication. p.297. 21 Burkhardt, Jr., R.W. Patterns of Behavior: Konrad Loren, Niko Tinbergen, and the Founding of Ethology. Chicago: University of Chicago Press, 2005.

13 biology and psychology. Although this work leans heavily on the social relations of domestication research, I have tried, like Burkhardt, to give scientific content its due attention throughout. This “integration of ‘content’ and context’” is best achieved through a careful look at individual investigators and the contexts in which they worked and which they sought to influence.22

The United States early adopted the experimental ethos. The outlay of state and federal provisions for agricultural experiment stations and land grant colleges, culminating in the Adams Act of 1906, provided opportunity for American leadership in burgeoning scientific fields such as genetics, while results were more uneven in others, like comparative psychology.23 The decades surrounding the turn of the 20th century were marked by great promise for American science but also significant vulnerabilities.

Establishing legitimacy and authority in emerging scientific disciplines remained a significant concern for American scientists. Experiment offered one means of securing credibility, the concurrent trend of specialization, another. Both reshaped the identities and affiliations of scholars and consequently, their authority within the scientific community and beyond. The primacy of experiment and specialization was not, however, a foregone conclusion.

The respective roles of experiment and specialization in American biology and psychology moving forward were contingent and negotiated subjects. A number of pressing questions remained unsettled: would specialization make for surer knowledge or

22 Phrase borrowed from Burkhardt, Jr., R.W. Patterns of Behavior. p.13. 23 See Kimmelman, B.A. A Progressive Era Discipline for the case of genetics, and for comparative psychology see Mitman, G. and Burkhardt, Jr., R.W. Struggling for Identity: The Study of Animal Behavior in America, 1930-1945 in, The Expansion of American Biology, Keith R. Benson, Jane Maienschein, and Ronald Rainger (eds.). New Brunswick, New Jersey, Rutgers University Press, 1991. p.164-194.

14 facts dissociated from one another and, thus, an inaccurate picture of nature? Was experiment antithetical to (or should it displace) pre-experimental philosophies about life, evolution, and behavior? Would it sever ties to earlier ways of knowing, such as the empirical observations of breeders during their long experience crossing and cultivating animals? Could it really make uniform and transferable data from natural phenomena, so variable and full of peculiarities? Would it safeguard against assumptions, biases, and prejudices?

Many early bio-behavioral researchers asked pointed questions of their peers committed to experimental research programs. Thomas Wesley Mills, for instance, said of Edward Thorndike’s seminal puzzle box experiments: “…he placed cats in boxes only

20x15x12 inches, and then expected them to act naturally. As well enclose a living man in a coffin, lower him, against his will, into the earth and attempt to deduce normal psychology from his conduct.”24 Mills’s criticism extended beyond Thorndike, as elsewhere he broadly warned that “it cannot be too well borne in mind that our experiments are very clumsy imitations of nature in a large proportion of cases.”25 Yet

Mills did not forsake experiment, only those “so fast bound in the grip of… experiment” to lose sight of science’s real object, an accurate and meaningful understanding of the natural world.26 To Mills and to others skeptical of the experimental enterprise, intellectual rigor was not subsumed beneath the narrow parameters of control and

24 Mills, T.W. The Nature of Animal Intelligence and the Methods of Investigating It. Psychological Review. 1899. Vol6(3):266. 25 Idem. Instinct. Science. 1896. Vol.3(64):441. 26 Idem. The Nature of Animal Intelligence. p.266.

15 methodology; it required the questioning of unacknowledged assumptions, the hard work of observation and analysis, and the perilous task of interpretation.

Domestication is in some ways an ideal subject for exploring alternative visions of American biology and psychology at the dawn of the experimental era and how these ideas shaped the fields in the decades to follow. Domestication raised questions about the effects of control and captivity, predicated in concern about the apparent dichotomy between nature and artifice. Its multifarious and concurrent effects on the body and mind challenged disciplinary boundaries and attracted those interested in their interaction and synthesis. Its cultural legacy, both in metaphor and in the literal tradition of breeding and keeping animals, forced an evaluation of the ideals of objectivity and epistemic purity that had intensified in the experimental era. For the relative rarity of experimental programs dedicated to its understanding, domestication had a disproportionate influence in the development of American bio-behavioral research to the middle of the 20th century.

It also held outsized importance for the understanding of contemporary social problems, not only for familiar cultural notions about domestication, but for the deliberate efforts of domestication researchers to promote the social implications of their findings.

I have arranged this study in three sections, with the overarching approach an attempt at comparative scientific biography. The investigators featured in this work are for the most part minor, marginal, or forgotten figures, unfamiliar to current biobehavioral researchers and likely to students of the history of biology and psychology.

This is no history of great men. And yet among them are the greatest organizer and explicator of early 20th century American biology (Charles Otis Whitman), America’s foremost neuroanatomist, a man whose research interests and personal and professional

16 connections situated him uniquely well at the nexus of biology, psychology, and anthropology (Henry H. Donaldson), and one of the most prolific experimental psychologists of the 20th century (Curt P. Richter). That their notoriety does not match their achievements is not entirely surprising since, almost to the last, these figures left few scientific heirs and relatively meager records of their scientific and extra-scientific activities.

Section I of this work traces the conceptual development and realization of the most comprehensive experimental program on domestication in the United States prior to midcentury, at the Wistar Institute of Anatomy and Biology in Philadelphia. I examine, first, how the philosophical foundations of the Wistar Institute influenced its organization, hiring, research priorities, and approach during those formative years around the turn of the 20th century leading to the initiation of the experimental program on domestication by Donaldson shortly after his arrival in 1906. In the latter portion of

Section I, I explore the details of Donaldson’s multifaceted attack on the problem of domestication and explain the challenges that he and the Director of the Institute, Milton

Greenman, had in maintaining control over the project and its interpretation as it grew, passed into the hands of other associates, and was digested by the world outside of the

Institute’s walls. Section II centers on one of the most suggestive findings to emerge from the Wistar Institute domestication program, the idea that mutations in coat-color genes may have far-ranging effects in anatomy, physiology, and behavior. It proceeds, however, by following the fate of this idea in the hands of Clyde Keeler, a sometimes itinerant and perpetually marginal geneticist whose pursuit of the idea upon leaving

Wistar reveals interesting instances of the persistence of cultural ideas in experimental

17 science and the relations of breeders and geneticists, the traditional and new arbiters of knowledge on domestication, respectively. Section III addresses the cultural significance of domestication, especially its use in metaphor, in bio-behavioral studies. I also analyze the wider importance of domestication study in shaping approaches and research practices in fields contributing to the understanding of behavior.

This study is in no way exhaustive. It does, however, provide a clearer window into the relations of scientific and broader cultural strains of knowledge, of practical and scientific experts, of trust in experiment and trust in experience during a crucial period in the development of American biology and psychology.

Dynamic Development in the Science of Man: The Culture of the Wistar Institute and the Origins of the Domestication Program.

In 1892 the leadership of Philadelphia’s newly established Wistar Institute of

Anatomy and Biology was charged with the preservation and administration of the aging

Caspar Wistar and William Horner anatomical collections, formerly the responsibility of the University of Pennsylvania. The care of the large and haphazardly arranged anatomical museum could have been viewed as a terrible legacy cost for the new

Institute, given the move toward experimental methods and standards of evidence at similar research institutions. The Wistar administrators viewed this, however, as an opportunity to reorganize the collections to reflect their biological philosophy rather than medical or pedagogical utility, to transform American anatomy from the handmaiden of medicine to a pillar supporting a holistic program of evolutionary morphology.

The enduring legacy of the Wistar Institute’s founding was the establishment of a dynamic, but largely directional view of life, which served as a guiding philosophy for the Institute well beyond the tenure of its founding members. This vision, which borrowed from idealistic morphology and more directly from contemporary Neo-

Lamarckian and ontogenetic traditions of American evolutionism, relied upon familiar embryological analogies, generalized biological types, and progressive trends in evolution. Researchers and administrators at the Wistar Institute upheld dynamic development as the vera causa of organic evolution, which had its apogee in man.

Following this supposition, they encouraged comparative and developmental studies on animals with the goal of revealing truths about contemporary human types.

The early years of the Wistar Institute yielded little in the way of original, experimental research. Prior histories of the Institute reveal that this disappointing record of achievement prompted a reorganization of the Institute just a decade after its founding.

The installation of Milton Greenman as Director in 1905 and Henry H. Donaldson as

Director of Research a year later signaled a methodological shift at the Institute. Rather than an ancillary goal, experimental research would become the primary responsibility of the Wistar Institute under Greenman and Donaldson. This did not, however, amount to the Institute “shaking off its Victorian origins.”27 While these accounts appropriately emphasize strategic and methodological changes, some attention is due for the substantial continuities that linked the biological philosophies, investigative ideals, and social goals at the Wistar Institute before and after its reorganization in 1905.

Shortly after Donaldson’s arrival, his research program was dominated by a search for factors to explain the differences between his chosen research material, the domesticated albino Norway rat, and the wild gray, its presumed progenitor. The subject of domestication seemed particularly well suited to the Institute’s philosophical leaven.

It was regarded as one of the most dynamic of biological phenomena, as amenable to the comparative approach favored at the Wistar, and by social scientists as relevant to the trajectory of human civilization. Donaldson’s domestication program resonated, rather than competed, with the philosophical tradition already established at the Wistar Institute, especially with the focus on human applications of the work completed there.

27 Clause, B.T. The Wistar Institute Archives: Rats (Not Mice) and History. Mendel Newsletter. 1998. No.7. p.3.

From Synthetic Museum to Research Institute: The Founding Philosophy of the Wistar Institute

The anatomical collections of Caspar Wistar had been used to establish an eponymous museum in 1808. Despite its care and expansion under Wistar’s successors to the chair of anatomy at the University of Philadelphia, including William Horner and

Joseph Leidy, the Wistar and Horner Museum fell into disrepair. A fire in Medical Hall, where the collections were held, compelled University Provost William Pepper to find a more suitable arrangement for the safekeeping of the Wistar collections. Pepper turned to affluent Philadelphians to help secure funding and a new home for the nation’s oldest and largest anatomical museum.

His most valuable conquest was General Isaac Jones Wistar, Caspar Wistar’s great nephew and a well-heeled Philadelphia attorney and businessman. Following a somewhat aimless youth, the younger Wistar achieved distinction, if not always success, in the Union Army where he reached the rank of brigadier general. Wistar’s accomplishments came steadily after the war, with key investments in canals and railroads, a burgeoning law practice, and a surer place among Philadelphia’s social and intellectual elite. Rescuing the museum from penury and neglect would not only establish Isaac Wistar’s philanthropic credentials, Pepper assured him, but also reaffirm his family’s pride of place within American and particularly Philadelphian anatomical science. In 1891, Gen. Wistar donated $20,000 to a trust fund to preserve the Wistar-

Horner Museum and followed that largess less than a year later with a plan to incorporate an entirely new institute charged with the creation of new and original knowledge. The new Wistar Institute of Anatomy and Biology, named for the General’s illustrious great-

21 uncle, was to be housed in a new, “fireproof” edifice rising near the corner of 36th and

Spruce on the University of Pennsylvania campus. General Wistar bankrolled the construction of the building and established an endowment and further trusts for the independent funding of the Institute.28

Independence- fiduciary, administrative, and epistemic- was central to Wistar’s notion of a modern research institute. General Wistar was not an oblivious deep pocket, content with immortalizing a family legacy. Even before money troubles and a timely fire presented him with the opportunity to build a research institute, he had a vision for what such a station might look like and how it might be operated. After his discharge from military service, Gen. Wistar pursued his interest in the natural sciences as aggressively as his business ventures. He inserted himself into the circle of

Philadelphia’s most prominent naturalists and medical men as a member of the Biological

Club, a fortnightly dining society led for the bulk of its years by Joseph Leidy. His closest associates in the sciences included Leidy, Horace Jayne, Edward Drinker Cope, Harrison

Allen, and John Adam Ryder. General Wistar’s involvement in institution building also preceded the call to preserve his family’s scientific heritage. As an administrator of the

Academy of Natural Sciences of Philadelphia’s Building Fund, Wistar visited the zoological gardens, natural history museums, and research institutes of Europe in 1888.29

28 Greenman, M.J. Appendix: The Wistar Institute of Anatomy and Biology in Wistar, I.J. Autobiography of Isaac Jones Wistar, 1827-1905 Volume II. Philadelphia: Wistar Institute of Anatomy and Biology, 1914, p.164. Also, Logan, C.A. “[A]re Norway Rats… Things?” p.305; Clause, B.T. The Wistar Rat as a Right Choice. p.333; Rovera, G. The Wistar Institute. Molecular Medicine. 1997 Vol.3(4):229; Brosco, J.P. Anatomy and Ambition: The Evolution of a Research Institute. Transactions and Studies of the College of Physicians of Philadelphia. 1991. Ser.5 Vol.1(1):1-28. 29 Wistar, I.J. Autobiography Vol.II, p.153-155. By 1891 Wistar had also been named President of the Academy of Natural Sciences and Director of the Philadelphia Zoological Society. See Greenman, M.J. Appendix p.162-166.

When William Pepper and James Tyson, Dean of the Medical School at Penn, solicited his help to rescue the Wistar-Horner collections, Isaac Wistar likely had greater designs than mere preservation. With the help of his scientific associates, he re-imagined the organization and use of the Wistar-Horner Museum around the model of independent research institutes like those he had visited in Europe. The new Wistar Institute was to be independent of the influence of any entity in Philadelphia’s established scientific tradition. It would be seeded, however, with a particular guiding philosophy, which shaped the Institute itself as well as the work and interpretations emanating from it.

Isaac Wistar saw through the incorporation of the Wistar Institute in April 1892.

The museum collections remained on the Penn campus until the completion of the new fireproof building in 1893 at a cost of $125,000. The Wistar Institute was christened the following May with a series of speeches that vacillated between the solemn responsibilities and the ripe hopes pinned on the nascent institution. Harrison Allen, the

Institute’s first director, spoke of both. He reminded his audience, “At least upon the anatomist rests the responsibility of explaining the structure of the animal kingdom and through a knowledge of this structure how human ills can be alleviated.”30

The biological philosophy that produced the particular shape and character of research at the Wistar Institute lay tacit in Allen’s declaration of duty. Isaac Wistar’s scientific associates in Philadelphia, especially Ryder and Allen, had detailed a grand new vision for anatomy, independent of medicine but complementary to its goal of improving the human condition. Their approach was comparative, seeking fundamental biological truths from a canvass of the diverse forms available to them. Their focus on

30 Allen, H. The Objects of the Wistar Institute. University Medical Magazine (University of Pennsylavania). 1894. Vol.6:587.

23 dynamic development and archetypal forms recalls the lasting influence of teleological morphology in the United States, primarily through the conduit of Louis Agassiz. Like

Agassiz, the Wistar group repudiated the overtly metaphysical elements of

Naturphilosophie, but their own conceptions of nature were strongly directional and relied on theoretically convenient correspondences. Nature, viewed progressively, likened itself to the developing individual, tempting knowledge seekers to uncover its underlying plan in the unfolding embryo. Life, viewed broadly, promised to reveal its essential forms to those who could generalize from its diversity. These perspectives appealed to Wistar’s Philadelphia circle in the late 19th century much as they had to

Romantic naturalists more than fifty years earlier.

By the time Wistar and his associates began planning their vision of a modern research institute idealistic morphology had given philosophical structure to a new and largely American evolutionism. Perhaps the most distinctly American model of organic evolution proposed that the stages added onto (or removed from) the normal ontogeny of a species were literally the means of ascension (or decline) in a perceived evolutionary scale. The proponents of this view were primarily students or disciples of Louis Agassiz, whose description of parallel series in nature betrayed his idealism:

[T]he phenomena of animal life correspond to one another, whether we compare their rank as determined by structural complication with their phases of growth, or with their succession in past geological ages; whether we compare this succession with their embryonic growth, or all these different relations with each other and with the geographic distribution of animals upon earth. The same series everywhere!31

31 Agassiz, L. An Essay on Classification. London: Longman, Brown, Green, Longmans, & Roberts and Trübner and Co., 1859. p.196.

Despite Agassiz’s vehement opposition to transmutationism, his followers in the younger generation were quick to see how his ideas could be applied to such a theory.

Alpheus Hyatt, a former student of Agassiz, and Edward Drinker Cope, who was firmly under Agassiz’s influence if not his direct tutelage, became the leading proponents of the acceleration of growth concept, whereby new species emerge by elaborations to the ontogeny of an existing species. Historian of science Peter Bowler suggests that to provide a source for the changes in ontogeny, the acceleration of growth was subsequently aligned with the Lamarckian concept of inherited acquired characters, especially with “internal directing mechanisms- physiology and behavior- that would explain the consistency of variation.”32 Another of Agassiz’s students, Alpheus Packard

Jr., dubbed this biological view “Neo-Lamarckianism” in 1885.33 Others, however, including Isaac Wistar’s close friend John Adam Ryder had identified the Lamarckian influence in this evolutionary concept years earlier.34

Ryder was perhaps the catalyst for the establishment of the philosophical tradition at the Wistar Institute. He was best known for his strongly mechanical view of evolutionary change, which attributed such modifications to “knowable and discoverable causes originating in the mechanical and dynamical conditions which surround living organisms and by which they are related to the cosmos.”35 He hoped that this conception would not only grant biology the same prestige as physics, but would also make it

32 Bowler, P.J. The Eclipse of Darwinism: Anti-Darwinian Evolutionary Theories in the Decades around 1900. Baltimore: Johns Hopkins University Press, 1983. p.119-120. 33 Pfeifer, E.J. The Reception of Darwinism in the United States, 1859-1880. Ph.D. dissertation. Providence, Rhode Island; Brown University, 1957. p.167. 34 Ryder, J.A. On the Origin of Bilateral Symmetry and the Numerous Segments of the Soft Rays of Fishes. The American Naturalist. 1879. Vol.13(1):43. 35 Ibid.

25 epistemically continuous with it. For this reason Bowler suggests that “Ryder does not appear to have been influenced by Agassiz’s idealism.” 36 Ryder’s mechanistic explanations and repudiations of some teleological speculations notwithstanding, his broad view of organic evolution was clearly progressive and contained many of the hallmarks of Agassiz’s idealistic philosophy (although Herbert Spencer’s universal evolutionism seems to have been a more direct influence on Ryder).37

In Ryder’s schema, mechanical factors did not reduce dynamic development to oblivion, but rather instantiated the concept, which remained almost indistinguishable from that of his more idealistic contemporaries. The developing individual is not subject to every capricious change of wind (or other physical force), but rather seems to follow patterns already established in the phylogenetic history of the species:

It passes… not by leaps, but from one stage to the next higher, and so on in an absolutely continuous manner… so that absolute continuity becomes a fundamental characteristic of the process of development… The being, in its evolution of the ovum, accordingly recapitulates the forms of its successively more and more complex, or more and more modified ancestral series- its paleontological history preserved in the rocks together with more or less note of its contemporary allies… This doctrine and its modifications is [sic] the motive force of modern Biology.38

W.P. Wilson, Ryder’s colleague and a botanist at the University of Pennsylvania, characterized the Weltanschauung driving Ryder’s biological theorizing: “[It] was his

36 Bowler, P.J. The Eclipse of Darwinism p.137. 37 That Ryder was not hostile to the basic idea of progressive evolution may be witnessed in a statement that concluded the first installment of his large study of mechanical forces and the evolution of tooth forms: “It is hoped that a better appreciation of what might, without violence to commonly received ideas, be called evolutionary teleology, may be attained by pursuit of similar inquiries in other directions.” In Ryder, J.A. On the Mechanical Genesis of Tooth Forms. Proceedings of the Academy of Natural Science of Philadelphia. 1878. Vol.30:45. 38 Ryder, J.A. The Gemmule vs. the Plastidule as the Ultimate Physical Unit of Living Matter. The American Naturalist. 1879. Vol.13(1):13.

26 intense desire to formulate a harmonious theory of evolution and development of life…

He hoped to make the aggregation of facts and principles explain more fully the thinking, moving, existing universe around us.”39 Certainly Ryder attempted this with an overtly mechanical theory, but his worldview was simultaneously organic and holistic. “Life,” he announced, “is continuous in the material succession and derivation of its individuals… Vital energy and cosmical energy are coexistant and continuously cooperative.”40 In his dynamic and directional transformism, Ryder seems to have reconciled his commitment to scientific rationalism and his abiding “trust in an infinite beneficence.”41 Mechanical principles helped him describe the spirit that he believed imbued life.

Early in 1893 Ryder published an outline for the institutional realization of his biological philosophy entitled, “The Synthetic Museum of Comparative Anatomy as a

Comprehensive Basis for Research.” Even as the walls of the Wistar Institute were rising on 36th Street, Ryder never mentioned the Institute by name. He indicated, however, that the essay had been fully formed since the autumn of 1887, but publication was demanded now by circumstances and the “solicitation of friends,” of whom only Harrison Allen was specifically named. Despite Ryder’s claims that his ideal museum would liberate the researcher from existing dogma, it is clear that his exhibits were meant to instill a particular view of nature: “Objects should be arranged on the shelves of an anatomical museum as to indicate the order of nature, the sequence of the phenomena of being; in

39 Wilson, W.P. Dr. Ryder as a Colleague. In In Memoriam: John Adam Ryder. Philadelphia: Committee of Edward Drinker Cope, Amos P. Brown, J. Percy Moore, and Philip P. Calvert, 1895. p.27. 40 Ryder, J.A. The Principle of Conservation of Energy in Biological Evolution: A Reclamation and Critique. Proceedings of the Academy of Natural Science of Philadelphia. 1892. Vol.44:460. 41 Allen, H. A Biographical Sketch of John Adam Ryder. Proceedings of the Academy of Natural Sciences of Philadelphia. 1896. Vol.48:239

27 short, it should aim to illustrate, in the broadest sense of the words, the philosophy of living things.”42 Students and researchers at the institute would ostensibly be intellectually free, and in their wanderings, “[i]t will soon be discovered that new modifications are superimposed upon old ones, and that the method of progressive modification is thus made palpable as it cannot be in any other way.”43 Objective research would bring them to those truths that Ryder held to be fundamental, and, if not, the arrangement of specimens would gently lead them to the correct interpretation. At

Ryder’s hypothetical museum the human brain was the citadel to be breached: “Can we ever hope to make out the history of an organ so intricate in structure and important in function? The answer… is to be sought in comparative anatomical and embryological research- in the process of becoming the brain.”44 Evolution had reached its zenith in the human brain and by use of the embryological analogy the historicized brain could be epitomized in the development of lower animals.

Harrison Allen, who had solicited Ryder’s essay for publication, was named the first Director of the Wistar Institute in 1892. He retired from the position less than two months after the Institute was formally opened in May of 1894. In his short tenure he was unable to realize his friend’s concept of a synthetic museum, but successfully inculcated the Institute with the substance and spirit of Ryder’s philosophy. Before he had convinced Ryder to publish his essay on the “Synthetic Museum,” Allen had published his own vision for an institute for research and teaching in the area of anatomy.

42 Ryder, J.A. The Synthetic Museum of Comparative Anatomy as a Comprehensive Basis for Research. Zoological Contributions from the Biological Laboratory of the University of Pennsylvania. 1893. Part 1, No.1 p.1. Italics original. 43 Ibid. p.6. 44 Ibid. p.9.

Allen’s hypothetical museum was similar in many ways to Ryder’s, especially his emphasis on comparative and developmental studies of human, other mammalian, and avian brains. He was clear about the need for sovereignty: “Independence in intellectual as well as in political life should be the object of American citizenship… If the course [in advanced anatomy] were well established, it would be well to institute a laboratory and museum distinct from any on the university grounds. I am of the opinion that the administrative success of such separation of collections would be assured.”45 This opinion was publicly expressed just months before Isaac Wistar reached a formal agreement with the University of Pennsylvania to establish the Wistar Institute as a body independent of any other on the University campus.

At the opening of the Institute Allen expressed his hope that “the Wistar Institute will be spared the affliction of displaying any specimen whatever which is not in exact illustration of an idea.”46 He expanded Ryder’s biological philosophy, connecting it more firmly with ontogenetic concepts about racial and individual senescence, a position he expressed most clearly in Science a few years later: “As the individual nears the time of its own extinction it experiences changes in the composition of its tissues and gross variation in characters. Groups of animals which are also approaching extinction behave in a similar manner.”47 Whatever qualms Ryder may have had with evolutionary teleology, Allen displayed none. Ontogeny and phylogeny both proceeded by increased specialization of parts. Specialization passed to rigidity and finally to extinction.

45 Allen, H. On the Teaching of Anatomy to Advanced Medical Students. Science. 1892. Vol.19(471):85. 46 Allen, H. Objects of the Wistar Institute. p.590. 47 Allen, H. On the Effects of Disease and Senility as Illustrated in the Bones and Teeth of Mammals. Science. 1897. Vol.5(112):294.

Morphology could thus be used to diagnose the organism or species’ stage of senescence.

Allen, in a portent of the research program that would take root at the Institute almost twenty years later, believed that domesticated animals were more specialized than their wild counterparts and displayed physical signs of this advanced development. He inquired, “After what manner may one expect taxanomic characters modified in these generations of prisoners?... What proportions of malignant growths, such as sarcomata, are met with in the feral state of quadrupeds as compared with those in the domesticated or the captive state?”48

Domestication in Allen’s mind was not limited as a model of diseased action, but was central to his efforts to reform anatomy at the time he assumed leadership of the

Wistar Institute. Significantly, he reminded the Congress of American Physicians and

Surgeons: “[T]he anatomical material coming under our notice is almost always that of a highly domesticated, or as one may say, over-acclimated nature, and that in order to study it with advantage we must recall what is common between man and the animals (both in a wild and domesticated state) about him.”49 That is, the physician and the anatomist must keep in mind that the modern human is a modified form, altered by a process parallel, if not identical, to animal domestication. That Allen considered most domesticative changes to be degenerate, diseased, or otherwise morbid from over-refinement made them no less progressive in his scheme of ontogenetic evolutionism.

48 Allen, H. On the Teaching of Anatomy. p.86-87. Also see Allen, H. On the Effects of Disease and Senility. p.294 49 Allen, H. Morphology as a Factor in the Study of Disease. Transactions of the Congress of American Physicians and Surgeons. 1894. Vol.3:8-9.

Allen’s pioneering work in modern physical anthropology further developed the analogy between domestication and civilization.50 He insisted, “Man has doubtless undergone changes in his organization resembling those which existed in a domesticated animal when his organization is compared to that of an ancestral form.”51 Thus, when he compared skulls from civilized and primitive peoples he noted, “what impresses me most in a series of skulls of civilized centers is the absence of correlation… The bones themselves appear to be individualized,” a finding which his contemporaries paralleled with the rise of the individual in modern cultures.52 His thinking about the anatomical effects of civilization (and domestication) evokes, again, the central progressive tenet of transcendental morphology: ontogeny (serving here as a model for organic as well as sociological telos) proceeds from the general to the specialized, from the archetype to the individual.

Allen intended that the domestication analogy and his own findings on various human groups would inspire active research and even experimentation in anatomy with an eye toward the source and direction of variation. He encouraged students in advanced anatomy to imagine the implications of their studies beyond the needs of practical medicine, to consider that a synthetic knowledge of anatomy would ultimately benefit humankind more than what was immediately practicable. He imagined that “[w]hen this phase of comparative anatomy shall have been formulated we shall for the first time have a reasonable hope that … the history and destiny of man himself may be in shape for

50 For Allen’s role in establishing modern American physical anthropology see Hrdlicka, A. Contribution to the History of Physical Anthropology in the United States of America, with Special Reference to Philadelphia. Proceedings of the American Philosophical Society. 1943. Vol.87(1):63. 51 Allen, H. Morphology as a Factor. p.9. 52 Brinton, D.G. Dr. Allen’s Contributions to Anthropology. Proceedings of the Academy of Natural Sciences of Philadelphia. 1897 Vol.49:526. Quotation from Allen, H. Morphology as a Factor. p.13.

31 elucidation.”53 Only months prior to the foundation of the Wistar Institute, Allen invoked

Bacon’s ‘New Atlantis,’ where “we read of ‘parks and enclosures of all sorts of beasts and birds, which we use not only for view or rareness, but likewise for dissection and trials, that thereby we may take light what may be wrought upon the body of man.’”54

Allen’s ambitions for the Institute, like Ryder’s, were rooted in the idea that a broad and comparative approach would reveal fundamental patterns in nature and thus a better understanding of its highest achievement, humankind.

The Wistar Institute came under the leadership of Horace Jayne, another of Gen.

Wistar’s associates and Professor of Anatomy at the University of Pennsylvania, upon

Allen’s retirement. Jayne held the directorship for ten years but published little to suggest how the biological philosophy at the Wistar Institute progressed in that time. He contributed much, however, to the size and arrangement of the museum collections. He took as his model the Hunterian Museum in London, which had influenced Allen and

Ryder’s conceptions of an illustrative anatomical museum.55 Jayne was also cited in support of the particular modes of evolution advanced by Ryder and Allen.56 Certainly,

Jayne seemed to be engaged in the philosophical program that had been established at the

Wistar. However, during his tenure the Institute produced little of the new and original research that Isaac Wistar had hoped would be its hallmark. Jayne left the Institute in

1905 to join the Board of Trustees at Drexel University, though unsatisfactory leadership

53 Allen, H. Morphology as a Factor. p.13. 54 Allen, H. On the Teaching of Anatomy. p.87. 55 Scientific Advisory Committee of the Wistar Institute, Memoir of Milton J. Greenman. Philadelphia: Wistar Institute of Anatomy and Biology. 1937. p.13. 56 Ryder, J.A. The Principle of the Conservation of Energy . p.462.

32 has also been cited as a reason for his departure.57 Perhaps supporting this interpretation, the Institute underwent a significant reorganization after Jayne ceded the directorship to his assistant, Milton J. Greenman.

Experimental research at the Wistar began under Greenman’s direction. He had served as assistant director under both Allen and Jayne before taking administrative leadership of the Institute. In the histories offered of the Wistar Institute, his promotion is viewed as the turning point of the Institute’s research program. This view is in large measure accurate.58 Greenman seized the opportunity to reinvent the Institute in the mold of a modern research center, unlike his predecessors who lacked either the time or the ambition to realize the Institute in this way. He immediately set to work refining the focus of research at the Institute and declared at the end of his first year as Director,

“T]his is the first year in the history of [the Wistar Institute’s] existence when the chief aim of the Institute has been research and when the museum and all departments of the

Institute have been made subservient to its aim.”59

With the help of Isaac Wistar, he convened a meeting of many of America’s premier anatomists to discuss new directions in the field and the role the Wistar Institute should play in their development. From attendees of the meeting he formed a permanent

Advisory Board to guide the Institute’s hitherto desultory research output. The Advisory

57 Brosco, J.P. Anatomy and Ambition. p.13; Clause, B.T. The Wistar Rat as a Right Choice. p.333. 58 On the methodological shift at the Wistar Institute see: Brosco, J.P. Anatomy and Ambition p.13-14; Clause, B.T. The Wistar Rat as a Right Choice p.333; Logan, C.A. “[A]re Norway Rats…Things?” p.305; Baatz, S. Biology in Nineteenth-Century America: The Wistar Museum of Anatomy in Renato G. Mazzolini (ed.) Non-verbal Communication in Science Prior to 1900. Florence: Leo S. Olschki, 1993. p.475-477; Pauly, P.J. The Appearance of Academic Biology in Late Nineteenth-Century America. Journal of the History of Biology. 1984. Vol.17(3):391. 59 Greenman, M.J. Report of the Director of the Wistar Institute of Anatomy and Biology Appendix IX in Report of the Provost of the University of Pennsylvania 1907.

Board decided that the primary role of the Wistar in the field of anatomy would be that of an original research institute rather than a museum for “a merely gaping public.”60 From among their ranks, they elected the neuroanatomist Henry H. Donaldson as Director of

Research at the reorganized Institute. In the style of similar institutes in Europe, the

Advisory Board declared that the Wistar should become specialized, concentrating in the areas of neurology and embryology. Greenman indicated that general calls for cooperation from the German embryologist Wilhelm His and his compatriot the neurologist Paul Flechsig had inspired the Advisory Board to focus on these fields.61 He neglected to mention, however, that more immediate and local influences suggested these areas of specialization to the Wistar leadership. John Adam Ryder had made the same fields the focuses of his hypothetical museum of comparative anatomy. Harrison Allen reiterated Ryder’s call just months before he became Director of the Wistar Institute.62

Despite the sense that the Wistar Institute’s 1905 reorganization was a complete departure from the traditions established thirteen years earlier, the philosophy guiding the work performed there survived the methodological shift.

Milton Greenman had been a part of the Wistar Institute since its founding, his reputation as a harbinger of change notwithstanding. His associations with Allen, Jayne, and Ryder reached back further still. He had graduated in 1889 from the Department of

60 Quotation from a letter by the late Isaac Jones Wistar to Milton Greenman in Greenman, M.J. Appendix . p.170. The Advisory Board consisted of Lewellys F. Barker (Johns Hopkins), Edwin G. Conklin (U. Penn.), Simon H. Gage (Cornell), G. Carl Huber (U. Michigan), George S. Huntington (Columbia), Franklin P. Mall (Johns Hopkins), J. Playfair McMurrich (U. Michigan), Charles S. Minot (Harvard Medical), Henry H. Donaldson (U. Chicago), and George A. Piersol (U. Penn.), all of whom were committed to a synthetic and biologically meaningful view of anatomy, apart from medicine. 61 Ibid. 62 See Ryder, J.A. The Synthetic Museum of Comparative Anatomy p.9, and Allen, H. On the Teaching of Anatomy. p.86.

Biology at the University of Pennsylvania, where Ryder and Jayne were on the faculty and Allen had recently become emeritus due to failing health. The junior Greenman then had at least nominal oversight of his former mentors as Director of the University’s short- lived Laboratory of Marine Biology at Sea Isle City, New Jersey, where all four men taught classes in the early 1890s.63 Greenman also followed in John Adam Ryder’s footsteps at the U.S. Fish Commission station at Woods Hole. Both men retained strong ties with the place through their involvement with the Marine Biological Laboratory where they spent most summers.64 In 1914 Greenman acknowledged the remaining influence of these philosophical leaders of the Wistar’s founding, especially Harrison

Allen, who “left his impression upon the future development of the institute which today is following his suggestions as closely as if he were still its Director.”65 Greenman’s words were more than empty plaudits for his late boss. The ideas of Allen, Ryder, Jayne, and Isaac Wistar were indeed implemented even as the Wistar Institute was recast as a center for modern experimental research. Greenman followed their suggestions to focus on neurology and embryology, to adopt a comparative approach, to delineate archetypes from those comparisons, and to interpret the human implications of those findings.

He published little research of his own at the Wistar Institute, especially after assuming the Directorship from his former Penn professor Horace Jayne. It is clear, however, that his thinking about anatomy and biology was strongly typological and,

63 “The University of Pennsylvania” The University Magazine. 1892. Vol.7:124. 64 For Ryder’s involvement with the Fish Commission and MBL, see: Dean, B. Dr. Ryder’s Work with the United States Fish Commission. In In Memoriam: John Adam Ryder. p.12 and Moore, H.F. Dr. Ryder as a Teacher. In the same volume, p.22. For Greenman’s involvement see: Stockard, C.R. Minutes on the Death of Milton J. Greenman. Biological Bulletin. 1940. Vol.79(1):16 and Clause, B.T. The Wistar Rat as a Right Choice. p.334. 65 Greenman, M.J. Appendix. p.167.

35 furthermore, was attuned to contemporary social problems. For instance, while still the

Assistant Director of the Institute in June of 1898, Greenman explained to the press that physiognomy foretold an American victory in the war with Spain: “Among other traits the Spanish face denotes is a lack of power of endurance and a notable deficiency in the lines of determination… According to his physiognomy, the Spaniard is brave only by impulse, and is not naturally possessed of the courage to face an enemy on equal terms or at close fighting quarters.”66 As Director, he devoted some portion of his time to a project to define the criminal type from an analysis of crania.67 He also supported research on the Institute’s collections of brains from eminent men, particularly scientists, to identify the neural peculiarities associated with social and intellectual distinction.

Greenman’s training in medicine and interest in evolution were jointly engaged by the budding American eugenics movement in the second decade of the 20th century. He was named chairman of Philadelphia’s Committee on Provision for the Feebleminded, a signal that the work of the Institute gained broader, more sociological implications under

Greenman.68

It is true that Greenman’s most earnest ambitions were in establishing a thoroughly modern research tradition at the Wistar. It is equally true that, at least initially, he sensed no reason why this goal would be incompatible with the prevailing biological philosophy of the Institute.

66 “American Superiority: Reasons Why We are Able to Thrash Spain.” The Evening Times. (Washington, D.C.) June 2, 1898. Greenman’s focus on the facial characteristics of race may demonstrate the influence of Allen who specialized in facial anatomy in his anthropological studies. 67 White Jr., William J. What Does Your Face Tell Your Boss? Popular Science. 1927. Vol.112:48. 68 Author unknown A Committee to Eradicate Feeblemindedness. The Survey. 1915. Vol.34:369. See also the following chapter in this volume for a fuller illustration of how the ties between the Wistar Institute and the American eugenics movement grew stronger under Greenman’s direction.

Uniting Philosophy and Experiment- Henry H. Donaldson and the Domestication Program at the Wistar Institute

“Concentrated and coordinated efforts according to a program might be our slogan [at the Wistar Institute],” wrote Milton Greenman in 1925.69 He felt, by then, that he had successfully given direction to the study of anatomy as had been the goal of the

Institute since it was merely a vision in the minds of Ryder, Allen, and Isaac Wistar.

Greenman had committed himself to transforming the 19th century museum to a 20th century center for experimental investigation. In so doing he determined that he needed an equally committed partner to guide the research proper while he took care of the general administration of the Institute. In 1905 he hired Henry H. Donaldson, at the time on the faculty of the University of Chicago, as Director of Research and Professor of

Neurology at the Wistar Institute. Donaldson’s election by the Institute’s Advisory

Board may have been little more than a formality; Greenman felt that he had already found an ideal fit for the Institute in Donaldson. Greenman’s promotion to the directorship of the Wistar coincided with the publication of a paper by Donaldson that was similar in scope and disciplinary vision to the missives written by Ryder and Allen more than a decade earlier.

In the paper Donaldson asked his readers to inquire: “[W]hat is the purpose of all this anatomical work; for without a strong guiding idea we are liable to repeat the errors of earlier generations, and merely accumulate observations.”70 He called for institutes to allow their researchers to devote themselves to investigation rather than teaching. His

69 Greenman, M.J. The Wistar Institute of Anatomy and Biology and its Advisory Board. Science. 1925. Vol.61(1584):473. 70 Donaldson, H.H. Problems in Human Anatomy. Science. 1905. Vol.21(523):20.

37 discussion of anatomical problems was devoted exclusively to the nervous system and its development, his area of expertise. He was interested, also, in questions that “bring anatomy into connection with the problems of sociology” as well as with “the broader problems of zoological relationship and susceptibility to modification.”71 Perhaps most important, Donaldson had a research program concerning the growth of the brain and employing a live animal model, the albino rat, rather than resorting to the dead anatomy of museum study. Greenman might also have been piqued by Donaldson’s desire to apply the fruits of his research to humans: “We must necessarily work to reach human life either through medical practise [sic] or through other avenues of approach, for in the end the object and purpose of all science is to ameliorate the unfavorable conditions which surround man.”72 Greenman knew that Donaldson was fully prepared to carry out exacting experimental research. With the publication of the 1905 paper, Greenman was convinced that Donaldson was philosophically amenable to the Institute.73 Greenman

71 Ibid. p.19. 72 Ibid. p.20-21. 73 Donaldson’s paper was to be read as part of the International Congress of Arts and Sciences at the Louisiana Purchase Exposition in St. Louis (September 19-25, 1904). Personal tragedy, however, kept Donaldson from presenting his paper. His wife, Julia Vaux Donaldson, was suffering from a severe bout of depression, which ended with her suicide by hanging less than six weeks after the Congress. The paper was subsequently published in Science where it was noticed by Clara Perine, Librarian at the Wistar Institute, and brought to the attention of Greenman. William Addison credits the published paper with convincing Greenman to hire Donaldson as Director of Research. The opportunity at the Wistar may also have seemed more attractive to Donaldson following the trauma of his wife’s death in Chicago. Greenman’s courtship of Donaldson still included at least seven meetings between the two men before Donaldson accepted the position. Brosco, J.P. Anatomy and Ambition. p.15 footnote. Addison, W. H. F. Henry Herbert Donaldson 1857-1938. Bios. 1939. Vol.10(1):14-15; “Professor Donaldson’s Wife, Insane, Hangs Herself” New York Times November 11, 1904; Donaldson to Adolf Meyer November 11, 1904 Adolf Meyer Papers Series: I, Folder: 960/8, Alan Mason Chesney Medical Archives, Johns Hopkins Medical Institute, Baltimore, Maryland; Donaldson to Franz Boas November 11, 1904 Franz Boas Papers B.B61 Folder: Donaldson, Henry H. #6, American Philosophical Society, Philadelphia, Pennsylvania.

38 wrote to Edwin Conklin of the institute’s Advisory Board, “We must have the best man and so far as I am able to judge it seems to me that Dr. Donaldson is that man.”74

By 1905 Henry Donaldson had built a solid career and reputation as a neurologist.

He was born May 12, 1857 in Yonkers, New York to Louisa Goddard and John Joseph

Donaldson. His father, a moderately successful banker, moved the family to New York

City when Henry was only two. Henry Donaldson’s biography bears few of the early childhood memories of life on the farm or romps through the woods that seemed formative to many of his contemporaries in American biology. Rather, he remembered his aunt Gertrude supplying him with “endless kittens, which usually ended their career by running around in fits… We never achieved a cat.”75 To rectify this situation and to allow his son some time in the countryside, Henry’s father sent him to spend the summers of 1870 and 1871 with a farmer named Whitney in Maine. “I was to live like a farmer,”

Donaldson recalled, “and, if possible, to regain my health. I fed the cows and milked them… I planted corn and helped about the farm in general.” Finally, his father bought him a pony so that he “should learn to care for a horse.”76 Mercifully, the horse fared better than his kittens.

Donaldson grew to love the outdoors and the study of natural history. His interest in the sciences blossomed during his undergraduate years at Yale, particularly during his independent researches with R.H. Chittenden in physiological chemistry. Following the path that seemed most practical in his generation, Donaldson pursued a medical degree

74 Greenman to Conklin May 12, 1905. Edwin G. Conklin Papers. Series II, Box 9, Folder: Greenman, M.J. 4. Princeton University, Firestone Library, Rare Books and Special Collections. 75 Donaldson, H.H. Memories for My Boys. 1931. p.4. Henry H. Donaldson Papers B.D713M APS. 76 Ibid. p.13-14.

39 from the College of Physicians and Surgeons in New York, but dropped his course when

Johns Hopkins offered him a fellowship in biology in 1881. He specialized in physiology under H. Newell Martin, but, like many early biology students at Hopkins, divided his time between Martin and the more openly theoretical morphologist William Keith

Brooks.77 Thus, Donaldson was exposed not only to exacting research methods, but also to broad theoretical trends in contemporary zoology.78

It was, however, another of the intellectual leaders at Hopkins who would guide

Donaldson to neurology. Professor of Psychology G. Stanley Hall recognized the growing relations between neurology and experimental psychology and needed the help of researchers trained in physiological methods. Hall was committed to the view that psychology was a natural science rather than a branch of metaphysics. He was not, however, prepared to surrender all speculation. Hall’s biographer Dorothy Ross noted that his “tenure in Germany coincided with the great debates over Haeckel’s biogenetic law. Hall was attracted to Haeckel’s ideas despite the conflict they incited with his mentors in physiology: du Bois-Reymond, Ludwig, Helmholz, etc.”79 Hall was enchanted with the historicized evolution implied in German idealistic philosophy.

“Evolution always implied normative conceptions for Hall,” wrote Ross, “and always centered on the historical process of development over time, rather than on the functional

77 Maienschein, J. Transforming Traditions in American Biology 1880-1915. Baltimore: Johns Hopkins University Press, 1991. p.37-59. Donaldson also spent a portion of 1884 in special investigations with Brooks in his semi-nomadic summer laboratory, at the time stationed in Beaufort, North Carolina. Conklin, E.G. Biographical Memoir of Henry Herbert Donaldson, 1857-1938. National Academy of Sciences of the United States of America Biographical Memoirs. 1938. Vol.20:230. 78 Donaldson, H.H. Memories for My Boys. p.72. 79 Ross, D. G. Stanley Hall: The Psychologist as Prophet. Chicago: University of Chicago Press, 1972. p.89-90.

40 process of selection and adaptation.”80 Like the German idealists and the evolutionists that prevailed in his home country, Hall fixated on development as the key to understanding the past and future trends of evolution. His most famous work concerned the psychology of the developing infant, child, and adolescent. His essay “Evolution and

Psychology” equated childhood with the psychological state of humankind’s phylogenetic ancestor: “…the child is vastly more ancient than the man… adulthood is comparatively a novel structure built upon very antique foundations.”81 What was needed was an “embryology of the soul” to understand the present condition and evolutionary trajectory of the human mind.82

Hall spent much of his early career trying to create a fundamentally biological psychology. His approach attracted some of the brightest students at Hopkins, including

Donaldson, James McKeen Cattell, Joseph Jastrow, E.M. Hartwell, and John Dewey. Of these, Donaldson is remarkable for his joint publication of experimental results in physiological psychology with Hall, who published little else in the field despite his fierce advocacy for it.83 Donaldson took his Ph.D. under Hall and adopted his mentor’s interest in the relations of neurology and psychology. Hall encouraged Donaldson to expand his studies of the nervous system with a visit to the leading neurologists of

Europe, including Brown-Sequard, von Gudden, Nissl, Forel, Golgi, and Flechsig among

80 Ibid. p.93. 81 Hall, G.S. Evolution and Psychology. In Fifty Years of Darwinism: Modern Aspects of Evolution. American Association for the Advancement of Science. New York: Henry Holt and Company, 1909. p.262. 82 Ibid. p.263. 83 Hall, G.S. and Donaldson, H.H. Motor Sensations of the Skin. Mind. 1886. Vol.10(1):557-572.

41 others.84 Shortly after Donaldson’s return Hall was named President of the new Clark

University in Worchester, Massachusetts. Hall acquired Donaldson for his selective and highly regarded faculty. Money troubles and administrative disagreements came between

Hall and his faculty shortly after the university’s founding, which precipitated a mass exodus of leading scholars, including Donaldson, from Clark to the new University of

Chicago in 1892. Despite the strain of their later relationship, Donaldson remembered

Hall as “a man of unusual mental gifts and the most suggestive and stimulating person with whom I ever came in contact.”85

Hall may also be credited with inspiring the research question that defined

Donaldson’s career. In 1889 Hall gave Donaldson the opportunity to dissect and describe the brain of Laura Bridgman, an educated, blind, deaf, and mute woman who had died at the age of 60. Bridgman had been relatively famous in her life, due in part to the attention she received from the scientific community, including Hall. After an exhaustive study of Bridgman’s brain and the relevant literature, Donaldson noted that no clear account of the developing human brain existed: “Here was a gap- a gap covering the period of the development of the mind. It caught my attention, and the desire to help in closing this gap dominated my subsequent work.”86

Donaldson set to work accumulating the existing knowledge on the development of the human brain and the brains of other animals. The resulting publication, The

Growth of the Brain: A Study of the Nervous System in Relation to Education, debuted in

84 Addison, W.H.F. Henry Herbert Donaldson. p.12. Von Gudden accepted Donaldson into his laboratory, but Donaldson’s time with von Gudden was cut short by the latter’s mysterious drowning along with his most famous patient, King Ludwig II, on June 13, 1886. 85 Donaldson, H.H. Memories for My Boys. p.90. 86 Donaldson, H.H. Research at the Wistar Institute, 1905-1925. Science. 1925. Vol.61(1584):480.

1895 and highlighted Donaldson’s familiarity with biological analogizing and his inclination to see the human sociological implications of his research. In the book

Donaldson showed more sensitivity to the functional aspects of evolution than his mentor, but the discussion remained rooted in ontogenetic analogy: “[T]he suggestion here is that species, like individuals, have a period of growth followed by specialization, which latter is in the end the cause of their extinction, through the loss of adaptability to new conditions.”87 Donaldson also extended the analogy by linking the fate of species with the fate of cells in normal development.88 He retained his mentor’s verve for evolutionary explanations and sociological applications, but was generally more reserved in his interpretations of neuroanatomical research. The Growth of the Brain may not have achieved Hall’s “embryology of the soul,” but it was, fairly, the embryology of the seat of the soul. This, though, was precisely the goal that John Adam Ryder had imagined for his synthetic research museum: to grasp “becoming the brain.”89

The development of the human nervous system became the focus of Donaldson’s subsequent career. His comparative-developmental approach was intended to historicize neuroanatomy as G. Stanley Hall had done with developmental psychology. At its heart,

Donaldson’s concept of evolution was driven by incremental changes in the pattern of development. Thus, when in his early studies he used a broad array of species including sharks, bony fishes, frogs, owls, dogs, and monkeys in addition to human material, it seemed that he was approaching his desired subject, the human brain, step by step.

87 Donaldson, H.H. The Growth of the Brain: A Study of the Nervous System in Relation to Education 2nd edition. New York: Charles Scribner’s Sons, 1907. p.38. 88 Ibid. p.32. 89 Ryder, J.A. The Synthetic Museum of Comparative Anatomy. p.9.

Donaldson suggested to his close friend Franz Boas that human races could likewise be arranged in such a progressive phylogeny, with “the great point of divergence

[beginning] at adolescence, and the inference is fairly good that we shall not find in the brains of the lower races the post-pubertal growth in the cortex.”90 It was through his animals that Donaldson was able to speculate about human evolution. Near the time that

Donaldson was picked to lead research at the Wistar Institute, he felt he had found the ideal comparison for the developing human: the developing albino rat.

Donaldson is best known for his efforts to standardize the albino rat for its use in the laboratory. His laboratory at the University of Chicago was using rats by 1896, though he claimed that he had been introduced to the animals three years earlier by the

Swiss-American psychiatrist Adolf Meyer. Historian of science Cheryl Logan argues that Donaldson’s comparative-developmental perspective lay behind his interest in developing the albino rat into a laboratory standard. “Donaldson’s rats,” notes Logan,

“were not a generic vertebrate standard, they were a model for humans.”91 He was most intrigued by the parallels in the growth patterns of rats and humans, as indicated in a note to Boas just prior to his arrival at the Wistar Institute:

The growth curve for [the albino rat] goes through all the phases characteristic of the human curve… It is the humanist [sic] thing you could wish- of course it is for me merely a basis for the study of the growth of the central nervous system, but it is a great satisfaction to learn that the two animals are so similar in the way they gain weight for it facilitates the further comparisons greatly.92

90 Donaldson to Boas, June 30, 1894. Franz Boas Papers B.B61, Folder: Henry H. Donaldson #1, APS. 91 Logan, C.A. The Altered Rationale. p.10. 92 Donaldson to Boas July 9, 1905 Franz Boas Papers B.B61, Folder: Henry H. Donaldson #7, APS.

His respect for biological diversity intact, Donaldson continued to use other animals, particularly frogs, in his comparative research after his move to the Wistar

Institute. Norway rats, however, came to dominate his subsequent studies. Once

Donaldson had committed to the rat, he sought an encyclopedic knowledge of the animal.

Before leaving Chicago he initiated investigations along with some of his brightest students, John B. Watson, Irving Hardesty, and Shinkishi Hatai, into the provenance, physiology, development, and variability of the rat. Hatai, who accompanied Donaldson to the Wistar Institute, completed some of the most substantial research into the nature and background of the albino rat. His investigations highlight some of the early concerns with the animal, including of course nervous growth, but also nutrient cycling, whole- body growth curves, nervous structure, and taxanomic placement. Hatai published a brief paper, “On the Zoological Position of the Albino Rat,” in 1907.93 In it he identified the albino rat as a sport of the common gray rat (Mus norvegicus)94 rather than of the black rat (Mus rattus) by a comparison of external and skull morphologies. He also suggested that some of the albino’s peculiar morphologies could be associated with captivity and confinement, but stopped short of claiming that the animal had been domesticated.

Donaldson published an update to Hatai’s paper five years later. He reaffirmed Hatai’s taxonomy, but he noted more of the particulars, behavioral as well as morphological, that distinguished the albino even from the gray rat. He declared, “We are inclined to regard

93 Hatai, S. On the Zoological Position of the Albino Rat. Biological Bulletin. 1907. Vol.12(4):266-273. 94 The taxonomy of the Norway and black rats has shifted since Donaldson’s earliest work on the species. Ned Hollister found that priority for a distinct generic epithet for the rat belonged to Fischer (1803), thus in 1916 the genus name changed to Rattus and the specific epithets remained the same for both the Norway and black rats. Hollister, N. The Generic Names Epimys and Rattus. Proceedings of the Biological Society of Washington. 1916. Vol.29:126.

45 these alterations as the result of domestication.”95 Donaldson was well aware that

“domestication” described a complex of phenomena rather than a singular biological or psychological event. The term, he felt, aptly described the co-occurrence of behavioral and morphological differences between the albino and gray Norway rat. In the five years between the publication of Hatai’s paper and of Donaldson’s, the differences between the albino and gray Norway rats had become a central concern at the Wistar Institute and domestication a key explanation for these changes.96

The work to determine the zoological position of the albino rat precipitated the comparisons between the albino and gray forms of the species. Hatai had noticed that the albino was slightly smaller overall than the gray. His subsequent work found that the brain and spinal cord weights of the albino were substantially less than those of grays caught near Philadelphia, “animals of like body weight and body length being compared.”97 Donaldson and Hatai reported that these data agreed with the findings of

Darwin (1883) on domestic and wild rabbits and those of Louis Lapicque and Pierre

Girard (1907) on domestic and wild rabbits, ruminants, ducks, and fowl.98 The distinctions between Donaldson’s chosen research vehicle, the albino rat, and its wild

95 Donaldson, H. The History and Zoological Position of the Albino Rat. Journal of The Academy of Natural Sciences of Philadelphia. 1912. Vol.15:368. 96 Donaldson had mentioned the potential importance of domestication as early as 1905 in an internal memorandum to the Advisory Board. He and his associates at the Wistar Institute, however, spent several more years cataloguing the basic differences between albino and gray Norway rats before they publicly cited domestication as a root cause of these distinctions. Donaldson, H.H. To the Advisory Board of the Wistar Institute of Anatomy and Biology. Enclosed in a letter from Greenman to Conklin May 12, 1905 Edwin G. Conklin Papers, Series II, Box 9, Folder Greenman, M.J. #4, Princeton University, Firestone Library, Rare Books and Special Collections. 97 Donaldson, H.H. Appendix C. In Director’s Report of The Wistar Institute for the Year 1909. Wistar Institute Library. Philadelphia, Pennsylvania. 98 Donaldson, H.H. and Hatai, S. A Comparison of the Norway Rat with the Albino Rat in Respect to body Length, Brain Weight, Spinal Cord Weight, and the Percentage of Water in Both the Brain and Spinal Cord. Journal of Comparative Neurology. 1911. Vol.21(3):455.

46 progenitor, the gray rat, now squarely affected his research program on the development of the nervous system.

By 1909 Donaldson seemed resolved that these distinctions were best explained by domestication. In April of that year he left Philadelphia for a four-month tour of

Europe that included the second meeting of the International Brain Commission in Zurich and stops at research stations in Vienna, Paris, and London. At the latter three stations he visited the laboratories of Heinrich Obersteiner, Hans Przibram, Lapicque, and Victor

Horsley to determine whether Norway rats varied appreciably between Philadelphia and cities in Western Europe. Donaldson sought to demonstrate that his albinos were not simply derived from an anomalous population of Norway rats but were, rather, domesticated.99 He found that the populations of gray rats were not substantially different for any of the measurements taken, but they regularly differed from the condition of albino rats from the same region. Furthermore, the differences between albinos and grays were largely in the same direction. “A very satisfactory result from my point of view,” he wrote later.100 Donaldson concluded such regularity, irrespective of location, could best be explained by a common cause, domestication. Meanwhile, in

Philadelphia, Hatai initiated preliminary experiments to determine the effects of various factors of domestication.

From the beginning Donaldson conceived of the experiments on domestication as part of an integrated research program. Although he found it convenient to assign the

99 Donaldson, H.H. A Comparison of the European and Albino Rats (Mus norvegicus and Mus norvegicus albinus) with Those of North America in Respect to the Weight of the Central Nervous System and to Cranial Capacity. Journal of Comparative Neurology. 1912. Vol.22(1):71. 100 Donaldson, H.H. Memories for My Boys. p.118.

47 myriad changes between the gray and albino Norway rat to “domestication,” Donaldson recognized that the term likely summarized a host of factors:

Darwin has already called attention to the smaller cranial capacity of domesticated animals and it seems fair to look on our albino as a domesticated form. If we analyze the problem then, the nervous system in the Albino may be smaller 1. As the result of albinism. 2. From lack of exercise. 3. As an effect of interbreeding. 4. As a result of the absence of the “struggle for existence.101

To these Donaldson and his associates later included inbreeding and other potential factors. Donaldson frequently reminded his staff that the domestication program, though crucial to the work of the Institute, was subsidiary to the larger goal of describing the growth of the central nervous system. The new program, however, proved difficult to contain. The work on putative factors of domestication demanded more and more of the Institute’s time and resources, much of it for the subject’s intrinsic appeal.

Donaldson was surprised that since Darwin’s observations on domestic and wild rabbits few had pursued the question of domestication’s effects on growth and structure until

Lapicque and Girard (1907). In truth, others had been interested in the question, but their contributions often relied on empirical observations, analogies, and speculation. If domestication was responsible for the changes leading from the gray to the albino rat, as

Donaldson was convinced, the collective wisdom concerning domestication would need to be deconstructed, demythologized, and treated as a series of testable hypotheses. “For this purpose we have attempted to analyze the condition of domestication into its

101 Donaldson, H.H. Appendix C. In Director’s Report of The Wistar Institute for the Year 1909.

48 elements with a view to testing these one by one,” Donaldson wrote in 1911.102 The existing wisdom, however, framed Donaldson’s understanding of domestication and that of his colleagues and contemporaries.

Domestication was not merely a convenient explanation for the varieties of

Donaldson’s rats. It seemed to suggest itself as a subject of study amenable to both the biological philosophy of the Wistar and its relatively recent experimental research tradition. It appealed especially to two longstanding emphases of the Institute: dynamic development and the human and sociological relevance of research findings.

Donaldson had selected rats because, like humans, their development was altricial

(i.e. gradual, from a helpless infancy through adolescence and finally into adulthood). He thought that this period of extended plasticity made rats (and humans) such malleable and adaptable creatures. Domestication, it was widely assumed, rendered animals extraordinarily variable through its effects on development. Donaldson pointed to a “lack of growth” to explain the differences between his gray and albino rats and thought it

“most natural to attribute the lack of growth to the whole set of conditions summed up in the word ‘domestication.’”103 Donaldson’s prior concern with the rigidity that accompanied specialization perhaps convinced him that domestication preserved the plasticity of an incomplete development. Changes continued to accumulate in the course of his studies, changes that he claimed, “…indicate that there are fluctuations in character both on age and on body weight, and also that there may be a drift or trend in the form and functions of the albino rat as domestication progresses.” He added, “Such a drift is

102 Donaldson On the Influence of Exercise on the Weight of the Central Nervous System of the Albino Rat. Journal of Comparative Neurology. 1911. Vol.21(2): p.129. 103 Donaldson, H.H. The Rat: Data and Reference Tables. Second Edition. Philadelphia: Wistar Institute of Anatomy and Biology, 1924. p.340.

49 sometimes designated as orthogenesis.”104 Harrison Allen, whose suggestions for experiments on domestication preceded Donaldson’s own by more than a decade, likewise found domesticated animals useful for illustrating orthogenesis, a teleological evolutionism that likened the development of the individual to the fate of the species. He recommended comparisons between domesticated cattle and their progenitor, the aurochs, to uncover symptoms of impending extinction in the latter and also suggested that the domestic dog “is of exceptional value in the philosophic study of senility and diseased action.”105 The assumedly recent occurrence of domestication and persistence of the ancestral types seemed to make the comparison between ontogeny and phylogeny not only theoretically possible but also practicable.

What appealed most to Donaldson, and to Allen before him, was the notion that domestication was a dynamic and largely directional phenomenon. That life was never static was one of Donaldson’s most steadfast beliefs, from the work that launched his career: “The living world of which we are part is ever changing;”106 to the project that consumed his later years: “In the very nature of [producing standard laboratory animals] such accuracy and constancy is unattainable for all animals are at all times in flux.”107

He approached domestication with this view in mind. His experiments were meant to illuminate the process that separated the wild and domesticated rat rather than the static differences between types. The synthetic anatomy favored by Donaldson’s intellectual

104 Donaldson, H.H. The Rat. 2nd edition. p.viii-ix. 105 Donaldson and Allen seemed to have held different opinions about the specific effects of domestication, and it is probable that Donaldson never read about Allen’s proposed experiments on the subject. Both, however, found domestication useful for the reason that it ostensibly illustrated orthogenetic evolution on a shortened time scale. Allen, H. On the Effects of Disease and Senility. p.292. 106 Donaldson, H.H. The Growth of the Brain. p.21. 107 Donaldson, H.H. The Rat. 2nd Edition. p.xi.

50 predecessors at the Institute, Ryder and Allen, stressed the becoming of structure and function rather than their present conditions. This view made use of dynamic processes, namely development and its cognate evolution, to explain the past and future course of a particular morphology. Donaldson was of a similar biological philosophy. For him, domestication offered a like and related process by which to understand the change- still in progress- that distinguished his albino rats from their wild, gray counterparts. Given

Donaldson’s refrain of “human applications,” domestication also became a window into the change that he and others perceived in the human species.

Donaldson wrote in the preface to the second edition of The Rat (1924): “[I]n pointing out some of [the rat’s] similarities to man, it is not intended to convey the notion that the rat is a bewitched prince or that man is an overgrown rat, but merely to emphasize the accepted view that the similarities between mammals having the same food habits tend to be close.”108 He was not, however, as vigilant to guard against unwarranted analogizing as this statement might suggest. Much more common were the exuberances of Donaldson and his staff that “there is every reason to prepare for the application of these results [from rats] to man.”109 He emphasized not only the anatomical significance of his findings, but also their evolutionary and sociological relevance. His sociological interests were directed primarily toward improvement of human mental health and ability, and the rat was involved from an early point in his career with his eugenical ambitions. In the 1905 paper that convinced Greenman to bring

Donaldson to the Wistar Institute, Donaldson suggested that his early work on the albino

108 Ibid. p.xiv. 109 Donaldson, H.H. Problems in human Anatomy. p.23.

51 rat “would lead ultimately to the attempt to breed animals with improved nervous systems in which we shall know the nature of the improvement in considerable detail.”110

In the same paper he identified as one of the goals of modern science “to produce a human individual more capable of resistance to disturbing influences, and better suited for the enjoyment of the world in which he lives.”111 Human and social concerns influenced much of Donaldson’s work with the Norway rat, and the knowledge thereby gained was, not surprisingly, interpreted to have human and social significance.

Donaldson’s tenure at the University of Chicago had prepared him to examine the sociological significance of his work on the nervous systems of animals. Scholars at the

University stressed continuity rather than distinctions between biological and cultural phenomena, seeking fundamental concepts that underlay both.112 The members of the early faculty were, for the most part, broadly philosophical and committed to some manner of progressive evolutionism. These shared commitments and the formation of the faculty-wide Quadrangle Club in 1893 eased efforts toward collaboration and cross- disciplinary approaches. Many on the University faculty, most notably John Dewey and

George Herbert Mead, were also intimately engaged in Chicago’s civic life. Donaldson, despite his reputation as a reticent and “almost calculatedly uncontroversial figure,” played a crucial role in fostering this unique atmosphere of scholarship and

110 Ibid. p.25. 111 Ibid. p.21. There may have been a personal dimension to Donaldson’s call to produce a human less susceptible to “disturbing influences.” Two months prior to the publication of “Problems in human Anatomy” his wife succumbed to “torturing thoughts- the fictions of an unbalanced mind” in Donaldson’s words and hanged herself. Donaldson to Boas November 11, 1904 Franz Boas Papers, APS. The preface to Donaldson’s Growth of the Brain also suggests that he viewed the relationship between nervous development and modern civilization through the lens of contemporary theories concerning nervous strain and neurasthenia. Donaldson, H. H. The Growth of the Brain. p.5. 112 Pauly, P.J. Controlling Life: Jacques Loeb and the Engineering Ideal in Biology. Oxford: Oxford University Press, 1987. p.67. Ironically, it was Donaldson’s friend and fellow Clark apostate, Boas, who ultimately undermined this approach.

52 application.113 He was among the founders of the Quandrangle Club and served as its second President. With his wife, he became involved in the University of Chicago

Settlement, whose objective was to “[introduce] the student of theories and principles to the laboratory of human experience.”114 He also served with Mead and Dewey on the

Board of the Chicago Hospital-School for Nervous and Delicate Children (formerly the

Chicago Physiological School), the curriculum of which was established through a collaboration of Donaldson’s Department of Neurology and Dewey and Mead’s

Department of Philosophy.115 Once in Philadelphia, Donaldson sought similar ways to combine his neurological expertise and progressive social ideals.116

As Donaldson began to formulate his experimental domestication program at the

Wistar Institute, he took cues from experimentalists and social scientists who had already explored the putative similarity of animal domestication and human civilization. Willard

S. Small, a comparative psychologist in Donaldson’s former department at Clark

University, attributed the behavioral differences between albino and gray Norway rats to domestication in 1900.117 The point particularly intrigued him because “[i]t suggests the

113 Characterization quoted from: Logan, C.A. “[A]re Norway Rats… Things.” p.301. 114 McDowell, M.E. The University of Chicago Settlement. Chicago, s.n., 1901. p.24. Donaldson delivered lectures at the Settlement and his wife served as Vice President of the University of Chicago Settlement League. 115 American News and Notes: The Chicago Hospital-School for Nervous and Delicate Children. The Philadelphia Medical Journal. 1901. Vol.2(13):596. 116 In the year of his arrival in Philadelphia, Donaldson was named a member of the “Feeble-minded Club,” an informal society whose leaders included E.R. Johnstone and H.H. Goddard of the Vineland Training School in New Jersey, Philadelphia educator Earl Barnes, and soap magnate and social reformer, Samuel S. Fels. See Zenderland, Leila Measuring Minds: Henry H. Goddard and the Origins of American Intelligence Testing. Cambridge: Cambridge University Press, 2001. p.61-62 117 Small, W.S. Experimental Study of the Mental Processes of the Rat. American Journal of Psychology. 1900. Vol.11(2):133-165. Donaldson knew of Small’s work by 1915 when he published his book The Rat complete with an exhaustive bibliography of researches completed on the animal. He may have known of the work at an earlier period thanks to his correspondence with Adolf Meyer

53 very live question whether civilization, which is a kind of domestication, operates similarly with the human species.”118 The conceptual link between domestication and civilization, however, predated its use by comparative psychologists. Anthropologists, including Johann Friedrich Blumenbach, James Cowles Prichard, William Lawrence,

Gustav Fritsch, and Donaldson’s predecessor at the Wistar, Harrison Allen, had developed tropes and sometimes more literal suggestions that compared primitive and civilized humans with wild and domesticated animals, respectively, from the late 18th century to the close of the 19th.

This line of thinking came closer into Donaldson’s purview in 1911, when his friend Franz Boas published The Mind of Primitive Man. Boas followed Fritsch in declaring that “[the] environment has an important effect upon the anatomical structure and physiological functions of man… It seems plausible that one of the most important causes of these modifications must be looked for in the progressive domestication of man incident to the advance of civilization.”119 Boas also mentioned that domestication modified mental traits as easily as physical traits, something that would have appealed to

Donaldson’s twin interests in neurology and psychology.120 Boas’s book did not inspire

Donaldson to think of domestication. By 1911, he had already begun preliminary experiments to determine its effects. Later that same year, however, Donaldson noted that the his morphological comparisons between albino from gray rats “correspond with

who worked as an adjunct professor in the Clark Psychology Department at the time Small was conducting his studies on rats. 118 Small, W.S. Experimental Study of the Mental Processes of the Rat II. American Journal of Psychology. 1901. Vol.12(2):212. 119 Boas, F. The Mind of Primitive Man. New York: MacMillan & Company, 1911. p.75. 120 Ibid. p.74-75.

54 what anthropologists usually find in the different races of man.”121 This exchange of ideas (anthropologists, including Boas and Ales Hrdlicka, recognized the work on rats coming from Donaldson’s laboratory as relevant for their own work in the science of man) reaffirmed for Donaldson that his work held special significance for modern humans and their conditions of life.122

Donaldson was committed to his program of experimental domestication by 1912, though it was then only in its early stages. The subject seemed to condense the broad and historical concerns of transcendental morphology into an observable and, indeed, testable set of phenomena. Thus, Donaldson believed that he was bringing the Wistar Institute, and with it the field of anatomy, into the modern era of biological research. In so doing he preserved much of the biological philosophy that preceded him at the Wistar Institute, including concern for diversity over generality, the temporal space described by long- term dynamic processes (rather than briefly imposed and seemingly static conditions), phylogenetic context, and human implications. He did this, of course, with little conscious effort because they were the same concerns that drove him, that described the biological questions that interested him. That he shared these views with the present and former leadership of the Wistar Institute was not likely a coincidence, but was probably the reason Milton Greenman hired him to guide the research arm of the Institute, thus fulfilling the vision of Greenman’s friends and mentors: Isaac Wistar, Harrison Allen, and John Adam Ryder.

121 Donaldson, H.H. and Hatai, S. A Comparison of the Norway Rat with the Albino Rat in Respect to Body Length, Brain Weight, Spinal Cord Weight, and the Percentage of Water in Both the Brain and Spinal Cord. Journal of Comparative Neurology. 1911. Vol.21(3):417-458. 122 See Hrdlicka, A. Physical Anthropology Its Scope and Aims: Its History and Present Status in the United States. Philadelphia: Wistar Institute of Anatomy and Biology, 1919. p.110-111; Boas, F. Race, Language, and Culture Revised Edition. Chicago: Chicago University Press, 1982. p.48, 162.

This should not be taken to mean that Greenman and Donaldson (or for that matter, any of their predecessors) were of exactly like mind. As Bonnie Clause has shown, Greenman had by 1908 adopted the gospel of efficiency and standardization espoused by fellow Philadelphian Frederick Winslow Taylor and sought to apply these principles to the development of a standard albino rat.123 Cheryl Logan has further suggested Donaldson’s latent discomfort with Greenman’s approach toward standardizing the albino rat, which she identifies with Philip J. Pauly’s concept of the engineering ideal in modern biology (that is, an ideal that places prediction and control above the traditional concerns of explaining adaptive function and phylogenetic relations as the primary objects of modern biology).124 Donaldson worried privately in 1910 that

“G.[reenman] cannot adjust to the vacillations of biological work- the regularity of machines is more to his mind.”125

There is reason to believe, however, that Donaldson and Greenman would have understood the implications of the engineering ideal differently. Donaldson had been witness to the philosophical disputes at the University of Chicago that formed the backdrop for Pauly’s historiographic concept of the engineering ideal. These feuds pitted

Jacques Loeb and his emphases on control and generality against the University’s core of progressive evolutionists (including Donaldson) and their emphases on diversity, adaptive specificity, and phylogenetic history. Where Greenman could blithely embrace

123 Clause, B.T. The Wistar Rat as a Right Choice. p.341-343. 124 Logan, C.A. Altered Rationale. p.15-16. Logan and Clause both agree, however, that Donaldson gradually aligned himself with the goal of producing standardized, “perfect animals” for research purposes. After examining many generations of closely inbred rats whose conditions of life had been carefully controlled, Donaldson finally hedged and admitted that perfect standardization may not be possible with a living organism. For an explanation of the engineering ideal in modern American biology, see Pauly, P.J. Controlling Life. 125 Quoted from Logan, C.A. “[A]re Norway Rats… Things?” p.305, footnote.

56 the power of analogizing the standardized albino rat to the uniformity and interchangeability of industrial components, Donaldson could not. Indeed, Greenman may not have recognized the potential incompatibility of Taylorism and the tradition of transcendental morphology in which he was trained by Ryder, Allen, and Jayne at Penn and in the early years of the Wistar Institute. That is, in adopting the one, he did not abandon the other.

Greenman’s pet project at the Institute, aside from the effort to standardize the rat, was supporting research into the development of the Virginia opossum (Didelphys virginiana), regarded in the literature of the Institute as interesting for its extreme

“primitiveness.” He supported researches on the opossum for nearly 30 years at the

Wistar Institute and beyond, including the building of several breeding colonies, the last of which was established at a 150 acre experimental farm associated with the Institute, where he also made his home for the last years of his life. The rationale for the experiments consistently cited the opossum’s unique phylogenetic position, or as one of

Greenman’s beneficiaries explained: “The study of a particularly primitive mammal leads one almost irresistibly to the subject of mammalian phylogeny.”126 This rationale would have been unintelligible to Loeb.

How might we reconcile Greenman’s progressive evolutionary rationale and the ideals he adopted in order to establish an experimental tradition at the Wistar Institute? It may be that Greenman, never exposed, as Donaldson was, to the internecine fights over competing biological ideals at the University of Chicago, did not view his prior

126 McCrady Jr., E. The Embryology of the Opossum. American Anatomical Memoirs. 1938. No.16:207. McCrady also refers to Greenman’s special enthusiasm for studies on the opossum. Donaldson also cited this particular interest in his memorial piece dedicated to Greenman. Donaldson, H.H. Milton Jay Greenman. The Anatomical Record. 1937. Vol68(3):265.

57 philosophical preparation and newly adopted ethos of control and efficiency as incongruous- at least not initially. The institutional context of the Wistar Institute allowed Greenman to avoid the sense that these views were dichotomous. As Clause has indicated, “With its ties to medicine, museums, and experimental science, the Wistar

Institute was an interesting and somewhat enigmatic hybrid in the era when biology was being defined.”127 In its blend of medicine, museum work (i.e. morphology), and experimental research, however, the Wistar Institute was perhaps less idiosyncratic and more representative of the unsettled direction of American biology in the early years of the 20th century. The Institute’s fiercely guarded independence (a feature demanded by

Isaac Wistar and Harrison Allen upon its founding) and Greenman’s sometimes autocratic leadership style may have had more to do with the particular combination of

19th century biological philosophy and 20th century experimental ethos that prevailed there.

It is significant that Donaldson’s domestication program paralleled the standardization of the albino rat at the Wistar Institute in the first decades of the 20th century. Widespread concerns in contemporary biology about the place of basic inquiry into nature and the demand for control over the processes studied were reflected in both of these efforts at the Institute. As Clause has shown, the desire for control and predictability was central to the experimental ethos that Greenman instituted at the Wistar and to his rationale for standardizing the albino rat. However, by the early 1930s, as

Wistar rats had become generalized models for mammalian physiology and behavior in

127 Clause, B.T. The Wistar Rat as a Right Choice. p.333 footnote.

58 laboratories throughout the world, Greenman was forced to admit that the degree of uniformity and the condition of rats in the standard colony

…were not what we had expected [e]ven with a diet as well balanced and uniform as may be devised, with environmental conditions as uniform as may be secured,… and many other conditions of a subtle nature which influence a growing animal. All these together with the variable and undeterminable state or conditions of the albino rat itself in its reactions to food and to external influences, result in a constantly changing organism.128

Donaldson’s domestication program raised similar concerns over the extent to which control was appropriate or even achievable given the dynamic conception of life at the Institute. Perhaps this is not surprising, as domestication had long had a tenuous association with control. It was considered synonymous with mastery over nature, but as the experience of countless breeders demonstrated, this was an elusive ideal. Control, not only experimental, but philosophical and administrative as well, played a crucial role in the fate of the domestication program at the Wistar Institute, an account of which follows.

Analogy in Practice: The Wistar Institute Domestication Program and its Social Applications.

Biological interest in domestication was by no means new in the first decade of the 20th century when Donaldson settled on pursuing the topic. Several others had even anticipated the concept that the effects of domestication could be experimentally assessed, Charles Darwin chief among them.129 The Wistar Institute, in its brief history

128 Quoted from Clause, B.T. The Wistar Rat as a Right Choice. p.348. 129 Darwin, C.R. Origin p.96-97. Darwin suggested mating a “wild animal with a highly domesticated one,” specifically a fox and a hound to pointer-bitches and then breeding the progeny back to the

59 prior to Donaldson’s arrival, already boasted two advocates of intensive studies on domestication. In 1897, Harrison Allen proposed keeping wild animals in captivity to determine the effects of domestication on ontogenetic development and pathology.130

Simon Henry Gage of the Institute’s permanent Advisory Board had written about the promise of domestication research on the eve of the 20th century. Although Gage wrote on the subject seven years before his association with the Institute, the questions about domestication research that framed his discussion paralleled the modus operandi and agenda for human application that characterized the Wistar Institute: “(1) What has been and what is likely to be the influence of the study of the domestic animals upon the doctrine of the evolution of organic forms? (2) What has the study of them contributed in comparative anatomy, embryology, and physiology? (3) What has been the contribution in hygiene and preventive medicine? (4) And finally what should be their influence in theories of heredity and sociology?”131 He answered that their contributions would be great in all of these fields in the new century.

When Donaldson began thinking in similar terms, however, he could consult few precedents that had done more than speculate about the value of experiments on domestication. Perhaps this is understandable. Laboratory experimentation had only come into vogue in biology in Donaldson’s generation, expanding from its earlier use in physiology to address a range of problems in the fields of zoology, embryology, and comparative psychology among others. Donaldson had started his tenure at the Wistar pointers, in: Questions about the Breeding of Animals [1840] Facsimile. London: Society for the Bibliography of Natural History, 1968. p.3. 130 Allen, H. On the Teaching of Anatomy. p.86-87. Also see Allen, H. On the Effects of Disease and Senility. p.294 131 Gage, S.H. The Importance and the Promise in the Study of the Domestic Animals. Science. 1899. Vol10(245):306.

Institute believing that he would pursue a single overarching research question: the growth of the human brain. By then he had already determined that the albino rat would be the best model for human nervous development, owing to its similar but more expedient patterns of growth. His interest in domestication grew, first, from his endeavor to determine the nature and suitability of his model organism and, second, from domestication’s perceived value in answering basic questions about heredity and evolution. He was also aware of the analogy between domestication and civilization in anthropology, whose champions included Allen, his predecessor at Wistar, and one of his closest friends, the anthropologist Franz Boas. Domestication, thus, rose in Donaldson’s esteem not only in connection with the problem of neurological growth, but also in relation to his broader interests in evolution, anatomy, medicine, psychology, and sociology.

So little was known about the nature of albino rats when Donaldson began using them as models of human neurological growth that much of his early work with them sought answers to the most basic questions of provenance and zoological relation.

Shinkishi Hatai, Donaldson’s one-time student at the University of Chicago and who continued as his assistant at the Wistar Institute, determined by breeding tests and comparative anatomical methods that the albino rat that had come into use in

Donaldson’s laboratory and in a small but growing number of others worldwide was derived from the gray Norway (Mus norvegicus) and not the black (Mus rattus) or

Alexandrine (Mus alexandrinus) rat. This finding precipitated further comparisons between the gray and albino varieties of the Norway rat, which revealed marked differences in particular anatomical points. Albinos were somewhat smaller overall than

61 the grays, their adrenal glands were dramatically smaller, their pituitaries were slightly larger, and most significantly for Donaldson, their brains weighed on average 12 per cent less than those of the grays. He attributed these changes to one of the most evident distinctions between the forms: the albino was apparently domesticated while the gray was by all accounts a wild animal.

The concept of domestication as a metaphenomenon drove Donaldson to initiate a series of studies on its constituent and associated factors. The domestication program he developed at the Wistar Institute subsequently evolved in concept and practice as this complex problem was teased apart into its elements. In his first attempt to enumerate these factors in 1905 he named: albinism, lack of exercise, interbreeding, and “absence of the struggle for existence.”132 Donaldson and his associates designed experiments to clarify the roles of each of these factors in haste. Several approaches were required to evaluate some factors. Experimentally assessing “the struggle for existence,” for example, was realized by both releasing tame albino rats into the wild and bringing wild gray rats into captivity and was variously explained by selection pressures and the action of the environment, reflecting in part Donaldson’s mélange of evolutionary perspectives.

These experiments were developed almost simultaneously to address the questions raised by the key distinctions between Donaldson’s tame albino rats and their wild gray counterparts, and, thus, constituted a single program of research on domestication.

132 Donaldson, H.H. “To the Advisory Board of the Wistar Institute of Anatomy and Biology.” This memorandum is among the Edwin Grant Conklin Papers. Series II, Box 9, Folder 4 “Greenman, M.J..” Division of Rare Books and Special Collections, Princeton University Library. Princeton, University Library (Hereafter PUL).

Feral Rats, Captive Rats: A First Look at Domestication

While Donaldson may have determined the shape of the domestication program, its execution eventually fell to his assistants, primarily Shinkishi Hatai and Helen Dean

King. In the years around 1910 Hatai seemed especially overburdened with the responsibilities of the program. In addition to several fundamental studies on rat neuroanatomy, Hatai’s reports to Milton Greenman in the years 1910 and 1911 mentioned his role in four separate experiments linked to the domestication program: on hybridization, on inbreeding (with Helen Dean King), on breeding in captivity, and on feralization, or an attempt “to determine whether life in the open would restore the central nervous system of the albino to the relative weight which it holds in the grey rat.”133

This last experiment was conceived in tandem with the effort to breed wild gray rats in captivity in order to determine if the domesticative process would be 1) observable and 2) reversible. Its counterpart, variously named for caging, captive breeding, or the whole phenomenon of domestication, may give the false impression of encompassing the problem. It was, however, only one element of Donaldson’s program to understand the many effects of domestication. The feralization project initially received equal if not more attention than the captive breeding study. This may have been as much for practical expedience as for any other reason. It initially seemed easier, or at the very least, less costly to deposit colonies of albino rats in various open locations and to check on them

133 Greenman, M.J. Director’s Report of the Wistar Institute for the Year 1910. p.10. Wistar Institute Library, Wistar Institute, Philadelphia, Pennsylvania. Hatai later became involved in Donaldson’s research to determine the effects of exercise on brain growth, which was explicitly tied to the domestication program.

63 periodically than to incur the difficulty and expense of bringing gray rats into the laboratory, housing them, and inducing them to breed.

Feralizing rats was a unique approach to the problem of domestication, questioning whether or not the numerous effects of domestication were reversible. It also challenged the common assumption that domesticated animals were incapable of surviving under wild conditions, especially an animal made conspicuous by albinism.

Although it may have seemed the easier of the two projects on the captive-feral dichotomy, the feralization experiment still required careful planning and execution. Site selection was critical. The location would need to be free of Norway rats that might be confused for the progeny of the feral population, if they were in fact capable of rapid morphological change in the new surroundings. Preexisting Norways would also present the fiercest competition for the transplanted albinos. Donaldson thought that a small island might be the ideal location, where the field could be cleared of any existing rats, transplanted rats and their progeny would not find too many places to hide, and external interference, especially by humans could be minimized. Finally, the site would need to come with a landowner who would submit to having dozens of rats released on the property with the stated goal that they would propagate themselves.

In his diary, Donaldson recorded a dinner meeting in April, 1910 with Charles

Davenport, who raised the possibility of using Goose Island, a small skerry in Long

Island Sound, which the Carnegie Institution had purchased for the use of Davenport’s

Station for Experimental Evolution one year earlier.134 Less than a month later

Donaldson visited the island with Davenport and came away with a “plan to catch all the

134 Donaldson, H.H. Diary- April 23, 1910. Henry Herbert Donaldson Diaries and Papers. American Philosophical Society, Philadelphia, Pennsylvania.

64 rats there and restock the Island with Albinos to see whether they can regain their nervous weight.”135 This plan was to begin with intensive trapping; “After that,” wrote

Donaldson in late May, “we must have recourse to a ferret or chlorine gas or something of that kind.”136 On June 6 he recorded, “Hatai [and J. MacPherson] Stotsenberg started for Goose Island to exterminate the rats- a fine day.”137 Hatai reported back to

Donaldson that there were no wild Norway rats to be found on the island. Davenport, when he learned of Hatai’s report, was incredulous, but agreed that it was probably safe to release the albino rats. He added optimistically, “I shall be much surprised if we do not get some grays with short white patches on the belly by the end of the year.”138

Donaldson, though, was still concerned about human intervention and possibly about raising ire in nearby Norwalk, Connecticut by seeding the island with vermin. He requested that the rats be taken by boat from Davenport’s station at Cold Spring Harbor on the other side of the Sound, rather than routing them through Norwalk, “and thus doing it without exciting remark in the town.”139 Hatai performed the clandestine colonization and waited with the immigrant rats for a day before returning to

Philadelphia.

The first attempt at feralizing albino rats evidently failed, though Donaldson did not seem downcast about the outcome. He wrote to Davenport a month after the release,

“I enclose this first notice of the ‘Goose Island’ adventure. Harmless in a way- but

135 Donaldson, H.H. Diary- May 13, 1910. Henry Herbert Donaldson Diaries and Papers. APS. 136 Donaldson to Davenport May 20, 1910. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.1. APS. 137 Donaldson, H.H. Diary- June 6, 1910. Henry Herbert Donaldson Diaries and Papers. APS. 138 Davenport to Donaldson June 10, 1910. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.1. APS. 139 Donaldson to Davenport June 13, 1910. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.1. APS.

65 saddening if it were not so funny.”140 Davenport, though, relayed that a workman building a dock on the island “saw a large white rat and one young, that is small, gray rat.”141 The sighting, similar to Davenport’s earlier, sanguine prediction of gray progeny by year’s end, was never verified. Nevertheless, Donaldson and Hatai made plans to repeat the experiment the following summer.

On Hatai’s second attempt to establish a breeding colony of albino rats on Goose

Island, he brought 34 ear-marked albinos with him. A preliminary survey of the island yielded a surprising result: no albinos were found, but Hatai spotted several gray

Norways. Even with the ripe hopes surrounding the rapid transformation of the domestic albino to the condition of the wild gray, Hatai had to conclude that these rats were not derived from the colony he planted. Perhaps Davenport was right and preexistent grays had escaped detection in 1910. Norway rats are also notoriously strong swimmers and may have reached Goose Island from another neighboring island, Hatai ventured. He went ahead with his planned release and stayed on the island to monitor the albinos for several days. The outcome this time seemed less like a comedy of errors than a nightmare scenario. At dusk after the release Hatai could see albinos scurrying at a distance; at night he heard repeatedly “the screaming of rats.” The morning revealed several dead and mangled rats, all albinos. More albinos, dead and alive, appeared until the time of Hatai’s departure.142 Subsequent monitoring convinced Donaldson not that

140 Donaldson to Davenport July 20, 1910. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.1. APS. Donaldson’s report was not preserved with the letter in Davenport’s collection. 141 Davenport to Donaldson September 5, 1910. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.1. APS. 142 Hatai S. Untitled Report on Goose Island Experiment. August 1, 1911. Charles B. Davenport Papers B.D27 Series I, Folder “Hatai, Shinkishi” APS. The same folder also includes a letter from Hatai

66 feralization was impossible, but that Goose Island was not a suitable experimental site.

He wrote Davenport:

Taking all things into consideration, it has seemed to us on talking it over that we were against too heavy odds on Goose Island to hope for ultimate success; the odds being the scarcity of food and openess [sic.] of the Island to invasion by Norways. I should be pleased if you found something to contradict this impression, but for now we shall have to wait… The experiment gains in importance [however] by the fact that we have conclusive evidence at present that domestication combined with inbreeding brings down the weight of the nervous system in the Norway rat to that of an albino in about four generations.143 Scarce records indicate that the feralization project migrated north, briefly, to the

Ipswich Sands in northeastern Massachusetts before heading south to another Carnegie- operated site in the Dry Tortugas, Florida. Donaldson wrote of the latter attempt, “For the opportunity to set out the colony we are indebted to the Carnegie Institution- represented in this case by Dr. A. G. Mayer, director of the station on the Dry Tortugas, who most kindly placed the animals for us.”144 The willingness of the Carnegie staff to help allowed the experiment to be executed from a distance. Hatai and Donaldson learned that the rats placed on the small atoll of East Key in June 1914 were still alive more than a year later. With the promise of such success, they made plans to visit the

Tortugas in the summer of 1916: “The purpose of this visit was to examine a group of albino rats which had been placed for us in June 1914 on a remote Key in this group. The hope was that these rats would adapt themselves to a wild life and in succeeding

to Davenport (August 19 1911) apologizing for having sent a report unapproved by Milton Greenman. Hatai repeatedly asked Davenport to strike any mention of Greenman or the Wistar Institute from the report, suggesting the control that Greenman exercised over the research output of the Institute. 143 Donaldson to Davenport June 27, 1912. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.2. APS. 144 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year 1915. p.5. WIL.

67 generations undergo physical changes as a result of these free conditions. In all, seven rats were removed out if eight originally placed on the key, but we failed to get evidence that the rats bred. Unless they breed, the experiment fails.”145

Donaldson failed to mention that it was in fact impossible to determine if the albinos had bred and successfully reared young. He noted elsewhere that the handful of rats that Mayer released were not marked and were thus indistinguishable from any fully- grown progeny they might have produced.146 He found no modifications in the albinos retrieved from East Key in 1916.147 Presuming that the albinos on East Key were either very few in number or had been extirpated, Donaldson placed 30 pairs on the island and another 11 males and 13 females on Garden Key in the same archipelago. Hatai traveled to the Tortugas the following summer and found no rats on Garden Key, but managed to recover seven albino rats from East Key, including two young rats that were not earmarked.

A summary in the Carnegie Institution Yearbook for 1917 suggested that this trial had achieved nearly everything that Hatai and Donaldson had hoped it would: “[T]he animals bred on the key and were feeding upon Ocypoda crabs and grass seed. The weight of the brain and the spinal cord has become relatively heavier in these rats now living under the trying conditions imposed by the small semi-desert islet East Key.”148

Hatai’s detailed report was somewhat more qualified, noting that the relative increase in

145 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1916. p.6. WIL. 146 Donaldson, H.H. Experiment on the Feralization of the Albino Rat. Yearbook of the Carnegie Institution of Washington. 1916. Vol.15:200-201. 147 Donaldson, H.H. Diary- June 26, 1916. Henry Herbert Donaldson Diaries and Papers. APS. 148 Mayer, A.G. Department of Marine Biology. Yearbook of the Carnegie Institution of Washington. 1917. Vol.16:165.

68 nervous system weight was approximately the same in the original lot and in their progeny. He noted that he would require fully grown progeny, preferably in multiple generations, to determine whether the wild conditions conferred lasting and not merely temporary constitutional changes.149 Donaldson struck an even less triumphant tone in his report to Greenman in 1918: “I regret to report that this is our fifth failure to establish a colony of wild albinos… Rats released the previous summer [1916] had in some instances survived and in one instance at least reared young, but the food, water, and temperature conditions are too severe for a colony to thrive.”150 Donaldson added,

“Nevertheless I do not think that the effort to [feralize the albino rat] should be abandoned.”151 Despite his optimism, the Wistar Institute made no further attempts.

After more than eight years and at least five trials, the feralization project met a disappointing end. Donaldson was encouraged by the limited results of the experiment, though he acknowledged that the project was ultimately unsustainable in any of the locations attempted. Certainly it had proven more complicated in practice than in concept. Donaldson and Hatai’s naïve expectations for the feralization experiment were likely tied to the vagaries behind the question of how a “wild life” would revert the physiognomy of an albino rat back into that of its progenitor, the gray Norway. Although this experiment was devised to address the role of the “struggle for existence” in domestication, Donaldson and Hatai most often treated the “life under wild conditions” itself as the agent of morphological change. The notion of reversing domestication did

149 Hatai, S. On the Condition of the Albino Rats on East Key and Garden Key in the Dry Tortugas. Yearbook of the Carnegie Institution of Washington. 1917. No.16:179. 150 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s report for the Year Ending December 31, 1918. p.8-9. WIL. 151 Ibid. p.8.

69 not signal a departure from teleology either. The changes these wild conditions were meant to affect were expected to move toward known ends. That is, feralization was expected to follow a defined path from the altered condition of the albino to the ostensibly more natural form of the wild gray rat, one that paralleled its initial domestication. That this did not occur in five attempts did not dissuade Donaldson that it should.

In the wave of new experiments associated with Donaldson’s interest in the effects of domestication, Hatai also initiated a study complementary to the feralization project, this time taking wild gray rats into captivity and observing their morphological change under newly imposed conditions of life. This experiment seemed to address the problem of domestication in a more straightforward manner, though initial expectations seemed to be that this effort would be costlier and would perhaps take longer than the feralization study. Hatai’s first mention of the experiment noted that in 1910 “[s]everal litters [of gray Norway rats] were born in captivity and this encouraged us to follow this work which must necessarily extend over a number of years, as we desire to determine whether domestication affects the nervous system unfavourably.”152 He faced trouble early on in maintaining fecundity under captivity, however. “After several failures to breed the Norway rats in cages, we have succeeded in raising them in captivity by means of an improvement in the cages and diet, as well as in general treatment,” Hatai reported in 1912.153 The challenge of caging wild grays required some associated changes in diet and treatment, but Hatai also mated the subsequent generations, brother to sister, “as [he]

152 Hatai, S. in Director’s Report of the Wistar Institute for the Year 1910. p.12. WIL. 153 Hatai, S. On the Appearance of Albino Mutants in Litters of the Common Norway Rat. Science. 1912. Vol.35(909):875.

70 wished to combine the effects of close inbreeding and captivity,” perhaps to more closely emulate the effects of the metaphenomenon, domestication.154 This approach, it seemed, could only further complicate the interpretation of his experimental results. The changes he was able to affect by this method, however, exceeded his expectations. Donaldson recorded Hatai’s results with enthusiasm in his diary: “Hatai has succeeded in getting albinos in two litters of pure Norways. A great event for us.”155 Hatai elaborated,

“Although I have been anticipating that such albino mutants might occur sometime, nevertheless, it is a great surprise to obtain them within so few generations. So far as I am aware, this is the first instance in which the albino mutants have been obtained from the common Norway rats under laboratory conditions.”156 As mentioned in his letter to

Davenport, Donaldson attached great significance to another finding from the captive breeding experiment which he described in his diary: “Made first exam. of inbred domesticated Norway rats (Hatai). The nervous system less than in albinos. This is a most important result.”157 Again, Hatai and Donaldson presumed to know in which direction the morphological changes associated with captivity would take, extending even to color. The speed with which these modifications occurred, however, was a genuine surprise.

What is perhaps most surprising about Hatai’s captive breeding experiment was how abruptly it was aborted. Little mention was made of it after 1912. An explanation is difficult to come by, except for some suggestive entries in Donaldson’s diaries, which

154 Ibid. 155 Donaldson, H.H. Diary- May 3, 1912. Henry Herbert Donaldson Diaries and Papers. APS. 156 Hatai, S. On the Appearance of Albino Mutants. p.876. 157 Donaldson, H.H. Diary- June 21, 1912. Henry Herbert Donaldson Diaries and Papers. APS.

71 were spared any superfluous detail. Just two months after Hatai discovered albinos among his captive grays, Donaldson recorded: “Talk with Castle. He is skeptical about our sports. Must take this info.”158 He followed this entry three months later with:

“Serious talk with Hatai about rat work- he is much discouraged by difficulties in the conduct of the rat room. Must adjust this.”159 Perhaps another variable had marred

Hatai’s already complicated results. A general problem with insufficient protein in the diet of all the Wistar rats had altered the condition of most stocks at the Institute early in

1911. This problem was addressed prior to Hatai’s captive breeding experiment, but similar problems with care may have contributed to Hatai dropping the project. Still, following such encouraging results, the project’s disappearance seems remarkable.

Throughout the remaining decades of work on domestication at the Wistar Institute, this early experiment by Hatai was conspicuously absent from the institutional memory.

Donaldson strayed from his custom of reiterating the human implications of the work on the Norway rat with these early studies. He did group the feralization study (but not Hatai’s captive breeding experiment) in with other rat work to suggest the rat’s general applicability to human problems: “The rat is used for these studies and the further we progress with the study of it the clearer becomes the fact that in the nervous system at least the results obtained on the rat can be carried over to man.”160 Aside from this general allusion Donaldson gave neither of these early experiments specific human

158 Donaldson, H.H. Diary- July 5, 1912. Henry Herbert Donaldson Diaries and Papers. APS. 159 Donaldson, H.H. Diary- October 3, 1912. Henry Herbert Donaldson Diaries and Papers. APS. The 1915 Director’s Report alluded briefly to Hatai’s experiment. This description seemed to affirm the validity of the experimental results, though it offered little in the way of an explanation for why the project was cancelled. See The Wistar Institute of Anatomy and Biology Director’s Report for the Year 1915. WIL. 160 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1918. WIL.

72 implications, likely because problems with their execution prevented him and Hatai from publishing their results more fully. It thus remains an open question what might have been “carried over to man” at least from the attempt at feralizing domesticated albino rats and if one or both of these experiments were originally conceived in terms of the anthropological analogy between domestication and civilization with which Donaldson was familiar.

Experiments on Exercise and the Tone of Modern Life

Between 1910 and 1915, Hatai and Donaldson were also engaged in a series of experiments to determine the impact of exercise on the weight of the brain and other organs of the albino rat. Exercise, according to Donaldson, was another potential factor to explain the presumptively domesticative changes that separated captive albino and wild gray rats. By Donaldson’s explanation: “The diminution in the amount of exercise is… one of the several factors possibly contributing to the difference. The present experiments were undertaken therefore to see whether if they were given more exercise the albinos would improve in respect to the weight of their central nervous system.”161 In the first experiment, completed in two trials during 1910, Donaldson placed a small number of rats in revolving drum cages, the walls of which spun on a fixed horizontal axle from which food, water, and a nesting box were suspended. An approximately equivalent number of rats were placed in standard stationary cages with limited opportunity for exercise, constituting the control group. Following six months in cages

161 Donaldson, H.H. On the Influence of Exercise. p.129-130.

73 that either permitted or restricted free exercise, the brains and spinal cords of the experimental and control rats were weighed. Exercised rats’ brain weights increased, on average, 2.4 percent more than those of the controls in the first trial and 2.7 percent more in the second, this, despite a slight reduction in body length among 74 percent of exercised rats. Donaldson’s plans for future research suggested that he regarded these changes as more than transitory physiological effects. “If exercise can influence the weight of the nervous system,” he wrote, “it seems probable that its effects can best be studied on animals grown for several generations under the conditions here described.

Later we hope to study animals thus grown, as it has already been found that rats can breed in these revolving cages and bring up their young in them without difficulty.”162

He imagined that exercise, like many of the changed conditions of life under domestication, was capable of becoming internalized and permanent after successive generations.

In 1915 Hatai published an update and expansion of Donaldson’s initial exercise study. The expansion did not, however, conform to Donaldson’s plans of four years earlier. Hatai’s experiment used at least three generations of albino rats, but these were not successive and he gave no indication that the rats had been born and raised under the conditions of the exercise study. Rather, it seems, these were separate series. Hatai did expand Donaldson’s study, though, by comparing anatomical features other than the brain and spinal cord. He was encouraged by his discovery of several differences in the weights of ductless glands between gray and albino Norway rats, which he interpreted as

162 Ibid. p.136.

“a response to the complex conditions represented by domestication.”163 Hatai’s study on exercise, then, signaled that the subject of domestication had assumed an importance at the Wistar Institute independent of Donaldson’s original and overarching problem, the growth of the brain.

In his paper, Hatai reiterated the relation of the exercise study to the domestication problem: “Domestication… involves numerous interrelated factors which can only be slowly isolated by systematic study, and this experiment was devised to test the value of exercise, which seems to be one of the important factors forming the complex of domestication.”164 He found that long continued exercise (i.e. for 90 or 180 days) had little effect on the external morphology of the albino rat, not even the slight decrease in body length among exercised albinos that Donaldson noted. Internal viscera, however, were considerably more plastic. Like Donaldson, he noticed a slight increase in brain weight (4.1 percent) and no change in spinal cord weight with exercise. The hearts, livers, and kidneys of exercised rats were approximately 20 percent larger than those of the controls, while the spleen showed an equivalent decrease. The change in the adrenal glands and pituitaries was complicated by differences in sex, but Hatai maintained that with exercise they approached the condition of the wild gray Norway.165

The albino rat alone was studied in both the 1911 and the 1915 experiments.

No comparisons were made with exercised wild caught gray rats, for it was felt that “a considerable curtailment of the normal activity in the albino rats under domestication

163 Hatai, S. On the Weight of Some of the Ductless Glands of the Norway and of the Albino Rat According to Sex and Variety. Anatomical Record. 1914. Vol.8(12):522. 164 Hatai, S. On the Influence of Exercise on the Growth of Organs in the Albino Rat. Anatomical Record. 1915. Vol.9(8):663. 165 Ibid. p.650-657.

75 seems highly probable.”166 Donaldson and Hatai considered “normal” activity to be that of the wild and putatively more natural gray Norway. Thus the exercise treatment, like feralization, was not meant to modify a natural state, but to return the already modified albino to a condition like that of its wild progenitor. Wild gray rats did not need to be tested because with them the results of generations of heightened activity were inborn.

Donaldson had long expressed interest in the long-term effects of peculiar conditions, especially as they were found to be cumulative or inherited. This interest seems to have underlain not only his view of the effects of exercise and domestication more broadly, but also some of his attempts to suggest the importance of anatomical research to sociological problems. He once stressed the value of studying “the modifications of structure that come along with the lifelong pursuit of certain handicrafts… as, for example, the anatomy of a shoemaker,” which he hoped to tie to

“problems in sociology.”167 In describing another of Hatai’s experiments in which chronically undernourished albino rats showed a reduction in brain weight, Donaldson again revealed how the work appealed to some of the moment’s most pressing social concerns: “[W]e may expect to find a corresponding modification in man, when the human body during the period of growth is subjected to unfavorable conditions of a similar nature. As a matter of fact such conditions do exist in the crowded quarters of our largest cities, and it seems highly probable that we have there in progress examples of partial starvation quite comparable with the experiments conducted in the laboratory.”168

A decade later Hatai suggested the applicability of the results of his study on the effects

166 Ibid. p.650. 167 Donaldson, H.H. Problems in Human Anatomy. p.19. 168 Ibid. p.24.

76 of long-continued exercise: “[F]rom the anatomical side the question of exercise is taken rather lightly in the case of man, nevertheless when we consider its striking effect on some of the organs of the rat, a further careful investigation, not only in the rat but in man also, seems certainly worth while, both from the general biological standpoint and for its bearing on hygiene.”169 In the context of 1915, “hygiene” was a laden term. Affixed to the words “mental’ or “social,” it signified an immense social reform movement in the

Progressive Era. Hygiene took on both physical and moral significance as cleanliness, purity, and health became symbolic ideals under which social and political movements such as temperance and eugenics found common appeal. It is doubtful that Hatai intended to invoke the more moralistic resonances of the social hygiene movement, but the appeal to “hygiene” in this decade recalled social rather than individual concerns, such as the pressures of urban life and modernity more generally.

As Hatai was responsible for evaluating the effects of “the struggle for existence” and exercise, so he also participated in experiments to address the remaining factors of domestication that had come to Donaldson’s mind, namely, interbreeding and albinism.

Donaldson first mentioned “interbreeding” as a potential factor of domestication in 1905 and reiterated this point in 1909, but the term fell out of use at the Institute after this.

“Interbreeding” is now most often associated with the practice of crossing distinct varieties of animals or plants, whereas before the turn of the 20th century it was nearly as often used to mean the breeding of closely related individuals.170 By 1911 Hatai

169 Hatai, S. On the Influence of Exercise. p.663. 170 In her first summary paper on the effects of inbreeding, Helen Dean King cited Darwin’s repeated use of the term “interbreeding” and without further explanation translated it into her own preferred term “inbreeding.” See King, H.D. Studies on Inbreeding I. The Effects in Inbreeding on the Growth and Variability in the Body Weight of the Albino Rat. Journal of Experimental Zoology. 1918.

77 employed the terms hybridization and inbreeding, respectively, to distinguish these meanings and had found ways to evaluate the effects of these practices. He seems to have initiated the hybridization experiments himself before 1910 by crossing individuals of the tame albino stock with wild caught gray Norways. For the inbreeding study, however, he partnered with Helen Dean King, who quickly assumed greater responsibilities at the Institute, beyond the technical duties for which she was hired. In the course of the next decade leadership on the hybridization project and even the revived captive breeding experiment traded hands, from Hatai to King. This transference, which was by all indications orderly, preceded Hatai’s return in 1924 to his native Japan to serve as Professor of Biology at Tohoku University. He remained tied to the domestication program, especially the exercise experiments and various neurological analyses, but the bulk of the responsibility for the program fell to King by the latter half of the decade, with Donaldson at least nominally in charge of the program’s overall organization and direction.

Inbreeding, Hybridization, and Notions of Purity in Rats and Man

Hatai and King arrived at the Wistar Institute within two years of each other.

When Donaldson left Chicago for Philadelphia, he brought a small number of albino rats and Hatai with him as an Associate in Neurology. At Chicago, Hatai had proven himself capable of very fine technical work and of staggering workloads. In the 1905-1906

Vol.26(1):1. The popular press, when commenting on King’s work, found the terms similarly interchangeable. See “Wistar Institute Proves by Rats Consanguinity No Wedlock Bar” Evening Public Ledger (Philadelphia) September 29, 1915.

78 academic year, for instance, Hatai assisted with or was lead instructor for eleven neurology courses, in addition to his laboratory research.171 Donaldson had hosted several Japanese researchers at his laboratory in Chicago, and he carried on this tradition at the Wistar Institute, no doubt with Hatai’s help. A number of women had also found opportunities for research in Donaldson’s Chicago laboratory, so perhaps it is not surprising that Helen Dean King was hired as an Assistant in Anatomy at the Wistar

Institute in 1908. King had distinguished herself as a graduate student under Thomas

Hunt Morgan at Bryn Mawr. Her studies there reflected Morgan’s influence, especially his interest in embryology, including her Ph.D. dissertation (1899), a detailed explanation of “The Fertilization and Maturation of the Egg of Bufo lentigenosus.” She spent a combined seven years following graduation teaching at the Shipley and Baldwin Schools, preparatory academies in Bryn Mawr, before assisting Edwin G. Conklin as a Fellow in

Biology at the University of Pennsylvania from 1906 to 1908.172 It may have been

Conklin’s influence as a member of the Wistar Advisory Board as much as Donaldson’s willingness to hire women for his laboratory that won King entry into the Wistar

Institute. Her position in the first year came without pay, a sacrifice many women in

King’s generation made for the opportunity to do scientific work. Early in 1909 she was made an Assistant in Anatomy and was given technical duties including “the preparation of sections and the elaboration of new technical methods. She was also able to

“[conduct] her own researches in sex determination” in amphibians, a longstanding

171 Annual Register of the University of Chicago 1904-1905. p.313. 172 For a more detailed description of King’s early career, see Ogilvie, M.B. Inbreeding, Eugenics, and Helen Dean King (1869-1955). Journal of the History of Biology. 2006. Vol.40:469-471. See also, Ogilvie, M.B. and Harvey, J. (eds.) The Biographical Dictionary of Women in Science: Pioneering Lives from Ancient Times to the Mid-20th Century. New York, Routledge,, 2000. p.695-696.

79 interest of hers and a mark of independence, as historian of science Marilyn Ogilvie has characterized it.173 By 1912, the same year in which Hatai was named Instructor in

Neurology, King was promoted to Assistant Professor of Embryology, a selection that won acclaim in the Director’s Report: “Your decision to promote to an Assistant

Professorship, Dr. Helen Dean King will be met with earnest approval by men working in the biological field of science.”174 Newspapers in Philadelphia were keen to point out the rarity of her position as a female professor at a scientific institute.175

That King started two lines of inbred albino rats, ultimately to determine if gender differentials could be established, suggests that she was free to pursue her own interests in sex ratios from the beginning of the inbreeding project. Her quickly growing esteem and autonomy at the Wistar Institute notwithstanding, Donaldson was likely the one to charge her with breeding rats brother to sister over numerous generations in connection with his interest in “interbreeding” as a factor in domestication and in the differential growth of the brain. King only published on the inbreeding problem nearly a decade after the experiment was begun, owing to the need for multiple generations and the solution to problems in care before she could obtain clear results. Her introduction to this synoptic series of papers nine years hence omits some of the context within which the project was conceived. The effect of inbreeding on the growth of the brain, for instance, is presented as a secondary consideration behind King’s own interests in its effects on fertility, vitality, and the sex ratio. Hatai, however, recorded in 1910 that “[the inbred] series is

173 Ogilvie, M.B. Inbreeding, Eugenics, and Helen Dean King. p.471-472. 174 The Wistar Institute Director’s Report for the Year 1912. p.4. WIL. 175 M’Liss “Woman Professor Says Theory may Predetermine Sex: Dr. King Declares Her Experiments with Rats have Bearing on Human Problem- not Afraid of ‘Heresy.’” Evening Public Ledger (Philadelphia). September 30, 1915.

80 being followed to determine primarily whether inbreeding causes a deterioration of the central nervous system- at the same time we are hoping to throw some light on the recently much discussed theory of the pure line.” The inbreeding experiment did not merely involve “taking advantage of the opportunity which the animal colony of the

Wistar Institute of Anatomy and Biology afforded,” as King later claimed.176 Rather, it belonged, first, to a coordinated program of research on the problem of domestication and its associated factors with reference to the growth of the brain.

Inbreeding and hybridization were among the most critical problems in theoretical biology during the first two decades of the 20th century. These practices were defined historically by cultural factors, specifically taboos restricting the union of close relatives in societies the world over. They were given scientific voice by Charles Darwin whose work self-fertilizing plants and association with breeders convinced him that consanguinity had deleterious effects on the offspring of a too-close union. Atop private fears that his own family suffered from generations of close relations, Darwin helped to create the sense that inbreeding and crossing were as sharply opposed in effect as they were in practice: “[I]t is a great law of nature, that all organic beings profit from an occasional cross with individuals not closely related to them in blood; and that, on the other hand, long-continued close interbreeding is injurious.”177

Research by H. Crampe, J. Ritzema-Bos, August Weismann, and Georg Von

Guaita prior to 1900 attested to the negative effects of inbreeding on animals. George

Shull reinterpreted Darwin’s dichotomy of inbreeding and outcrossing with his

176 King, H.D. Studies on Inbreeding I. p.2. 177 Darwin, C.R. The Variation of Animals and Plants Under Domestication. Vol.II. London: John Murray, 1868. p.116.

81 experiments with maize in 1908. He affirmed that the practices generally had opposing effects, but suggested that these results were based on a common principle: “the relative vigor of biotypes and their hybrids.”178 Still Raymond Pearl lamented that, despite increased interest from the scientific community, “no phase of biology has been enveloped in a fog of superstition, old wives tales, and other sorts of misapprehension as has inbreeding.”179 He suggested in 1913 continuing the work started by “[Georg]

Lehndorff, [W. F.] von Oettingen, Bruce Low, [F. Hoesch], [A. de] Chapeaurouge, [R.]

Bunsow, [R.] Stranz, and others” who evaluated the effects of inbreeding by the use of breeder’s records, but disparaged that these studies had not “led to results characterized by the precision or the definiteness or the quality of getting at fundamentals demanded in the present state of the science of genetics.”180 Before inbreeding could gain any biological legitimacy, Pearl argued, it had to be rationalized, unencumbered by speculation, superstition, and tradition.

In the Progressive Era, when interest in inbreeding and hybridity was at its height in the U.S., rationalization of the phenomena was not enough, but “precise and definite” answers from natural science would need to guide their new social and cultural standing as well. These aims met in a 1919 book by Edward Murray East and his former graduate student Donald Jones, Inbreeding and Outbreeding: Their Genetic and Sociological

Significance. “Interest in the effects of inbreeding and of outbreeding is not confined to the professional biologist,” the authors opened their argument, “Historically, these are

178 Shull, G.H. The Composition of a Field of Maize. Report of the American Breeders’ Association. 1908. Vol.4:299. 179 Pearl, R. Review of East, E.M. and Jones, D.F. Inbreeding and Outbreeding: Their Genetic and Sociological Significance. in Science 1920. Vol.51(1321):416. 180 Pearl, R. A Contribution Towards the Analysis of the Problem of Inbreeding. American Naturalist. 1913. Vol.47(562):578.

82 old, old problems, practical problems bound up with man’s gravest affairs, his marriage customs and his means of subsistence.”181 The effects of inbreeding and outcrossing were not merely of historical interest; they remained two of the great, unsolved problems in biology. As the American eugenics movement was steeped in anxiety over proper unions for race improvement, these problems found an audience in sociology as well.

The sociological implications that arose from the scientific investigation of inbreeding and outbreeding were as extensive as the superstitions and customs they were meant to replace. The new understanding of these practices, “in accordance with biological facts,” could be used “in establishing a concrete scientific basis for marriage, divorce, and immigration laws… predicting the changes to be expected in the body politic due to differential fecundity, birth control, and other agencies by which the character of the population is shifted; [it even has] some relevancy to many problems which one might suppose were wholly of an economic nature, such as minimum wages and mothers’ pensions.”182 King’s work on inbred rats featured prominently in East and Jones’ account, along with their own work on maize. Her experiments on inbreeding and hybridization demonstrated not only the careful execution that East, Jones, Pearl, and others praised, but also the Progressive impulse to apply experimental results to social problems.

The first six generations of King’s inbred albino rats seemed to affirm the conventional wisdom about the ill effects of close inbreeding. After starting with two pairs of stock Wistar albino rats, she began breeding as closely as possible, that is brother

181 East, E.M. and Jones, D.F. Inbreeding and Outbreeding: Their Genetic and Sociological Significance. Philadelphia: J.B. Lippincott Co., 1919. p.13. 182 Ibid. p.15-16.

83 to sister, with strict selection for size and apparent vigor. Each mating pair was also established as the start of two distinct lines, one eventually selected for a preponderance of male offspring, and the other for an excess of females. After two years of breeding,

“[m]any females in both series were sterile, and those that did breed usually produced one or two litters that were generally of small size. A considerable number of rats were dwarfed, or stunted in their growth, and many of them developed malformations, particularly deformed teeth. The animals showed, also, a decline in vitality in succeeding generations and usually died at a relatively early age.”183 Later in the year Henry

Donaldson shared with Charles Davenport that “in a series of experiments on very close inbreeding now in progress by Dr. King, we find an astonishing loss in the relative weight of the nervous system.”184 The decline that King described, however, was a general one, affecting not only the inbred, but also the normal stock rats at the Wistar. In the spring of 1911 all of the rats at the Institute were given a more diverse and protein- rich diet. The inbred rats, along with the stock albino rats, demonstrated a marked recovery in growth, vigor, and fertility.

King’s first publication on the subject, one in a series of four papers over two years, used body weight as a general gauge for the effects of inbreeding on growth. In the paper she declared, “The bad effects of inbreeding per se, as far as they might manifest themselves as a decrease in the body size of the individuals, have apparently

183 King, H.D. Studies on Inbreeding I. p.4. 184 Donaldson, though, countered King’s description by noting that there was no “apparent loss in bodily vigor, size, or capacity for reproduction.” This discrepancy might be explained by the fact that Donaldson’s comments were made late in the year, at least five months after King’s inbred rats were given a more nutritious diet and many of these qualities were improved in them. The brain weight in inbred rats seems to have made a slower recovery than other traits. Donaldson to Davenport October 26, 1911. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.2. APS.

84 been entirely prevented through the use of a strain of animals that seemingly had no inherent defects and by a careful selection of breeding stock.”185 Body weight showed a general decrease in variability versus the stock control strain as the inbred strain approached homozygosity. Although changes in variability after the fifteenth generation of inbreeding forced King to admit that her rats were “far from ‘pure’ in the sense in which the term is used by Johannsen (’09) and his followers.”186

Two of King’s later focuses, fertility and constitutional vigor, recalled some of the longest standing biological objections to the practice of inbreeding. Against the claims of

Darwin, Crampe, Ritzema-Bos, and von Guaita, King asserted that inbreeding entailed no necessary tendency to infertility. She maintained that fertility was probably the greatest measure of “constitutional vigor,” but added the evident ability for growth, resistance to disease, and longevity to produce a clearer picture of overall vigor. In all of these capacities, King assured, her inbred rats were equivalent or superior to the stock controls.

One other measure, “mental alertness,” seemed the most closely connected to

Donaldson’s original interest in inbreeding as a factor in the diminution of brain weight in domesticated albino rats. Indeed, Donaldson had pursued outside collaborators to determine if the decrease in brain weight he had noted in the inbred rats in 1911 was correlated with a change in mental function. In that year, he approached his former graduate student, John B. Watson in the Psychology Department at Johns Hopkins, who turned the project over to his own graduate student, Gardner C. Basset. Before Basset’s study was even complete, Donaldson had also contacted Robert Yerkes at the Harvard

185 King, H.D. Studies on Inbreeding I. p.48. 186 Ibid. p.49.

Psychological Laboratory to run a similar experiment, which Yerkes put in the charge of his wife and, subsequently, the would-be ethnologist, Nenozo Utsurikawa.187 Basset and

Ada Yerkes both concluded that inbred rats did not learn as well as their outbred stock counterparts. Yerkes and Utsurikawa agreed that inbred rats were less active than stock albinos, and simultaneously more savage in response to handling Utsurikawa added.

Yerkes attributed the lesser activity of inbred rats to “a greater timidity,” which King suggested might have been fixed by inbreeding because timidity and savagery were not selected against in her scheme of breeding. Hatai investigated the brains of several of these rats and some from King’s later generations. He confirmed that the central nervous system of the rat had recovered from its nadir in 1911, much like many of the other ill effects of the insufficient diet that year, albeit more slowly.

King maintained that, taken together, “these experiments have fully demonstrated… that even in mammals the closest form of inbreeding possible, i.e., the mating of brother and sister from the same litter, is not necessarily injurious either to the fertility or to the constitutional vigor of a race even when continued for many generations. Success or failure in inbreeding experiments depends chiefly, it would seem, on the character of the stock being inbred, on the manner in which the experimental animals are selected, and on the environmental conditions in which the animals are reared.”188 These last points, King claimed, were what distinguished between the conclusion of earlier investigators that inbreeding was invariably detrimental in its

187 Yerkes, A. Comparison of the Behavior of Stock and Inbred Albino Rats. Journal of Animal Behavior. 1916. Vol.6(4):267 and Utsurikawa, N. Temperamental Differences between Outbred and Inbred Strains of the Albino Rat. Journal of Animal Behavior. 1917. Vol.7(2):111. 188 King, H.D. Studies on Inbreeding II. The Effects of Inbreeding on the Fertility and Constitutional Vigor of the Albino Rat. Journal of Experimental Zoology. 1918. Vol.26(2):376.

86 effects and her own that such a decline was in no way necessary. She hypothesized that the pessimistic conclusions of Ritzema-Bos, von Guaita, Weismann, Crampe, and others were due, variously, to poor environmental conditions, lack of selection for vigor, poor quality initial stock, overcrowding, and beginning the experiments with hybrid rather than nearly pure strains of rats. She found it ‘rather remarkable, that, of the many writers who have cited the results of the above series of experiments as proof that close inbreeding lessens fertility, not one, to my knowledge, has emphasized the fact that all of these experiments were made with hybrids and not with a pure strain.”189 She then reminded her readers that while hybridization was known to increase fertility, it just as often induced sterility. Her own experiments on hybridity seemed to emphasize the capriciousness of this strategy versus its counterpart, inbreeding.

Hatai had begun the hybridization experiments at the Wistar prior to 1909 as part of the general effort to understand the nature of the difference in brain weight between wild gray Norway rats and the domesticated Wistar albinos. The experiment addressed not only the effect of crossing distinct varieties, but also some basic questions about the inheritance of brain size and weight and the possible correlation of albinism with a diminished central nervous system. Hatai bred stock Wistar albino rats to gray Norways trapped around Philadelphia and then bred together their hybrid offspring for several generations. From analyses of the brain weights of the parental, first, and second filial generations he concluded that “the brain weight character appears to blend in inheritance,” i.e. the brain weight of the first and second hybrid generations fell evenly between those of the gray Norways and the domestic albinos. Furthermore, by

189 Ibid. p.371.

87 comparing rough groups of pigmented versus non-pigmented rats, he determined that

“there is no definite correlation between the brain weight and the characters of the coat under these conditions.”190 Hatai did, however, mention that in the course of his studies several of his hybrid lines had suffered from sterility.

King became involved with this project by using Hatai’s rats to investigate the effects of hybridization on the sex ratio in rats. As these rats were not bred with such an investigation in mind, King faced considerable challenges in reaching a firm conclusion about the effects of hybridity. Nevertheless, she noted in later years, with a nod to the study’s putative sociological significance, “Hybridization in the rat, as racial crossing in man, seems to produce a marked alteration in the sex ratio.”191 She also mentioned that the later generations of extracted and subsequently inbred hybrid rats faced increasing rates of sterility, a trend that continued as she took over the breeding of the hybrid series from Hatai. She reiterated this point in her much later report on the effects on inbreeding: “none of the ‘extracted’ [i.e. hybrid] strains that I have studied have even been as fertile as the Inbred Albinos. The increase in sterility and the diminution in litter size with continued inbreeding has [sic.] been very marked in some of these strains, but this lessened productivity has been due, I think, to hybridization, and it has not been influenced by inbreeding save in as far as inbreeding has intensified the tendencies which acted unfavorably.”192

190 The pigmented group included not only solid gray rats, but also black, black piebald, and gray piebald rats that were extracted in the F2 generation. Hatai, S. On the Brain Weights of Rats Descended from the Cross between the Wild Norway (Mus norvegicus) and the Domesticated Albino (Mus norvegicus albinus). Journal of Comparative Neurology. 1915. Vol.25(6):563. 191 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1925. p.10. WIL. 192 King, H.D. Studies on Inbreeding II. P.371.

King wished above all to dispel the common belief as expressed by H. Kraemer that “continued inbreeding must always result in weakened constitution, through its own influence.”193 She did not necessarily support the use of inbreeding over hybridization among livestock and claimed that both practices would continue to find important uses in the future and more rational breeding of stock animals. She recognized that the effects of inbreeding and hybridization acted on a single principle, the accumulation of beneficial or deleterious qualities through heredity, as Shull had roughly described ten years earlier and as East and Jones would clarify a year later in distinctly Mendelian terms. Still, the rationalization of these phenomena, at least in the examples of King and East and Jones, lent certainty to the outcome of inbreeding, while leaving hybridization mired with the same reputation for unpredictability. Hybridity always entailed the gamble of random segregation. Meanwhile, inbreeding promised surer results. In the words of East and

Jones: “Homozygosity, when obtained with all the most favorable characters, is the most effective condition for the purpose of growth and reproduction.”194 It was around this point that much of the perceived sociological importance of inbreeding and hybridization turned.

In her published work King was generally conservative in the interpretation of her results, certainly more so than East and Jones, and even more so than her pointedly uncontroversial boss, Henry Donaldson. A question posed at the beginning of her inbreeding study did, however, suggest a distinctly eugenical preoccupation: “If no evil results appear, can inbreeding be used to improve a race by combining the best of the

193 Quoted from Ibid. p.376. 194 East, E.M. and Jones, D.F. Inbreeding and Outbreeding. p.187.

89 dominant characters with any desirable recessive ones that may appear?”195 Her statements to the press and explanations in internal Institute reports further indicate that the biological and eugenical implications of inbreeding and hybridization were as intimately tied in King’s mind as in East and Jones’.

The public, or at least the scientific world, was probably introduced to King’s work on these subjects at the Philadelphia meeting of the American Association for the

Advancement of Science in December 1914. Donaldson recorded in his diary: “Miss

King reported this A.M.- It was splendid. A great card for herself and for the

Institute.”196 The Philadelphia Evening Public Ledger, and especially the pseudonymous reporter M’Liss, took an interest in King’s work on inbred rats in the following year. An article that appeared September 29, 1915 suggested that King thought not only of overturning the conventional wisdom about the injurious effects of inbreeding, but also mulled over the rationale and validity of social customs relating to close unions: “Doctor

King herself did not hesitate to declare emphatically this morning that the laws forbidding intermarriage and the traditions that have been handed down from time immemorial that have made the union of close relatives repugnant, cannot base their existence on any scientific fact.”197 In the article King moved with facility between the conclusions she reached through experiments on rats and the implication for human consanguinity. Said King, “I firmly believe, though, of course, it has not been tried out that if a brother and a sister, carefully selected of a higher type, were to marry, the result

195 King, H.D. Studies on Inbreeding I. p.2. 196 Donaldson, H.H. Diary- December 31, 1914. Henry Herbert Donaldson Diaries and Papers. APS. 197 “Wistar Institute Proves by Rats Consanguinity No Wedlock Bar.” Evening Public Ledger (Philadelphia). September 29, 1915.

90 of the union would be a higher type of offspring than from the intermarriage of two other people. The opinion regarding the degeneracy of the offspring of such a union exists because the only races who intermarry are degenerate races and therefore you get an even greater degeneracy.”198 This line of thinking was not peculiar to King, even at the Wistar

Institute. Milton Greenman’s response, solicited by the interviewer, suggests that the social and cultural implications of the work were settled in more minds than one at the

Institute, and had perhaps been a point of internal discussion: “I do not think it would be too much to say that as the result of Doctor King’s experiments it would be perfectly safe for close blood relatives to marry, if they are carefully selected. Of course, in an experiment with rats there is no way of telling what the effects might be on mentality.

The improvement noted has been purely a physical one.” Even as Greenman spoke this mild caveat, King’s inbred rats were tested by Ada Yerkes to determine the effects of inbreeding on mentality.

King anticipated the “cry of ‘heresy’ that [arose] on all sides as a result of the publicity of [her] research” and “[deplored] the fact that accounts of her discoveries

[were] apt to be regarded by the lay mind as a little ‘yellow.’”199 Of course, the reports of the Evening Public Ledger only helped to fan these flames. Sensational journalism, after all, was as much a part of the Progressive Era as the scientific rationalization of social practices. In its third article on the subject in four days, the Ledger noted that its previous report “caused much comment and discussion in clerical circles in the city yesterday.”

The paper claimed that most of the clergymen willing to discuss King’s interpretations

198 Ibid. 199 M’Liss “Woman Professor Says…”

91 believed “that the marital code of the world was too high to even entertain thoughts of such [close] marriages, even though science should prove that they would tend to produce a superman.” Even in the Progressive Era, scientific rationalization was not invariably the victor over moral considerations steeped in culture and tradition. The eugenics movement, for instance, was never fully able to confront the moral dimension of the cause of race improvement either in its detractors or within its own ranks. Thus, upon a moral objection, one of the clerics in the Ledger piece, Clarence Edward Macartney of the Arch Street Presbyterian Church, reversed King’s rationalized move from animal to human back to the animal: “The moral fence could not be broken down to permit

[consanguinity]… If such a condition should ever exist, although I am not fearing it ever will, all the happiness that comes through life and the romance that adds to the joys of the home will be killed… If there was an indiscriminate marriage of close relatives the people would live together like a lot of animals.”200

King was not chastened by the public reaction to her work. When she took full leadership of the hybridization project from Hatai upon his departure from the Wistar

Institute in 1924, her description of the project suggested that her motivations for the experiment were as much sociological as biological: “Primarily the object of this investigation was to determine whether or not there is a gradual decrease in the fertility of hybrids in successive generations leading finally to sterility and the extermination of the race- a fate that Dr. Alfred Mjöen, of Sweden [sic.], predicts for the American people

200 “Ministers Reaffirm Stand on Wedlock of Close Relatives: Assert Experiments with Rat Cannot Serve as Basis for Radical Conclusions on Marriage.” Evening Public Ledger (Philadelphia). October 2, 1915.

92 because of the continued intermingling of different racial stocks.”201 Mjöen, a

Norwegian pharmacist and popularizer of eugenic concepts, had warned about the dangers of “disharmonious combinations” resulting from the union of ostensibly distinct racial types. Several American eugencists sympathized with this view, whatever their opinion of Mjöen’s acumen as a scientist, notably Charles Davenport and Charles

Stockard.202 The concept hinged on the notion that natural selection had assembled in each race a set of complementary characters best suited to its environment. It was assumed that crossing between distinct racial types would break up these naturally

“harmonious” assemblages of traits and result in something ill adapted to any real environment, something unnatural. Hybridization of domestic animals and domestication itself, by creating a dichotomy between the natural and artificial among animals, were analogically important in the discussion of this problem. Harvard geneticist William E.

Castle sharply criticized the idea of disharmonious crossing in the middle 1920s, especially the misapplication of experimental data on animals for social purposes. Castle concluded, “So far as a biologist can see, human race problems are not biological problems any more than rabbit crosses are social problems. The rabbit breeder does not cross his selected races of rabbits unless he desires to improve upon what he has. The sociologist who is satisfied with human society as now constituted may reasonably decry race crossing. But let him do it on social grounds only. He will wait in vain, if he waits

201 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1924. p.14. WIL. 202 Davenport offered his opinion of Mjöen to Edwin G. Conklin in 1926: “He is trying hard to establish a reputation as a scientific man but, I should judge, he does rather little nose-to-the- grindstone work himself… He is sort of mid way between Madison Grant (shall I say) and Michael Guyer.” Davenport to Conklin August 26, 1926. Charles B. Davenport Papers B.D27 Series I, Folder “Conklin, Edwin Grant, 1863-1952” No.2. APS.

93 to see mixed races vanish from any biological unfitness.”203 In later years Davenport, who initiated a study ostensibly outlining the dangers of race crossing in Jamaica after

Castle’s withering critique, defended his position on “non-harmonious, or ill adapted combinations of traits” to Castle on the grounds that “we differ a little in that you stress the monstrosities and I stress the absence of so nearly complete adaptation as is found in the ‘wild’ species.”204 Davenport’s analysis of their differences is perspicacious.

Castle’s view was predicated on “monstrosities,” that is, a range of variations or individual differences, while Davenport maintained the concept of an idealized wild type, one almost perfectly suited to its life in nature. Although both men understood well the genetic basis of heritable variations, Davenport’s thinking about their contribution to adaptive fitness was less contingent and relational than Castle’s.

Helen Dean King avoided entering this debate in her published works, despite the perceived applicability of studies on inbreeding and hybridization in animals to the discussion. Again, she felt freer to speculate on the sociological implications of her work to the press. After explaining the results of her experiments on inbreeding and hybridity in rats to the Wilmington, Delaware Sunday Morning Star, she explained how these processes had shaped the history of human civilization. “There is no evidence,” she averred, “to show that consanguineous marriages were injurious to any of those early nations” which practiced it widely: the Ancient Greeks, Phoenicians, Jews, Incas,

Teutons, and Egyptians. She was more pessimistic, however, about the effects of hybridity: “The decline of the Greeks and the Egyptians came when they ceased to be an

203 Castle, W.E. Biological and Social Consequences of Race Crossing. American Journal of Physical Anthropology. 1926. Vol.9(2):156. 204 Davenport to Castle December 8, 1930. Charles B. Davenport Papers B.D27 Series I, Folder “Castle, William E., 1867-1962” No.13. APS.

94 exclusive people and the strength and vigor of the race were sapped by vices and luxuries introduced from other countries.”205 Through the use of questionable historical claims,

King made moral shortcomings and cultural differences synonymous with the specter of hereditary taint, a theme that had motivated the American eugenics movement to a strongly anti-immigrant political position.

Captive Breeding and the Elusiveness of Control

In 1919 Helen Dean King began her own experiments on captive bred gray

Norway rats. She explained in the Director’s Report for the year that her investigations of the “physiological relation between coat-color and body growth in extracted strains of rats” had to be stalled until she could compare the results to a growth curve for the wild gray Norway. King wrote, “It seemed advisable, therefore, to undertake a rather extensive series of breeding experiments with the Norway rat… It is planned to continue the breeding of the wild rats for several generations, if possible, and thus to study the effects of domestication on this form.”206

The long-term goal of this study, to reveal the effects of long continued captive breeding, may have come about through Donaldson’s influence, as he was listed as a co- investigator on the project in the Director’s Report for the following year.207 Although this experiment was cast in precisely the mold of Hatai’s abandoned captive breeding

205 Vaynow, Andrew “New Facts on Brother and Sister Marriages” The Sunday Morning Star (Wilmington, Delaware). November 29, 1931. 206 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1919. p.14. WIL. 207 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1920. p.9. WIL.

95 study, the precedent was never mentioned, as if expunged from the institutional memory.

Thus, King was able to write in the Director’s Report for 1921, “This study, apparently the first investigation of the kind ever undertaken, has already given some interesting results.”208 As with Hatai’s experiment, King’s was referred to internally as the

“domestication experiment.” In Donaldson and King’s published reports, however, the scope of the study was limited to “continued captivity” as a factor contributing to the features that distinguished the wild gray Norway rat from the domesticated albino. The effects of captivity were to be evaluated thus: “the alterations which prove to be similar to those now found in the Albino could be credited to captivity and treatment, while other modifications in the Albino not represented in the captive Norway could be referred to albinism.”209 They also reiterated, briefly, the key distinctions between the stock albino and wild gray Norway, all of which they planned to monitor in their strain of captive grays:

The wild Norways are more excitable and much more savage. They gnaw their cages. The body weight is less for a given body length, hence it is a slighter animal. The skeleton is relatively heavier, also the suprarenals (both sexes), and the testes and ovaries. The thyroid is of like weight, but the hypophysis is distinctly lighter, in both sexes- this last difference being more marked in the female. On the other hand, the brain and the spinal cord are both heavier than in the Albino.210

The experiment began by taking 36 (20 female and 16 male) wild Norway rats of various ages caught throughout Philadelphia into captivity. The staff of the

Institute knew from their shorter-term hybridization studies that breeding wild caught

208 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1921. p.8. WIL. 209 King, H.D. and Donaldson, H.H. Life Processes and Size of the Body and Organs of the Gray Norway Rat During Ten Generations in Captivity. Anatomical Memoirs. 1929. Vol.14:6 210 Ibid. p.7.

96 gray rats in captivity posed a challenge. Wild Norways often became sterile under captivity, and if they produced a litter, the slightest disturbance would cause them to kill their young. With the exception of one relatively tame gray mother who nursed her own litter, all other litters were fostered with lactating albino rats. This strategy caused no changes in growth, according to King, who divided the litter cast by the “tame” gray and gave half of the young to an albino foster; there were no apparent differences in development. Donaldson also made anatomical measurements of the grays after months in captivity and compared them to those of grays freshly caught. He declared that months-long captivity had not altered these anatomical points, suggesting that captivity limited to a single generation was not able to significantly change the structure of the rat.

Once these initial concerns had been settled, King noted her “interesting results” after only four generations. They seemed to “indicate that the rapid growth and early maturity of the albino is very evidently the result of domestication.” King also noted that many of the captive grays “are exhibiting characteristics found in animals of the early generations of the inbred Albinos, namely abnormal teeth, a tendency to sterility, and a decrease in constitutional vigor.” Diet, which King identified as the problem of the early inbred albino rats, was ruled out in this case. Instead she attributed this general decline to a

“period of readjustment to new conditions,” from which they recovered in subsequent generations, likely because of the investigator’s selection against litters containing “weak individuals.”211 King argued that this selection was necessary, especially in the early stages of the project, but it may also have served to complicate the interpretation of her results.

211 King, H.D. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1921. p.8. WIL.

In the division of duties between the investigators, King was charged with observing and analyzing changes in the “life processes’ of the captive grays, including body weight, fertility, sterility, sex ratio, reproductive period, mortality, and behavior.

Some of these factors, such as mortality, fertility, and sex ratio showed few or ambiguous changes as a result of sustained captivity. King claimed, though, that the remaining factors gradually approached the conditions prevailing in the domesticated albino. The reproductive period, for instance, began earlier in late generation captive grays than in those of the earlier generations. It also ended slightly later with a delay in menopause among females, extending the reproductive period a total of two months. Body weight increased in relation to body length. Sterility fell from a high of 70 percent in the first generation to less than 10 percent in the tenth. Perhaps the most crucial modification was in behavior. King credited nearly all of the other changes to the lessening of “nervous tension” in the gray rat after multiple generations in captivity. These results, unlike those in King’s inbreeding study, conformed well to conventional wisdom about domesticated animals.

Donaldson analyzed the changes in anatomical structure among the captive gray rats, focusing on those points which diverged most strongly between wild-caught

Norways and domesticated albinos and which had first brought his attention to the question of domestication. Donaldson offered his analyses of first ten generations of captive bred gray rats in the paper he coauthored with King in 1929. He agreed with her that, “these captive Grays… are rats which grow normally in length, but tend to become

98 fatter as the generations progress.”212 Some of the most critical glands and organs, the adrenal glands, the pituitary, and the brain, displayed an overall change in weight closer to that of the albino rat. In each case, however, the change was not progressive, but was either abrupt or without a distinct trend. The gonads, in turn, were almost entirely resistant to the effects of continued captivity, which, for Donaldson, “was unexpected, in view of the much lower weight values for the gonads shown by the Albinos.” Donaldson was forced to conclude that “[t]en generations of captivity have, by no means, served to give the captive Grays the organ constitution of the Albinos.”213 When his teleological expectations for the feralization project and the first captive breeding experiment are remembered, these results must have been a disappointment. He left the door open, however, for further change in the future: “To obtain the full effects of captivity on the body and its organs will probably require many more generations and, therefore, any conclusions from the present data must be largely tentative.”214

The decision to limit the first report to the first ten captive generations was not wholly arbitrary; shortly after the tenth generation the strain “broke,” issuing coat color mutants that were subsequently perpetuated as separate strains alongside the captive grays. When the first report was released in 1929 Donaldson already knew that he had little to expect from the next ten generations (King had, by then, achieved 19 generations in captivity). His report of these later generations, though, never surfaced. As he and

King began preparing the second paper (up to 25 generations), Donaldson died of heart

212 King, H.D. and Donaldson, H.H. Life Processes and Size of the Body and Organs of the Gray Norway Rat During Ten Generations in Captivity. Anatomical Memoirs. 1929. Vol.14:101. 213 Ibid. p.106. 214 Ibid. p.100.

99 failure early in 1938, just days after his last visit to the laboratory. Plans to finish his portion of the paper fell through. The recipient of his dissection records, the medical geneticist Clyde E. Keeler, left the impression that Donaldson had been disappointed by the failure of the captive gray rats to acquire the marks of domestication.215

King, however, suggested that in their “life processes,” the captive grays continued their progressive march toward the behavioral and reproductive features of domesticated albino rats in her manuscript on the first 25 generations of captive breeding.

In body weight, the captive grays actually exceeded any of the albino strains at the

Wistar. King highlighted the agreement between her results and popular thinking about domestication: “The effects of captivity on the rate and extent of body growth in gray

Norway rats are similar to those that have occurred in various domesticated animals, such as cattle, horses, and sheep. These animals, as Darwin (1875) has shown, grow more rapidly and attain a much larger size than did their wild prototypes.” The length of the reproductive period continued its steady climb and sterility virtually disappeared after the twelfth generation. Again, King claimed the changes in behavior to be some of the most significant. She maintained that while “there was no noticeable difference in the conduct of individuals in any two succeeding generations… a comparison of behavior in rats in early and in late generations indicates clearly the striking modifications that occurred,” principally in the reduction of fear and “nervous tension.”216 The sex ratio, meanwhile, stubbornly remained around the mean established in the first few generations. King accepted this as evidence of inherent differences in the gray and white strains and

215 Keeler, C.E. and King, H.D. Multiple Effects of Coat Color Genes in the Norway Rat, with Special References to Temperament and Domestication. Journal of Comparative Psychology. 1942. Vol.34:242. 216 King, H.D. Life Processes in Gray Norway Rats During Fourteen Years in Captivity. Anatomical Memoirs. 1939. Vol.17. p.49-50.

100 adopted a taxanomic equivalence between races in rats and races in humans to suggest that “it may be of interest to compare the sex ratios in the white races of man with those for colored races, chiefly of negro descent.”217 While many of King’s explanations for the changed features of the captive gray rats remained the same as in the 1929 report, she laid more emphasis upon physiological and genetic explanations in 1939, rather than adaptation to the altered conditions of life as she had earlier. King’s studies with inbreeding and hybridization brought her greater knowledge of genetics, and this was expanded by her investigations of the coat character mutants derived from the captive gray strain. Milton Greenman and Henry Donaldson, whose views of biology belonged to an earlier generation and tradition than King’s own, both died between the publication of King’s first report on captive bred rats and her second. This may have left her freer to interpret her results according to her own view. In her 1939 report she countered

Greenman and Donaldson’s basic position “that variability of every kind is caused directly or indirectly by changed conditions of life” by suggesting that “[i]n light of our current knowledge of genetics, it is probable that changed conditions of life in captivity had little effect” on the variability of certain features in the captive bred strain.218

The appearance of coat color mutants in the captive gray line was cause for renewed excitement for the captive breeding project and for the larger goal of explaining the differences between the wild gray and domesticated albino rat. The Director’s Report for 1926 declared that “the most spectacular of the results obtained in domesticating the

217 Ibid. p.41. Donaldson similarly equated rat and human races in a letter to Davenport, discussing the latter’s race-crossing research in Jamaica. See Donaldson to Davenport November 23, 1929. Charles B. Davenport Papers B.D27 Series I, Folder “Donaldson, Henry Herbert 1857-1938” No.2. APS. 218 Ibid. p.24.

101 wild gray rat are the indications that this animal… is beginning to show signs of genetic modifications by the appearance of ‘sports’ or mutations.”219 Early in the year a litter of true-breeding black rats were born to a gray female in the captive strain. In the next generation the mutation “dilute” appeared also in a litter born to a captive gray. Before year’s end several coat colors appeared in another litter, including an albino. Donaldson wrote of the discovery later, “This is the first time that original mutant albinos have been available for study, and the material is of fundamental importance for the determination of those characters associated directly with albinism as contrasted with the characters developed by long-continued captivity,” again without reference to Hatai’s claims to an identical discovery in 1912.220 These mutants, the key to distinguishing between the factors of albinism and the “struggle for existence” that Donaldson had hoped to isolate for more than 20 years, also further complicated the picture of the effects of domestication and drew King into more basic questions about the nature of their inheritance.

Both King and Donaldson refrained for the most part from making specious claims about the applicability of their work on captive bred rats to human problems, save for King’s suggestion that distinctions in differently colored strains of rats had bearing upon distinctions in human races. The annual Director’s Reports displayed more speculation on this line. The 1926 Report asked of King and Donaldson’s investigation into “the changes which take place in the Norway rat under the influence of domestication as the brain weight gradually drops from that of its progenitors toward that

219 Author unknown. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1926. p.18. WIL. 220 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1931. p.16. WIL.

102 of the albino rat… Will it eventually reach the low limit of the albino rat? Shall we learn from experiments on the rat what may be accomplished in the development of the human brain?”221 To anyone familiar with Donaldson’s work and his rationale for choosing the albino rat as the vehicle for his research, this last question was purely rhetorical.

Donaldson had been convinced, prior to his arrival at the Wistar Institute, that the albino rat was the ideal model for human nervous system growth. He averred, consistently, that the experimental results that he and his assistants achieved with the rat were directly applicable to humankind. The captive breeding experiment, the project that he believed most nearly addressed the entirety of the problem of domestication, was no different. By the early 1920s, Donaldson had “taken steps to obtain material for a more extensive study of [the] change in cranial capacity in other mammals.”222 At the festschrift for

Constantin von Monakow in 1923 Donaldson presented a paper on the diminution of cranial capacity from the wild cavy to the domesticated guinea pig. The paper also included data on lions reared in zoological parks (a study strikingly similar to Harrison

Allen’s plans to investigate “domesticative” changes in zoo animals).223 This was one of

Donaldson’s rare departures from the rat as a research animal after 1920. The reason for this unorthodoxy was to generalize the reduction in brain size as a result of domestication to all mammals, preparing the ground for the inclusion of civilized humans. There is also evidence that by the late 1920s, discussions at the Institute suggested that the application of these results should extend beyond the anatomical to the social. In Milton Greenman’s

221 Author unknown. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1926. p.16. WIL. 222 Donaldson, H.H. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1922. p.8. WIL. 223 Donaldson, H.H. On the Cranial Capacity of the Guinea Pig- Wild and Domesticated. Archives Suisses de Neurologie et de Psychiatrie. 1923. Vol.9:206-222.

103 report for 1927, following a description of the captive breeding project, he suggested the end uses of the knowledge gained from rats: “Much of this work has a direct bearing upon race betterment and an economic importance in connection with domesticated animals… The Institute will find it advantageous to combine such work with comparative psychology and pathology.”224

By the early 1920s Helen Dean King had been trusted with the execution of significant portions of Donaldson’s domestication program, including the inbreeding, hybridization, and captive breeding experiments. Her work, which excited discussion from within the Wistar Institute and without, was curiously punctuated, abruptly and unexpectedly, at the beginning of the year 1922. Greenman notified King that her position would be terminated. According to Conklin, who also reported that the

Advisory Board had not been informed of this decision, “Greenman [put] the whole matter on the basis of financial inability. He [said] they have had a big deficit during the past year and have got to save money. He furthermore [insisted] that Miss King’s appointment [had] always been temporary from year to year, and that her work, he thinks, does not fit in with the work which they are doing at the Institute as well as it would fit with the work in other Institutes or Universities.”225 A Miss Pearson, one of King’s friends, wrote on her behalf to friends and colleagues in the field of biology as part of a late campaign to either save King’s position or find her another. Word quickly reached a number of influential biologists, including Conklin, William E. Castle, Charles B.

224 Greenman, M.J. in The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31st, 1927. p.12. WIL. 225 Conklin to Pearl February 2, 1922. Edwin Grant Conklin Papers. Series II, Box 18, Folder 9 “Raymond Pearl.” PUL. See also copy of a letter from Conklin to Castle, January 28, 1922 in Charles B. Davenport Papers B.D27 Series I, Folder “Castle, William E., 1867-1962” No.12. APS.

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Davenport, Raymond Pearl, and C. E. McClung. Castle, like all of the correspondents, was surprised and dismayed by the news. While he could not evaluate “the merits of the two sides to the case,” he opined to Davenport, “I do entertain a high regard for Miss

King’s scientific work, and I think it would be a great pity to have it interrupted… It is also clear that she can not work to the best advantage under irritating conditions. Hence it would be well for her to work elsewhere, if she can, rather than patch up a modus vivendi at the Wistar.”226 Conklin met with Donaldson and noted that “he was plainly much troubled because of the matter but was evidently not free to express himself.”

Conklin added, “I am confident the whole matter is some incompatibility between

Greenman and Miss King.”227

Ultimately, this rift was smoothed over, but its cause remains unclear. The

Minutes of the Board of Managers meeting on February 16, 1922 echoed Greenman’s earlier reasoning: funding was short and “the time had arrived when it appeared necessary to decide whether the Institute could afford to continue a piece of work which, through Dr. King’s unusual ability, had grown to considerable proportions in an important field somewhat outside the Institute’s present program of research.”228 King’s research, especially the perpetuation of multiple lines of experimental rats (inbred albinos, several hybrid strains, various mutant strains, and captive grays) required increasingly large portions of the Institute’s budget. By 1922, the litters she had produced for experimental work numbered in the thousands, and the individual rats in the

226 Castle to Davenport January 30, 1922. Charles B. Davenport Papers B.D27 Series I, Folder “Castle, William E., 1867-1962” No.12. APS. 227 Conklin to Castle January 28, 1922. 228 Minutes of the Board of Managers of the Wistar Institute Meeting February 16, 1922. WIL.

105 hundreds of thousands. Her particular interests, in the sex ratio, in fertility and sterility, and in other “life processes,” departed from the focus upon the brain that Greenman had established as the hallmark of the Institute when he became its Director in 1905. Thus, his concerns over King’s work seemed legitimate. He neglected, however, that

Donaldson had suggested most of these studies in order to isolate the factors of domestication and used King’s rats to evaluate the effects of these treatments on the growth of the brain. King’s researches were, according to Donaldson’s broad and circuitous approach to biological problems, still part of a program, albeit one that was less “concentrated and coordinated” than Greenman idealized. Funding was always a limitation for the Institute. However, months before threatening to fire King, Greenman had been able to set aside $30,000 for a new wing to house the Institute’s rat colonies, replete with, according to the press, the kind of luxuries that would make most humans envious.229

Conklin’s suggestion, that the matter was more personal than practical, may have hinged on both actors’ reputations as difficult and controlling personalities. Much of

King’s reputation came from her attempts to control the uses of her rats and her data.

When Franz Boas requested data from King’s early generations of inbred rats, in his own attempt to find the human implications of this data, King refused. Donaldson offered by way of explanation to his friend: “It strikes me as possibly a case of ‘nerves’ due to excessive work [and] will remedy itself soon, I hope. In the meantime my apologies for the existing state of affairs.” William E. Castle’s letters to L.C. Dunn in later years suggest that King’s reputation was familiar beyond the walls of the Wistar Institute:

229 “Will Build $30,000 Home for Super-Rats in Philadelphia, Pa.” The Columbia Evening Missourian. August 24, 1921.

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“…Dr. King, as you doubtless know, has peculiar ideas about cooperation, and has to be handled gingerly.”230 Clyde Keeler, who was party to an entirely different controversy involving King almost 20 years later, complained bitterly of her jealousy and vindictiveness toward him.231 The issue of King’s gender and the general struggles of women in science at this time were undoubtedly reflected in these somewhat patronizing portraits, in the protectiveness and insularity that prompted them, and, indeed, in the tentativeness of her position at the Wistar.

Greenman’s own intransigence came to public light well after his death in C.

Judson Herrick’s biography for his departed friend and former Wistar professor, George

E. Coghill. The story of Coghill’s employment and ultimate dismissal from the Institute parallels that of King’s near demise 13 years earlier. Herrick, who worked alongside

Coghill for two seasons at the Institute’s Effingham B. Morris Experimental Farm in

1929, recalled Greenman’s “dictatorial attitude and sometimes stupid interference… He was constitutionally unable to delegate details and overtaxed himself with many minor tasks. I have a vivid mental picture of him on a raw spring day when the outside wires were being strung on poles. With a pocket knife he was vigorously cutting off the insulation from the end of a thick wire, while two competent electricians stood by with their hands in their pockets.”232 For Coghill, this grew from a minor inconvenience- “I am sure he means well; he is simply driven by goads of detail, and sometimes, I think,

230 Castle to Dunn November 28, 1938. L.C. Dunn Papers B.D917 Series I, Box 4, Folder “Castle, William Ernest” APS. 231 See the following section in the present volume for more on the nature and result of this controversy between King and Keeler. 232 Herrick, C.J. George Ellet Coghill: Naturalist and Philosopher. Chicago: University of Chicago Press, 1949. p.53.

107 nearly broken,”- to an insufferable flaw in Greenman’s personality.233 Greenman became only more anxious and controlling as it became clear that the Institute was overextended programmatically and especially financially. Herrick counted both Coghill and

Greenman among his personal friends and ascribed the controversy between them to mistakes made by both men. In Greenman’s case alone, however, were the mistakes

“pathological.” The result of Greenman’s autocratic abuses was an “atmosphere… surcharged with covert animosities.”234 A similar atmosphere may have prevailed in 1922 as King’s fate at the Institute was being weighed.

Prior to this controversy, of course, King had made incautious claims about the sociological implications of her work on rats, statements that were sensationalized by the press. King obliged interviewers who sought the opinions of this “interesting woman

[who] draws conclusions about all sorts of interesting things just from experimenting with rats and tadpoles and whatnot” on a range of topics, usually eugenical, including consanguineous marriage, miscegenation, and the impact of the Great War on “the race.”235 It is doubtful that these opinions played a role in Greenman’s decision concerning King, as he echoed her pronouncements in several of the same newspaper articles. Rather, King’s eugenical statements were part of a long tradition of speculation about the human implications of the work performed at the Wistar and a more recent trend of finding ways to apply these implications to social problems.

233 Ibid. 234 Ibid. p.54. 235 M’Liss “Banish Bachelor and Revive Race: Dr. Helen Dean King, Embryologist, Thus Would Solve After-war Problem.” Evening Public Ledger. February 1, 1917.

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Greenman, for his part, encouraged the conceptual leap from experiments performed on rats to their human implications, especially with a social ideal in mind. He attempted as much himself in an interview with a reporter from the United Press in which he described the expenses undertaken to provide Wistar rats with the right conditions for exercise, mental development, proper health, and breeding to make them truly “super- rats.” The reporter gave some indication of the social ideal that Greenman maintained for his rats: “The super-rat, like the super-man, is not, as Nietzsche would have it, of the warrior type, but is a gentleman or a gentlewoman, an aristocrat at heart, although democratic in his ways.” Greenman clarified the goal in his own words: “Experience has shown that it takes about three generations to develop the aristocratic gentleman rat from the wild, rough-neck variety that is most familiar. Figuring on the relative rate of living of the rat and man, it would probably take ten years of intensive training to bring about the same change to a marked degree in the human being.”236 Thus, as late as 1921,

Greenman maintained that a remarkable measure of control was possible over modern man (justified, of course, by work on the albino rat), however, the controversy that arose with King and later with Coghill suggests the difficulty Greenman faced in maintaining control of his institutional vision.

236 The Columbia Evening Missourian. August 24, 1921. Greenman’s estimate of ten years to reform a human being overlooks the distinction between change across generations and change in an individual lifespan, an indication of how intimately linked phylogeny and individual development were in his biological thinking.

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Institutional Extension, Social Application

The human focus of the Institute was determined first by its role as the guardian of the Wistar-Horner anatomical collections. Next, the philosophical ferment of transcendental morphology, established by Isaac Wistar, John Adam Ryder, and Harrison

Allen, favored the view that the history of life was one of dynamic and directional development, with the general progressive force culminating in humankind. Finally, under Milton Greenman’s direction, the Institute fortified its ties to physical anthropology, despite the methodological shift to performing experiments on animals.

One of the first proposals from Greenman’s new Advisory Board was to “initiate a study of the racial anatomy of the brain,” blurring the line between physical anthropology and the Institute’s stated field of neurology.237 Ales Hrdlicka, who praised Donaldson for his contributions to anthropology, convinced the Wistar Institute to publish his American

Journal of Physical Anthropology in 1918. In the Journal, Hrdlicka identified the

Institute as an important, though underutilized, resource for anthropologists.238 Henry

Donaldson’s longtime friend was, incidentally, the anthropologist Franz Boas, who helped Donaldson to link his albino rats to humans by appealing to an assumed common experience, domestication. In later years, the museum of the Institute displayed an

“[e]xhibit showing the principles of variation found in several breeds of dogs, translated into terms of variation in man,” resulting from the work of Dr. Charles R. Stockard, a

237 Greenman, M.J. Report of the Director… Appendix IX. p.179. 238 Hrdlicka, A. Physical Anthropology: Its Scope and Aims; Its History and Present Status in America. American Journal of Physical Anthropology. 1918. Vol.1(3):110-111.

110 longtime associate of the Wistar Institute and, later, a member of the Advisory Board.239

In his work on purebred and hybrid dogs, Stockard championed the analogy between civilized humans and domesticated animals and this interpretation was preserved as his collections were displayed at the Wistar Institute.

Despite the Wistar Institute’s mark of independence, Greenman became an advocate for collaboration early in his tenure as Director. Some of the earliest attempts were directed at growing the Institute’s credentials in basic biological research, including a failed bid to acquire the Marine Biological Laboratory (MBL) at Woods Hole in

1907.240 The setback, though, did result in a longstanding partnership between the MBL and the Wistar. Less than ten years later, Greenman sought affiliation with a clinic or school where the neurological focus of the Institute could find wider application to social issues. He found a willing partner in Edward R. Johnstone, administrator of the Vineland

Training School in neighboring New Jersey, one of the first institutions devoted to the

“education and care of feebleminded children” in the United States. Johnstone had institutional ambitions similar to Greenman’s. Since taking over the Vineland in 1900, he had begun planning to expand the School’s role from education to research and ultimately to political action. These efforts began with the meetings of the informal

“Feeble-minded Club,” the first held in Philadelphia in March 1902. Ties with the Wistar

Institute were forged almost as early: Henry Donaldson was listed as a member in 1906, the year of his arrival in Philadelphia.241 The Feeble-minded Club was instrumental in

239 Untitled Inventory of the Wistar Museum March 3, 1953. p.8. Collection 5 The Wistar Institute Series IX, Box 2, Wistar Museum Exhibits 1954-55. WIL. 240 File “Marine Biological Laboratory” WIL. 241 Zenderland, L. Measuring Minds. p.62.

111 establishing a new psychological research laboratory at the Vineland School, especially through the largess of one of its members, the Philadelphia soap magnate Samuel S. Fels.

Johnstone selected another Club member, the psychologist Henry H. Goddard, to lead the laboratory. As the biannual meetings of the Feeble-minded Club shifted to Vineland, a new, more formal Committee on Provision for the Feeble-minded was formed in

Philadelphia in 1915 with Greenman as chairman and Johnstone as secretary.242 Despite the Committee’s official title, its stated purpose was not provision, but “‘[t]o disseminate knowledge concerning the extension and menace of feeblemindedness and initiate methods for its control and ultimate eradication from the American people.’”243 In a nationwide lecture campaign, the committee, represented by Johnstone’s brother-in-law

Alexander Johnson encouraged the view that feeblemindedness was a brooding menace.

Johnson was aided by the findings emanating from the laboratory at Vineland, especially

Goddard’s dour assessment of the mental quality of immigrants arriving at Ellis Island and his landmark study on the persistence and heritability of feeblemindedness in the pseudonymous Kallikak family. Fels, for his part, helped to foster a distinctly eugenical research agenda at Vineland. As one supervisor recalled, “As far as the Training School was concerned, [Fels] made it clear that his sole interest was in research. His concern

242 Quoted from an official release of the Committee in: A Committee to Eradicate Feeblemindedness. The Survey. 1915. Vol.34:369. For further information on the origin of the Feeble-minded Club and the later Committee on Provision for the Feeble-minded see Zenderland, L. Measuring Minds. p.61-63 and Trent, J.W. Inventing the Feeble Mind: A History of Mental Retardation in the United States. Berkeley: University of California Press, 1994. p.155-157, 172-183. 243 Ibid.

112 with the feebleminded was not with their training and welfare, but as he frankly put it- in getting them off the earth.”244

In 1919 Wistar Advisory Board issued a special statement of the activities of the

Institute, including its extension beyond the laboratory. The Board members wrote,

“[f]or some years past the Institute has been cooperating with the Training School at

Vineland, N.J. in the study of certain phases of neurology presented by the mentally defective,” and indicated that in 1918 the collaboration was made more formal, including a semi-permanent staff of Wistar researchers placed at the Vineland School. The increased Wistar presence in Vineland coincided with Goddard’s departure from the psychological laboratory. Stanley Porteus, Goddard’s replacement as Director of

Research, conjectured in his 1969 autobiography that Greenman had been involved in

Goddard’s ouster after growing skeptical about the validity of his work on the

Kallikaks.245 Whether Greenman played such a direct role is questionable, though

Goddard clearly felt that his relationship with his patron, Fels, had been poisoned by interlopers. In a letter to Fels upon his departure, Goddard wrote:

It was painful for you to come to the decision that you made known in your letter. It is painful for me to have you come to such a decision, but it is vastly more painful to realize that I am largely responsible for your decision and to always have to regret not making the work appeal to you… [H]ad it not been for outside influences I think that you would have gone on [and] we would by now have had results that would have satisfied you… I am responsible. I know I should have been able to continue, but I believe that there were subtle influences at work that I did not understand and hence could not counteract.246

244 Porteus, S.D. A Psychologist of Sorts: The Autobiography and Publications of the Inventor of the Porteus Maze Test. Palo Alto, California: Pacific Books, 1969. p.64. 245 Ibid. p.68. 246 Quoted from Wehmeyer, M.L. and Smith, J.D. Leaving the Garden: Reconsidering Henry Herbert Goddard’s Exodus from the Vineland Training School. Mental Retardation. 2006. Vol.44(2):154.

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Implicitly supporting Porteus’ conjecture, Greenman and the Wistar Institute stood to gain significantly from the reorganization of the psychological laboratory at

Vineland. After Goddard’s departure in 1918, research at the Training School was divided into more autonomous groups, instead of falling under the immediate supervision of the Research Director. This change and funding from Fels allowed for “extending the neurological research [of the Wistar Institute] in cooperation with the Vineland Training

School.”247 The extension consisted not only of maintaining a staff of Wistar Institute researchers at the Training School, “fitted to follow and record the growth, behavior

(psychological), and clinical history of the four hundred and ninety children in the institution,” but also of finally applying the work on the rat to the solution of a pressing social problem, the menace of the feebleminded. The Advisory Board explained, “At the same time it was planned to extend the present work on the growth of the nervous system of the rat, as the results obtained with the rat are applicable to man, and also to lay stress on the collection of data on the normal human nervous system.”248 It then described a tripartite research agenda involving clinical observations on feebleminded children at

Vineland, post-mortem measurements at the Wistar, and experiments with rats suggested by the comparison of normal and defective human material. By 1920 Greenman had assured that the Wistar Institute and its work on the rat were well placed to find a real application to the social, and particularly eugenical, concerns of the moment.

Historian of science Jeffrey Brosco observed, as an aside in his history of the

Wistar Institute, that “Greenman was probably Jewish, Hatai was Japanese, King was

247 Activities of the Wistar Institute April, 1917 to April, 1919: A Statement by the Advisory Board. The Anatomical Record. 1919. Vol.17(3):179. 248 Ibid. p.178-179.

114 female, and Donaldson was permanently disabled by tuberculosis in the 1890s. It is interesting that this socially marginalized group- given the sensibilities of the early twentieth century- sought to make their institute the ‘center’ of academic anatomy and biology” in the United States.249 It is equally intriguing that the research, interpretations, statements, and associations of this core of the Wistar staff resonated with the

“sensibilities” of the time, sensibilities that, as Brosco intimated, could be unkind to the disabled, women, ethnic minorities, and immigrants. Recent historical work has expanded and nuanced the view of these early 20th century attitudes, especially as they related to the causes of eugenics and social hygiene. Scientific supporters of eugenics were a far more heterogeneous group than was once recognized. Proponents of the eugenics movement hailed from diverse backgrounds and harbored different prejudices and ethical and political viewpoints. What seemed to unite them, and has since fallen out of fashion, was the moral stance that individual desires should be sacrificed for the rationally determined social good. In the context of a socially directed morality and an ongoing dialogue about the best course toward social improvement, advocates of eugenics did not think of themselves as morally blind, but rather, as earnest reformers.

The increasing alignment of the work of the “socially marginal” core staff of the Wistar

Institute with the goal of race improvement during the 1910s and 1920s becomes more explicable in this view. As late as 1936, Helen Dean King suggested the unity between this Progressive Era ethic and the eugenical implications of her research:

When the time comes that an enlightened people take thought of the wellbeing of the generations to come and realize the great value of favorable combinations of genetic factors that produce unusual ability,

249 Brosco, J.P. Anatomy and Ambition. p.17.

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marriage will be based not only on the physical fitness of the individuals but also on their recorded pedigree for several generations. The race can then be vastly improved through consanguineous marriages in families in which the members show exceptional mental and physical endowment in ways that are of value to them and to the community at large. Many of the ills to which man is at present subject will vanish. Superior and desirable traits will appear in an ever increasing number of individuals and eventually become the heritage of the race.250

Conclusion

Milton Greenman had, with the aid of Henry H. Donaldson, given the Wistar

Institute a new, experimental research direction in 1906. Meanwhile, the philosophical orientation of the Institute remained influenced by the tradition of transcendental morphology in which both Greenman and Donaldson had been trained. Tensions emerged between these two approaches, but until Greenman and Donaldson’s deaths in

1937 and 1938, respectively, there was little sense that the two were incompatible.

Greenman and Donaldson continued to emphasize comparative methods so as to reveal patterns of directed development, especially with the goal of applying their work to the zenith of progressive evolutionism, humankind. As Donaldson tried to set the research agenda in his early years at the Institute he began to focus more closely on one particularly valuable comparison, between the patterns of growth in the brains of humans and albino rats. The similarity of these processes suggested to Donaldson that the albino rat was the ideal model for human development. His determination that domestication was to blame for the pace and extent of the development of the albino rat’s nervous system made the domestication of the rat a too-tempting corollary for the nature and fate

250 Quoted from Ogilvie, M.B. Inbreeding, Eugenics, and Helen Dean King. p.498.

116 of modern humans. As Donaldson’s experimental program to determine the causes of domestication expanded through his own efforts and those of Hatai and King, so did its claims to social relevance. During this expansion Greenman often worried about philosophical and programmatic coherence, at one point contemplating King’s removal from the Institute for straying too far from its established focus and philosophy.

C. Judson Herrick painted an unflattering portrait of Greenman as an administrator: controlling, harried, and (to borrow a phrase from their generation of neurological experts) bordering on neurasthenic. Some 25 years earlier C. E. McClung, a member of the Institute’s Advisory Board, had given a very different characterization of

Greenman:

It is possible to adopt an autocratic method and put a man in charge of strong personality and intense convictions and authorize him to carry forward a personal program…There is another method of reaching out and making numerous contacts and thereby profiting from the abilities of many people, while at the same time a corresponding return is made to them. This is not so easy as the one-man method, because the involvement of numbers… naturally complicates matters. But it can be done, and The Wistar Institute has shown the way by its last twenty years of operation. As a result it has had an influence all out of proportion to its size, because, through the foresight of its Director, it elected to render the service that many desired, rather than the one he determined.251

There appears to be some truth in both depictions of Greenman (and consequently of the Institute he directed). He was praised for his liberal support of varied research at the Wistar, but according to his own decree, this was done in the service of a single, overarching program. He extended the Institute in directions his predecessors had not imagined and yet worked incessantly to ensure that the Institute fulfilled its original goals

251 McClung, C.E. in Bulletin No.6 of the Wistar Institute. 1925. p.72.

117 and spirit. It seems that for most of his 30 years as Director, Greenman balanced these demands ably, but at those points when extension threatened the identity of the Institute,

Greenman was at his most intolerable. The Wistar Institute, at least according to Herrick, followed closely Greenman’s tenor.

By midcentury the approach to research at the Wistar seemed as antiquated and obsolete to outsiders as the moldering anatomical collections occupying its basement.

Leonard Hayflick, doctoral student at the University of Pennsylvania, recalled his impressions of the Institute where he began working in 1952. “Wistar,” he relayed, “was right out of the last century.”252 Hilary Koprowski, an ambitious researcher who replaced

Edmond Farris (Greenman’s less ambitious successor), said of his decision to come to the

Wistar, “I liked it because it was dead.”253

For the first three decades of the century, however, the Wistar Institute was far from moribund. The typological and teleological thinking at the Institute was not out of place in the debate over evolutionary views in American biology at the time. Donaldson,

Hatai, and King maintained a robust research agenda during those years, dominated by

Donaldson’s interest in separating domestication into its constituent factors. The human and social implications derived from this work grew alongside Greenman’s efforts to expand the reach of the Institute to places where its research could find real world applications.

In fact, the work of the Institute scarcely enjoyed wider relevance than it did late in 1941. In those months, King and Clyde Keeler, who had inherited much of

252 Vaughan, R. Listen to the Music: The Life of Hilary Koprowski. New York: Springer-Verlag, 2000. p.80. 253 Ibid. p.78.

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Donaldson’s data on the rat, reported on an apparent breakthrough in the domestication research program: a radically new interpretation of the domestication phenomenon, spelled out in discrete Mendelian terms rather than inchoate notions of captivity, confinement, and breeders’ preferences. The novelty of King and Keeler’s discovery would seem to place it in stark contrast to prior understandings of domestication. An in- depth look at the fate of their idea, however, reveals the extent to which this scientific explanation relied on earlier aesthetic, practical, and philosophical notions of domestication and suggests the value that experience retained in an era and in a field

(genetics) that increasingly demanded experiment.

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The Color of Temperament: Cultural Exchange and Identity Between Breeders and Geneticists

Throughout his seventy-year career in genetics, Clyde Keeler carried a reputation as a man who saw relations to which others were blind and who followed them vigorously. He also bore the stigma of a dilettante. Following so many leads with such intensity, it seemed, made for poor product in the end. His former graduate advisor,

William E. Castle confided to Keeler, “You might qualify as an artist, a skilled technician in half-a-dozen artizan [sic] employments, cabinetmaker, sculptor, painter, an anthropologist, a geneticist, etc. You are a victim of the multiplicity of your talents.”254

Keeler was also perhaps too willing to lend credulity to ideas that others felt were beyond the pale of scientific research. Castle again, as summarized by Frank B. Hanson of the

Rockefeller Foundation: “A man with an unfortunate personality… [Keeler] lacks critical judgment and needs to be under [an] able leader, then does first rate work.”255 By the end of his career Clyde Keeler was cast as, at best, creative to a fault, at worst, adrift from a scientific culture moored by reason and discipline.

A review of Keeler’s published work and professional correspondence, however, reveals a dogged pursuit of a single line of inquiry. This logically coherent research program, which he nurtured and for which he proselytized for more than 40 years, was predicated on the discovery that a relative few coat-color gene mutations were responsible for the vast complex of morphological, physiological, and behavioral changes

254 Castle, W.E. to Keeler, C.E. December 31, 1954, quoted from Howard, I.K. Clyde Edgar Keeler (1900-1994): Geneticist, Artist, Cultural Historian. Journal of Heredity. 1995 Vol. 86(6):490. 255 Hanson, F.B. Memorandum of Interview with W.E. Castle, Boston, August 7, 1936. RG 1.2, Series 200A, Box 127, Folder 1126. Rockefeller Foundation Archives, Rockefeller Archive Center, Sleepy Hollow, New York (hereafter designated RAC).

120 ascribed to domestication. Few failed to recognize the significance of his findings: the ancient idea of correlated body and behavior was given a strikingly simple and eugenically relevant explanation. Keeler clothed his prize in the best scientific parlance, giving it the status of a general principle known as pigment gene pleiotropy. However, the seed of his discovery was the breeder’s art, domestication, and it was the store of knowledge and material to which he returned time and again in his attempt to solidify his theory. He compiled evidence from every source, sometimes uncritically accepting the wisdom of breeders, of fanciers, and of practically minded men and women who had daily experience with animals and with these phenomena. This attempt at fact gathering and the cavalier way that he relayed anecdotes to colleagues (and occasionally the reading audience) surely damaged his credibility. He pleaded wryly late in his career,

“For this may the gods of science forgive me.”256 But in the adoption of material, data, and even wisdom from breeders, Clyde Keeler and the science of genetics may well have been of the same faith.

It was not uncommon in the first half of the 20th century for geneticists to complain about the baseless ideas to which breeders held, especially the elaborate notions of fancy breeders. Nevertheless, the two cultures kept up a steady trade of animals, money, and advice. The transfer of ideas was an inevitable part of the dialogue. Keeler’s mentor, William Castle, was a staunch adherent to the idea of epistemic purity in science.

He was also engaged in establishing an active trade of material and knowledge with breeders, whose ideas he often regarded as suspect. Castle’s interaction with breeders demonstrates the ambivalence that some geneticists felt toward maintaining too close an

256 Keeler, C.E. Behavior Synthesis Through Pigment Gene Pleiotropy: Reprints and Essays. Atlanta: Publisher Unknown, 1968. p.7

121 association with the traditional arbiters of the knowledge of heredity. It seemed, however, an essential part of his work. Castle’s example is not merely a prelude to

Keeler’s intellectual preparation. Rather, it depicts the sometimes troubled but often necessary interaction between the culture of breeders and that developing around the science of genetics.

Clyde Keeler may have been unique in his candor and likely in the degree to which he adopted ideas and evidence from the culture of animal breeders. His case, however, is illustrative of the exchange, not only of medium, but of wisdom between breeders and geneticists. Keeler’s interpretation of the pleiotropic effects of coat-color genes drew from a deep cultural well concerning the significance of inborn color and held implications regarding its continued social relevance. This account seeks to understand what became of an idea: how Keeler’s social context, professional status, and temperament influenced the development and acceptance of the concept of pigment gene pleiotropy. It demonstrates that these social factors were related to and not distinct from the nature and quality of Keeler’s evidence. In the process, we may also see the interaction of different understandings of domestication and claims to expertise.

Coat-color in History and Culture

British zoologist Arthur Darbishire reported on the results of his experimental crosses between albino and Japanese waltzing mice in 1912: “… it is a fact that, in the case of my mice, reversion to the ancestral condition in regard to disposition is just as invariable a result of the cross as the reversion in regard to colour. The difference

122 between the disposition of the hybrid and that of the albino is as great as that between the colour of the two.”257 Most striking in his account was not only that changes in color and temperament of the animals had similar intensities and frequencies, but that they co- occurred. Darbishire was one of a handful of biologists around the turn of the 20th century, albeit a “scientific commoner” following Georg von Guaita’s example in this case, to lend experimental evidence toward the idea that an animal’s constitution, particularly its color, and temperament could be linked.258

The concept, however, had a long history in medicine and subsequently in the wider Western culture. Under Hippocratic and Galenic medical traditions, elements of physiognomy and behavior could only be explained by reference to a common cause, the balance of humors within the individual. Etymology can be helpful here. Complexion, in its original meaning, referred not to skin color, but to disposition or temperament. Our modern word temperament recalls the Latin term for the mixing of humors. Humoral balance or imbalance, broadly influenced by geography and climate, would determine an individual’s disposition, physical health, and other traits. ‘Sanguine’ and ‘melancholy’ survive in the modern vernacular; ‘choleric’ and ‘phlegmatic’ are archaic, yet they continue to evoke irreducible images of color, health, and character. Distinctions based on skin color date from the Hellenic period, though these generally served to distinguish

257 Darbishire, A.D. Breeding and the Mendelian Discovery. London: Cassell and Company, Ltd., 1912. p.82. 258 Ankeny, R.A. Marveling at the Marvel: The Supposed Conversion of A.D. Darbishire to Mendelism. Journal of the History of Biology. 2000 Vol.33: 315-347. Ankeny identified Darbishire as a “scientific commoner,” one who could clarify the dispute between the Biometric and Mendelian camps by virtue of his relative anonymity in the study of heredity and his ambivalence toward siding with either group. Georg von Guaita earlier reported the on the same cross, with the same general results in coat color and behavior: von Guaita, G. Versuche mit Kreutzungen von Vershiedenen Rassen der Hausmaus. Berichte der Naturforschenden Gesellschaft zu Freiburg. 1898 Bd. X, 3 Heft: 317-332.

123 specific tribes and ethnic groups, not the race as conceived today. Historian Joseph

Graves Jr. establishes the beginnings of modern race theory, laden with concepts of intrinsic inequities and underlain by biological theory, as a social construction between the Age of Discovery and the middle 19th century, not inherent in humor theory.259

As humor theory guided the philosophical antecedents of physiology and psychology for over 1500 years, breeders, particularly from the late 16th century forward, adopted and further developed its ideas to interpret their experiences with plants and animals. Hippology, the study of horses and theories important to their management, had since classical times employed humoral physiology to explain why certain points of an animal’s constitution were telling of its temperament. The Naples school, under the leadership of Frederico Grisone, formalized and popularized many of the concepts of classical hippology in the 16th century. The Englishman, Thomas Blundeville, drew heavily from the Neapolitans in his explanation of personality in horses:

For if he hath more of the Earth than the rest [the other three elements], he is melancholy, heavy, and faint-hearted, and of colour a black, russet, a bright or dark dun. But if he hath more of water, then he is phlegmatic, slow, dull, and apt to lose flesh, and of a colour most commonly milky white. If of the air, then he is sanguine, and therefore pleasant, nimble and of colour is most commonly a bay. And if of the fire, then he is choleric, and therefore light, whote and fiery, a sterer, and seldom of any great strength, and is wont to be of colour bright sorel. But when he doth participate in all the four elements, equally and in due proportion, then he is perfect, and most commonly shall be one of the colours following. That is to say, brown bay, a dapple gray, a black full of silver hairs, a black like a moor or a fair roan, which kinds of horses are most commendable, most temperate, strongest, and of gentlest nature.260

259 See Graves, J.L. Jr. The Emperor’s new Clothes: Biological Theories of Race at the Millenium. New Brunswick, New Jersey: Rutgers University Press, 2002. 260 Blundeville, T. Fower Chiefyst Offices Belongyng to Horsemanship. Seres, London, 1565 quoted from Russell, N. Like Engend’ring Like. Cambridge: Cambridge University Press, 1986. p.70.

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Fancy traits, including coat or plumage color, were central to breeders’ practice as well as their understanding of heredity. Economic traits like girth or milk production were relatively inscrutable, generally continuous, varying in the slightest degrees, and yet prone to apparently capricious reversions or other unpleasant results. Even with the advent of new breeding methods, which gave the breeder greater assurance of the inheritance of economically meaningful characters, fancy traits continued to serve a crucial function in determining which pairing to mate. A 1916 editorial in the Breeder’s

Gazette claimed: “Color is as much a characteristic of a breed as type. The genius of the breeder has set for itself the working out of a problem whereby breeds are identified and distinguished clearly by their color schemes.”261 The meaning of color for breeders, however, was not limited to the identification of more practically valuable traits.

Historian Nicholas Russell has also emphasized the importance of the emotional satisfaction derived from breeders following fancy traits within their stock:

Horn form, coat colour, tail length, colour marks, beauty rather than usefulness of conformation, have always fascinated breeders… This position was reinforced by classical concepts of aesthetics, which implied that what looked beautiful possessed virtue, that external beauty was linked to internal character. The concept could presumably be applied to productive agricultural characters as well as points of personality.262

Harriet Ritvo’s account of the Victorian regard for the white Chillingham and other supposedly aboriginal bands of cattle in Great Britain perhaps sees this identification of color with character and human with animal to its furthest cultural explication yet. The

Chillingham cattle’s particular whiteness became emblematic of their antiquity,

261 Anonymous. Untitled. Breeder’s Gazette May 16, 1916 Vol.69:620. 262 Russell, N. Like Engend’ring Like. p.217.

125 indigeneity, wildness, and Britishness, such that when they occasionally threw a calf of another color or pattern, the misfit young was immediately dispatched by the private owners of the herd, apparently without regard for the irony of having to cultivate a

“natural” state.263 For animals whose domesticity was not so ambiguous, color and other fancy traits continued to play an important role in their breeding, identity, and worth to the end of the Victorian era and beyond.

Domesticated animals were most often classified, if at all, according to their specific utility or, alternatively, by color. Thus, Thomas Fairfax, in his sportsman’s guide, was able to describe red spotted dogs generally as “fiery, and hard to be managed” and yellow dogs as “of a giddy nature, and impatient.”264 Cats, too, were grouped arbitrarily by color. This did not prevent experts from ascribing reliable physiognomies and temperaments to these groupings.265 As the fancy trade around certain, otherwise useless domestic animals developed, pedigree became an increasingly important part of classification, along with breed standards. Coat color was essential to each standard and thus, was a signifier of the purity of the breed. Although the importance of pedigrees and standards reflected existing human social institutions more than innate biological differences, some agricultural scientists believed that these elements of the fancy were finally bringing scientific rigor to the art of breeding. William H. Brewer, Chair of

263 Ritvo, H. Race, Breed, and Myths of Origin: Chillingham Cattle as Ancient Britons. Representations. 1992 No.39:9-10. For further discussion of the Chillingham cattle and other cultural relations of humans and domestic animals see her larger work, The Animal Estate: The British and Other Creatures of the Victorian Age. Cambridge, Massachusetts: Harvard University Press, 1987. 264 Fairfax, T. The New Complete Sportsman or The Town and Country Gentleman’s Recreation. London: Alexander Hogg, 1770. p.125-126. Quoted from Ritvo, H. Pride and Pedigree: The Evolution of the Victorian Dog Fancy. Victorian Studies. 1986 Vol.29(2):238. 265 Huidekoper, R.S. The Cat, or a Guide to Classification and Varieties of Cats and a Short Treatise Upon Their Care, Disease, and Treatment. New York: D. Appleton and Company, 1895. p.64.

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Agriculture at Yale’s Sheffield Scientific School, noted that the benefits of established standards were clear: “the economic production of animals is now placed on a very much surer foundation, excellence is made more uniform, the chances for failure are enormously lessened and the methods of improvement placed on a philosophical basis.”

The praise belonged, not to the practical breeders, but to the fanciers, who “had their associations and set their standards long before the breeders of the more useful farm animals did, and to that Darwin turned his attention... The gain to science has been correspondingly great and numerous unresolved problems in biological science find here their material for use.”266

Of course, there were plenty of critics of the use of standards within the community of breeders and without. Their complaints covered some range, but they objected most frequently to the arbitrary nature of most breed standards and to the seemingly wasteful expense of time and effort in judging the finest points of beauty, which, they claimed, were unrelated to traits of more immediate value. Erwin Hopt, the plain-spoken professor of agriculture at the University of Nebraska, called for greater candor in the 1918 report of the Nebraska Corn Improvers’ Association: “[A] notice… should be displayed in a prominent place above the corn show [reading] something like this: ‘This is all a joke folks. Don’t take the thing too seriously. It should be distinctly understood that this is just a little game we boys have gotten up among ourselves. It has no relation to utility.’”267 He could have delivered the same message to animal breeders as easily. Biologists were often critical of the “notions” that breeders employed in

266 Brewer, W.H. The Mutual Relations of Science and Stock Breeding. Science. 1893 Vol.22(556):170. 267 Quoted from Multiple unidentified authors, Livestock Breeding at the Crossroads: A Foreword on the General Results of the Survey of Germ Plasm in Beef Cattle, Dual-purpose Cattle, Sheep, Swine, and Horses in Yearbook of Agriculture. Washington: U.S. Department of Agriculture, 1936. p.834.

127 preparing their crosses. “The breeder,” wrote Walker van Riper, a naturalist in the employ of the Research Laboratory at the University of Denver, “never fails to fall into the error of correlating the outward and visible signs with inward and spiritual grace…

The tyrannous power of these notions… are doubtless of the same nature as style and fashion. They have the same aesthetic basis, are equally irrational, may either promote or oppose welfare, may be irrelevant and foolish, and they operate with the greatest authority.”268 The standards, van Riper and other scientists alleged, were merely formalized aesthetic notions, simultaneously arbitrary and doctrinaire. It is not entirely clear, however, where these critics believed the authority lay. They blamed the thought collective of breeders’ associations to be sure, but the aesthetic notions themselves appeared to have a special power over breeders: “[Man] continually demonstrates in this field how incorrigibly subject he is to his aesthetic ideas.”269

The dissemination and elaboration of Mendel’s rules of heredity after 1900 gave many scientists confidence that heredity was or would soon be set upon firmer principles than those derived from breeders and fanciers. While interaction between breeders and the new geneticists was no great secret, geneticists had an interest in maintaining a measure of intellectual distance from their subordinate partners in the study of heredity.

Geneticists who later reflected on the early history of their discipline often marginalized or wholly neglected the role of breeders. The flow of useful information ostensibly proceeded in one direction, with geneticists the source and breeders the sink.

268 van Riper, W. Aesthetic Notions in Animal Breeding. The Quarterly Review of Biology. 1932 Vol.7(1):85-87. 269 Ibid. p.88.

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Historian of science Charles Rosenberg revised this history to a degree: “Mendel added nothing to the technical armamentarium of the skilled empirical breeder. As geneticist Raymond Pearl noted sharply before the First World War, all that the new genetics had really done thus far was to help the breeder interpret his accustomed techniques.”270 Perhaps this discounts the contributions of geneticists too much. The exchange between breeders and geneticists fostered change in techniques and philosophies but also permitted the maintenance of existing concepts as they were deemed still valuable. This selective, and sometimes unconscious, uptake of ideas seems to have varied according to the goals of the person carrying out the breeding and his or her experience with the animals to be bred (or with domesticated animals in general).

Reflecting this discernment, the Scottish fancier W. Mackintosh Kerr noted that “[in] the

Mouse Fancy… young and old seem eager and anxious to know more about the inheritance of colour. The newer outlook does not mean the scrapping of older methods but their modification. All the theoretical knowledge in the world will never make one a really successful fancier unless one trains one’s eye to detail.”271 Color was the prize of most fanciers, and they were eager to learn more definite rules for its inheritance. They were unwilling, however, to abandon tried and true practices that determined not only the outcome of their crosses, but also the culture of the fancy. They sought to incorporate the

270 Rosenberg, C.E. No Other Gods. p.167. For an expanded analysis of the relation of practical breeding and genetics, using the example of Raymond Pearl, see Cooke, K.J. From Science to Practice. 62-86. Using an institutional approach, see Kimmelman, B.A. A Progressive Era Discipline. 271 Kerr, W.M. Colour Inheritance in Fancy Mice. Idle, Bradford, England: Watmoughs Limited, n.d. p.6. Barbara Kimmelman also notes such discernment among breeders in the American Breeders’ Association as they readily took up Mendel’s postulates and generally neglected De Vries’ Mutationstheorie because it did not fit with their experience with sporting varieties. Kimmelman, B.A. The American Breeders’ Association. p.204

129 ideas of the new genetic sages into their existing frames of reference. In a similar way, scientific students of heredity did not approach their subject with a tabula rasa.

Familiarity nurtured the desire to establish an epistemic distinction between geneticists and breeders, especially fanciers. Many early geneticists had come from agricultural backgrounds, unsurprising in the late 19th century of their childhoods, but it meant that they entered their roles as geneticists with extensive experience observing and caring for domesticated animals and plants. From careers as experimental zoologists, morphologists, and embryologists, many of the new geneticists changed their experimental material from a more heterogeneous mix of domesticated and wild-sourced animals to more exclusively domesticated varieties prior to 1910. Aside from the perceived applicability of experimental findings, domesticated animals offered a number of benefits to the student of heredity: tractability due to tame behavior, the recognized plasticity of domestic varieties, co-occurrence of various traits, ready supply from an established body of breeders and fanciers, and the immediate availability of already distinct breeds. Although large livestock, by virtue of their economic importance, remained important subjects of study, scientists seeking general hereditary rules turned their attention to faster breeding and more compact animals. This, in turn, drew their attention to fanciers who, for their own interests in heredity and novel traits, had developed dozens or even hundreds of varieties of small mammals and poultry from which geneticists could select. It was not mere coincidence that geneticists found such material readily at hand. They shared a number of interests and goals with fanciers, including preserving and “extracting” interesting variations, or “sports” in the shared lexicon.

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Color was a particularly valuable currency in their exchange. It was one of the most easily noticeable traits of an animal. It also seemed to be one of the simplest and most reliably inherited traits of an animal, hence its use as an earnest for latent, more complex, or slowly developing traits among breeders. Geneticists and other scientists interested in heredity often employed color as a marker of hidden traits. John W. Gowen, a student of Raymond Pearl’s at the Maine Agricultural Experiment Station, argued that,

“the intimate association which exists between the hereditary units necessitates the studying [of] such things as coat color and tongue color for the full analyses of the economically important problems.”272 J.A. Weir, geneticist at the University of Kansas, noted as late as 1954 that “There is a prevalent view, not without evidence, that color and temperament are associated,” though he further qualified, “study of morphology is important but offers no shortcut of success to the breeder.”273 Even scientific heavyweights such as Charles Otis Whitman believed that color serves as a “tell” for other traits that were more complex, more ambiguous, or ill defined. “Weakness in common pigeons is frequently expressed in white color… In a pair of young pouters from gray (white-barred Brünn) pouters, one… is gray with dark bars and in form is apparently well developed; the other… is white with poorly developed wings, the primaries being imperfectly developed. This bird has the ‘shakes’; that is, it trembles all over and can not control the movements of the head. (I have had two such birds from a dealer, and they are

272 Gowen, J.W. Studies in inheritance of Certain Characters of Crosses between Dairy and Beef Breeds of Cattle. Journal of Agricultural Research. 1918 Vol.15(1):4. 273 Weir, J.A. On the Concept of Inherited Constitution. Transactions of the Kansas Academy of Science. 1954. Vol.57(2):118.

131 evidently of the same nature).”274 Whitman added red coloration and dilution of barring pattern as signatures of “weakness” as well. Having no clear definition of weakness, nor any certain analysis beyond anecdotal observation, Whitman’s correlation of color and weakness was qualitatively identical to the relations that many breeders and fanciers maintained as guides to their own practice. Charles Darwin reported famously on the co- occurrence of white coat, blue eyes, and deafness in cats in his treatise on the effects of domestication.275 The report was made on the authority of men familiar with the breeding of such cats.

Suggestive of the breadth and staying power of these notions, in 1945 the

Argentine doctor of medical science, Almanzar Marrero y Galindez, described a similar phenomenon in his Cromohipologia [sic.], or the science of color inheritance in horses.

Not unlike classical hippology, Galindez’s Cromohipologia placed great importance on the coat color of the horse for illuminating much more about its general constitution.

Each color variant was arranged in a hierarchy according to the number of

“degenerations” it had accumulated. “Every degeneration is immediately translated into a diminution of chromogenes, which makes it and all its descendants vulnerable, and on its body it carries its constitutional tally and the coats are a page in nature’s book.”276

Galindez placed the origin of the science squarely in the experienced hands of “the

274 Whitman, C.O. Posthumous Works of Charles Otis Whitman, Volume II, Inheritance, Fertility, and the Dominance of Sex and Color in Hybrids of Wild Species of Pigeons. Oscar Riddle (ed.) Washington, D.C.: Carnegie Institution of Washington, 1919. p.184-185. 275 Darwin, C.R. The Variation of Animals and Plants Under Domestication. Vol. 2 London. p.329. 276 “Cromohipologia” in William E. Castle Papers, Ms.Coll.14, Folder “Deering, Davis- Cromohipologia #1,” APS. For an explanation of the relation of domestication study to degeneration theory, see Section III this volume and Burkhardt, Richard W. Jr. Patterns of Behavior.

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Gaucho, who, after his own manner, knew anatomy of horses, and [due] to his profound observation, it is possible to individualize one horse among thousands.”277

It should be stated that the scientific study of heredity was not equivalent with the art of breeders and fanciers. Biometricians, geneticists, cytologists, embryologists, and experimental zoologists all had distinct intellectual heritages, distinct to varying degrees from one another and from the breeders’ world. Geneticists, especially, had reason to differentiate their activities from those of breeders. Theirs was a new science set upon solving an old problem, one that had been dominated by those with practical experience, ancient traditions, and a store of accumulated wisdom. They were in the process of establishing their own research agendas, standards of evidence, and foundational theory, consciously different from what breeders were using in practice. Breeders did not codify the rules of color inheritance, despite its perceived importance to their work. Fanciers could predict with some accuracy the result of a cross between two individual animals of known pedigree, but this reliability fell sharply as they moved to generalize their predictions. Color inheritance instead became one of the great victories for Mendelism in the first decade of the 20th century. “A breeder who was an expert on rabbits, guinea pigs, rats, or mice before 1900,” remarked historian of science William Provine, “could only marvel at the giant steps taken toward the understanding of color inheritance in only nine years.”278

These steps had been taken most notably by William E. Castle and his students at

Harvard’s Bussey Institute. The culture that prevailed at the Bussey encouraged the

277 Ibid. 278 Provine, W.B. Sewall Wright and Evolutionary Biology. Chicago: University of Chicago Press, 1986. p.57.

133 rigorous investigation of inheritance by controlled experiment and its explanation in mechanistic, if speculative, terms. This culture also encouraged mimicking many practices of breeders and fanciers, seeking out their unique varieties and, sometimes, advice. The cultures of geneticists and breeders were distinct, but they were not wholly independent. They were familiar trading partners, especially in the early years of the 20th century.

William E. Castle and the Bussey Institution: The Culture of Coat Color Genetics

William E. Castle held an abiding commitment to the idea of a “pure” science of genetics and another to its practical application in the field of agriculture. These aims were not by any means inimical to one another. They did, however, require the interaction of two cultures sharing a common interest in the rules of heredity: one whose claims to knowledge in heredity were borne out in their ability to improve the average condition of their animals over many centuries; the other claimed to have general and reliable rules of heredity founded on basic physiological truths rather than experience.

Castle’s philosophy regarding heredity always hewed close to the latter epistemic basis.

His training under Charles Davenport and E.L. Mark at Harvard had imbued him with a strict materialist interpretation of living nature. His practice, however, in carrying out his work in genetics, encouraging students to follow independent but fruitful paths, and applying his knowledge to practical concerns brought him into close association with breeders, their methods, and their notions. The place of both cultures at Castle’s

134 workbench may be illustrated by the peculiar nature of the Institution where he and his students completed the work on the inheritance of coat color and other traits.

Harvard’s Bussey Institution in the Forest Hills section of Jamaica Plain, eight miles from Cambridge, had an earlier life as a private farm for its namesake Benjamin

Bussey. Bussey, the man, bequeathed his estate into the care of Harvard University in

1835 with the purpose that it should provide “instruction in practical agriculture, in useful and ornamental gardening, in botany, and in such other branches of natural science, as may tend to promote a knowledge of practical agriculture.”279 Instruction did not begin at the Bussey Institution until 1871, once the money became available and a building was constructed. The facilities, however, never fully reached Mr. Bussey’s vision of undergraduate instruction in practical agriculture. The Institution relied on funds from rental income from other properties that Bussey had willed to Harvard. Massive fires in the fall of 1872 in Boston, however, left the Institution with meager funds. Its operations became increasingly practical and less instructive. Vegetables from the grounds were sent to Harvard’s dining hall, undergraduates split wood and sold it to locals, and privately owned horses occupied many of the outbuildings, providing income for the students and the ailing organization. By 1908, the situation at the Bussey had become sufficiently embarrassing for its negligent caretaker across the Charles River that the school was to be closed.

William Castle, who had been called back to Harvard to serve as a professor of zoology in 1897, campaigned to have the Bussey Institution reorganized into an advanced center for graduate training in the biological sciences, with an eye toward the practical

279 Quoted from Weir, J.A. Harvard, Agriculture, and the Bussey Institution. Genetics. 1994 Vol.36:1227

135 agricultural application of its researches. The effort to reform the Bussey came as part of physicist Wallace Clement Sabine’s conversion of the undergraduate Lawrence Scientific

School into the Graduate School of Applied Science, of which he was dean. Sabine used

Castle’s growing reputation in the field of genetics as leverage in talks with Harvard

President Charles Elliot: “The University has Professor Castle to start with; to lose him will be to lose the best man in the country in genetics.”280 The Bussey Institution was repurposed as a graduate training institute, and Castle was retained along with his substantial external funding from the Carnegie Institution of Washington.

Research in the reformed Bussey Institution was varied, not only between the divisions of animal and plant genetics headed by Castle and Edward M. East, respectively, and entomology headed by William Morton Wheeler, but even within disciplines. Castle encouraged his students to pursue their own research interests and to find solutions to problems independently. “There was a stamp of independence on nearly everybody who went through there. Nobody had to wear any yoke. If you disagreed… you said so, and you got a good tart reply. You see, everybody virtually looked out for himself,” relayed Bussey alumnus L. C. Dunn.281

Although Bussey students were given ample freedoms, Castle was sure to impart his mechanistic approach to explaining gene action and inheritance to his pupils. In order to better understand the physiology of coat color, a signature issue at the Bussey, Castle and his students regularly read and discussed the papers of pigment biochemists such as

280 Ibid. 281 Quoted from Rader, K.A. “The Mouse People”: Murine Genetics Work at the Bussey Institution, 1909-1936. Journal of the History of Biology. 1998 Vol.31(3):342. Rader notes that independence extended not only to Bussey students, but to the Institution itself, given its distance from Cambridge and the Zoology Department. This status helped to breed resentment at the Harvard campus.

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H. Onslow and Oscar Riddle.282 Students were also united by experience on the practical side. Many of Castle’s students were given charge of an animal or a particular strain of animals during their tenure at the Bussey. Castle trained the majority of his animal caretakers personally, his knowledge of the animals kept keen by regular turnover among graduate students. Stewardship of the mouse colony, for example, passed from C. C.

Little to Dunn to Horace Feldman to Keeler and William Gates to George Snell, with

Castle present to manage the transitions. Bussey students bred animals and noted the rare occurrence of “sports” with interest. They also attended fanciers’ shows, not only as spectators but also as participants.283 Immersing themselves deep within the culture of breeders seemed to facilitate the exchange of animals and advice. Castle saw value in this interaction, for he frequented many breeders’ and fanciers’ shows himself. He wrote to Dunn early in 1922: “I have been to the dog-show today with my ‘seminar class’ (zool

10) a nice bunch of some ten men, all of them but two working at the Bussey. It was very instructive. We had just been going over Darwin’s chapter on dogs and providentially this show came on. I jumped at the chance to give them something concrete and first hand on the subject. Over a thousand dogs benched, most of the recognized breeds represented.”284 Suggesting Castle’s reverence for the wisdom of practical experts,

Keeler remembered Castle telling him of the Bussey’s longtime caretaker David Patch

282 Provine, W.B. Sewall Wright and Evolutionary Biology. p.91. 283 Keeler, C.E. How it Began. p.185. in Morse, H.C. (Ed.) Origins of Inbred Mice. New York: Academic Press, 1978. Karen Rader provides an excellent account of the generally fruitful, but sometimes tenuous, relationships between Bussey “inmates” and mouse fanciers, especially through the example of C.C. Little. See: Rader, K.A. Making Mice. p.31-41. 284 Castle to Dunn February 21, 1922. L.C. Dunn Papers B.D917, Folder “Castle, William E.,” Box 15, Series I, APS. Also see: Castle to Dunn January 24, 1922. Ibid.; Castle to George Rommel November 22, 1916. Sewall Wright Papers Ms. Coll.60, Folder “Castle, W.E.,” Box 5, Series I, APS; and his correspondence with various horse and pony breeders in William E. Castle Papers, Ms.Coll.14

137 that, “if their circumstances had been exchanged, probably Patch would be a better professor and Castle a poorer caretaker.”285

Castle’s appreciation for breeders and their skill not withstanding, he held publicly and privately that his science remained objective and pure. His early background in experimental zoology (specifically embryology), his materialist, mechanist philosophy, and place among the first generation of geneticists trying to claim the mantle of leadership in studies of heredity contributed to his drive to distinguish his knowledge of inheritance from that of breeders. Castle maintained an interest in inheritance prior to the rediscovery of Mendel’s 1866 paper on hybrid peas, likely the influence of his research mentor Charles Benedict Davenport.286 It appears to have been William Bateson and not the triumvirate of Mendel’s rediscoverers who inspired Castle to begin experimenting with heredity.287 By 1902 Bateson was already entrenched in favor of the interpretation that characters varied in a discontinuous fashion, in opposition to the continuously variable traits observed by Karl Pearson’s biometric school. He declared,

“[T]here is one class of notable exceptions [to continuous variation], those in which colour is the character considered, whether the coats of horses and dogs, or the human iris.”288 Color inheritance, evidently discrete and discontinuous, gained a special importance for early Mendelians in their epistemic dispute with the biometricians.

285 Keeler, C.E. How it Began. p.182 Quoted from Rader, K.A. “The Mouse People” p.343. 286 Castle expressed interest in experimenting with a family of polydactylous cats that he and Harvard colleague Herbert Neal had discovered on the roadside in Castle to Davenport August 26, 1894. Charles B. Davenport Papers B.D27 Folder “Castle, William E.- 1,” Series I, APS. 287 Provine made this determination and the parallel of Castle’s early work and interpretation in genetics would seem to bear this out. Provine, W.B. Sewall Wright and Evolutionary Biology. p.37-38. 288 Bateson, W. Variation and Differentiation in Parts and Brethren. In Punnett, R.C. (ed.) The Scientific Papers of William Bateson. Vol.I Cambridge, Cambridge University Press, 1928. p.444. Emphasis original.

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Fulfilling Bateson’s prophecy and perhaps justifying the expense of reorganizing the Bussey Institution, Castle scored one of the great early victories for the school of discontinuous variation in 1908, by this time roughly associated with the legacy of

Mendel. His short paper in Science, “A New Color Variety of the Guinea-pig,” did not merely describe a new “cinnamon-agouti” sport arisen within a captive breeding population. It, rather, predicted its appearance despite the fact that such a strain was utterly unknown to guinea pig fanciers. Castle, who had known of cinnamon-agouti mice, made his prediction based on what he believed were fixed, broadly applicable, and distinctly Mendelian rules of color inheritance. His discussion at the end of the paper seemed to simultaneously assume leadership in the study of color heredity for the

Mendelians and to free geneticists, breeders, and fanciers from the superstitions and facile explanations about the inscrutable power of domestication to produce endless variation:

A moment’s study of this case shows what a really great advance in the theory and practice of breeding has been obtained through the discovery of Mendel’s law. What a puzzle this case would have presented to the biologist ten years ago! Agouti crossed with chocolate gives in the second filial generation (not in the first) four varieties, viz., agouti, chocolate, black, and cinnamon. We could only have shaken our heads and looked wise (or skeptical).

Then we had no other explanation for such occurrences other than “the instability of color characters under domestication,” the “effects of inbreeding,” “maternal impressions.” Serious consideration would have been given to the proximity of cages containing both black and cinnamon- agouti mice.

Now we have a simple, rational explanation, which anyone can put to the test. We are able to predict the production of new varieties and to produce them. 289

289 Castle, W.E. A New Color Variety of the Guinea-pig. Science. 1908 Vo.28:251-252.

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Castle’s reflection on the ideas held by breeders and shared by earlier naturalists suggested their absurdity in comparison with the new Mendelian principles. The orthodoxy was foolishness. Indeed, he had a more generally applicable program defined by unit characters, and if he was successful in determining their physiology he would be able to explain why, and not simply how, they appeared in new generations. His work, including that on coat color, was directed not toward finding novel color varieties, but toward clarifying some point of greater significance to heredity, variation, and ultimately evolution.

Castle’s 1908 report exemplified his commitment to a mechanist philosophy and to the concept of science unadulterated by unqualified “notions.” Despite his regular contact with breeders, he was always ready to defend the purity of his science from their influence. Castle was instrumental in reorganizing the American Breeder’s Association from “an original mixed assemblage of scientists interested in the study of heredity, agricultural experts guided by scientific knowledge, and rule of thumb horticulturalists, and animal breeders governed by old wives’ tales,” to a strictly scientific body by

“[eliminating] the third unprofitable constituent.”290 While he was accepting of agriculturalists (his examples were limited to university trained men, such as Willet Hays and Edward M. East), he vigilantly defended the science of heredity from uncritical thinking. To Charles Davenport, with whom he was engaged in an attempt to review the best books on domestication, Castle gave his impressions of a volume on captive bred birds: “I am somewhat disappointed in it, but I suppose I expected too much. These

290 Castle, W.E. The Beginnings of Mendelism in America. In Dunn, L.C. (ed.) Genetics in the 20th Century: Essays on the Progress of Genetics During its First 50 Years. New York, MacMillan Company, 1951. p.60.

140 practical breeders are not primarily men of science, and they get a good many notions mixed in with their science.”291 Breeders could, however, stay in Castle’s good graces by adopting the genetic faith and loosing their tradition-bound ideas, as in the case of E. C.

Richardson, “an English fancier who is something of a geneticist- a pupil of Punnett’s.

He has tried to do missionary work by informing the English fanciers about genetic principles.”292

From Castle’s constant interaction with breeders and his public and private comments about their “notions,” his prejudice seems to fall in opposition to the culture of breeders, their collected methods, traditions, and wisdom, rather than the practitioners themselves. This inclination is predicated on his vision of a pure science. His distaste for opinion or speculation is apparent in a letter to Davenport, near the end of his long defense of the idea that heritable factors could be modified themselves by selection: “Of course you don’t expect one, in matters scientific to accept any views except as I am convinced of the soundness of the evidence in which it rests. One can’t trust even his own views, much less another’s. We want incontrovertible facts, no matter whose views they favor… Indeed I have been getting used to scientific solitude. Not that I like it, quite the contrary, but a scientist should seek the truth first of all.”293 Evidence and not influence, Castle felt, determined his interpretation of nature.

Castle’s vision of a pure science was easier to maintain in theory than in practice.

He was clearly aware of breeders’ ideas, whether he received them appreciatively or

291 Castle to Davenport February 26, 1904. Davenport Papers, Folder “Castle, William E.- 1,” Series I, APS. Emphasis original. 292 Castle to Sewall Wright March 4, 1935. Wright Papers, Folder “Castle, W.E.,” Box 5, Series I, APS. 293 Castle to Davenport March 19, 1918. Davenport Papers, Folder “Castle, William E.- 10,” Series I, APS. Emphasis original.

141 disdainfully. His reputation and his science were, in part, built upon showing that the understanding of heredity among breeders was unsatisfyingly vague. Signatures of the breeders’ culture, though, found their way into Castle’s science. In a 1940 discussion of the genetics of coat color in mammals, Castle evoked both the esoteric and practical regard that breeders had long maintained about color and its relations to other, more latent traits:

When only one or two or even three gene differences are involved [in a cross], the respective gene combinations of two breeds are readily interchangeable, but such exchanges are rarely made, and in general are not advisable, since the distinctive coloration of a breed constitutes a kind of trademark guaranteeing the integrity of the assemblage of less obvious genes… When a color mutation is dominant in character, as the white face of Hereford cattle, it has a great trademark value, since it gives the same general appearance as purebreds.294

Castle had long maintained an interest in gene and character interaction and nurtured this idea among others. Robert M. Yerkes credited Castle with giving him the idea to study “the heritability of savageness and wildness in certain strains of rats which were being bred for studies in the heredity of structural characteristics at the Bussey

Institute” in 1910. Yerkes published on the subject in 1913, claiming to have “[proven] conclusively that savageness, wildness, and timidity are heritable behavior complexes.”

The greatest changes in behavior were seen between wild-caught rats and their progeny from crosses with ostensibly tame hooded rats. However, the extent of hooding in the tame rats was said to correlate with emotionality as well. Yerkes claimed impartiality in his psychological assessments, noting that Castle and his student John Phillips had bred

294 Castle, W.E. Mammalian Genetics. Cambridge, Massachusetts: Harvard University Press, 1940. p.143.

142 and selected the individuals to be tested for him.295 Thus, Castle had chosen different coat color varieties of rats for an evaluation of the inheritance of their emotionality.

Given that this was a preliminary report (the full length report never surfaced) and that

Yerkes was not involved with the selection of the rats or developing the original idea, it is difficult to determine Castle’s motivations for this study. The selection of coat color varieties for emotional testing, though, at least paralleled the notion popular among breeders that color was a key signal of such complex traits as temperament.

In 1905 William Castle himself described the relation between the scientist and breeder: “The aims of the biologist are so different from those of the practical breeder that to solve the theoretical problems in the formation of breeds the biologist must himself turn breeder, and see new organic forms arise out of material with which he is thoroughly familiar, and under conditions that he can control.”296 The differences between them required the highest level of familiarity, imitation. Thus, in training Little,

Wright, Dunn, Keeler, and others, Castle insisted on an intimate knowledge of the living material, the knowledge gained by husbandry. He was known to engage in even the most tedious aspects of breeding himself and made constant reference to domestication and domesticated animals in his published works.

Domestication seemed a risky subject for the sober scientist. It was regarded as inscrutable, likely a complex of interrelated phenomena, not one of which was well understood. Earlier attempts to explain it largely belonged to the cultural wisdom of

295 Yerkes, R.M. The Heredity of Savageness and Wildness in Rats. Journal of Animal Behavior. 1913 Vol.3(4):286-288 and 293-296. Yerkes seems to attribute all differences in emotionality to the amount of “wild blood” in the stocks, this term is not sufficiently explained, nor is the association of wideness of hood with wild blood. 296 Castle, W.E. The Mutation Theory of Organic Evolution, from the Standpoint of Animal Breeding. Science. 1905 Vol.21(536):522.

143 breeders, often referred to derisively as superstitions or old wives’ tales. Castle felt, however, that domestication offered the best opportunity to reveal the secrets of organic evolution. Invoking the ghost of Darwin, he declared: “[T]here is no essential difference between breeds and species, and if we can ascertain how breeds originate we can infer much as to the origin of species.”297 The discerning and objective scientist could perhaps minimize the influence of unfounded “notions”.

The subject of domestication became even more prevalent in Castle’s work after

Harvard unceremoniously closed the Bussey Institution in 1936. Castle found work, at the age of 70, as a research associate in mammalian genetics at the University of

California, Berkeley. There, certainly in the twilight of his career, his collaborators were as often breeders as they were geneticists. He coauthored papers with breeders, including

Larry Moore who operated a domestic mink farm in northeastern Wisconsin.298 He also maintained an extensive correspondence with horse and pony breeders in his later years.

The economy between Castle and breeders may be exemplified by an exchange with

Wayne A. Munn of the American Shetland Pony Club. Castle addressed Munn: “About

[ponies] I am told you ‘know a lot’ and am hoping you will be willing to share your knowledge with students of genetics… enlighten me with your wealth of personal observations.” Critically, it was not material but knowledge Castle sought, knowledge brought by experience. In exchange for information, Castle provided Munn with “genetic recipes” for his ponies.299

297 Ibid. 298 Castle, W.E. and Moore, L. Mutations in Mink under Domestication. Journal of Heredity. 1946 Vol.37(5):137-145. 299 Castle to Munn December 7, 1951 and December 2, 1952. Castle Papers, Folder “American Shetland Pony Club,” APS. Davis Deering, a Texas horse breeder, sent Castle a translated partial copy

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Castle shared his interest in domestication not only with breeders, but also with researchers at the Wistar Institute in Philadelphia. Despite his initial skepticism about the direction of rat research at the Wistar, Castle ultimately contributed to their domestication program by providing wild cavies and domestic guinea pigs for their comparison to gray and albino Norway rats, respectively.300 With Helen Dean King, he coauthored ten papers on linkage phenomena among mutant rat lines primarily derived from the Wistar

Institute’s rat domestication project. These mutant lines were distinguished by coat color and texture, the heredity of which Castle was unquestionably expert in explaining. He wrote a review of domestication-related phenomena in the rat in 1947, using the results of physiological and genetic work on domesticated rats to suggest their probable origins.301

William Castle was quite successful in his efforts to establish a culture of genetics around his mechanist, materialist philosophy of science. His landmark studies of coat color demonstrated not only the potential outcomes of specific matings, but helped to codify rules of color inheritance and of heredity more generally. He sought the physiological underpinnings of such color phenomena and encouraged the same pursuit among his students. All the while, he remained skeptical of reports that did not satisfy his criteria of mechanistic explanation. He heaped criticism on the naive ideas of breeders, despite his appreciation for their expertise and skill. Nevertheless, Castle found

of Galindez’s Cromohipologia in 1951. We are not given the pleasure of Castle’s reply. Deering to Castle November 27, 1951. Castle Papers, Folder “Deering, Davis- Cromohipologia #1,” APS. 300 Henry H. Donaldson wrote in his diary December 31, 1909: “Forenoon- To Castle’s Laboratory at Forest Hills- he misses our problem.” Henry H. Donaldson Papers and Diaries B.D713M, APS. Castle’s participation in the Wistar domestication program is mentioned in Donaldson, H.H. On the Cranial Capacity of the Guinea-Pig – Wild and Domesticated. Archives Suisses de Neurologie et de Psychiatrie. 1923. Vol.13:206. 301 Castle, W.E. The Domestication of the Rat. Proceedings of the National Academy of Sciences of the United States of America. 1947. Vol.33(5):109-117.

145 that the work of genetics in its infancy was nearly impossible without the material and epistemic aid of breeders and fanciers. He participated in an active exchange with these common sense experts throughout his career and found value in mimicking their practices in his scientific work and pedagogy. Practical experience developed the skills and discernment necessary to become a successful breeder or geneticist. Rare instances in

Castle’s published work and correspondence suggest that he may have seen value in some of the ideas popular among breeders, even before he had accomplished experiments to verify the accounts. From Castle we may see that the culture of geneticists was distinct from that of breeders but most intimately engaged with its practices, products, and knowledge.

Domestication and the Genesis of Pigment Gene Pleiotropy

By the time William Castle began his collaborative work with Helen Dean King on linkage in mutants from a line of domesticated Norway rats, his student Clyde Keeler had begun a career in medical genetics, a field whose aims seemed distantly related to the breeding intensive work that prevailed in the Bussey Institution. Since his days as a graduate student at the Bussey, Keeler seemed keen to demonstrate the importance of his work for humans, particularly in medicine. His former employer at the Eugenics Record

Office, Charles Davenport, had directed Keeler to Castle and to research work when

Keeler realized he would not have enough money to attend medical school.302 Keeler had professed an interest in natural science, but most of his work at the Bussey was directed

302 Keeler to Davenport April 15, 1923 and Davenport to Keeler April 18, 1923. Davenport Papers B.D27.2, Folder “Keeler, Clyde E. #2,” Series IIB, APS.

146 toward medical subjects. Indeed Keeler later offered the rationale for his pursuit of research above medicine: “If I went into medicine, I might save the lives of two hundred or more persons, but if I went into research and was lucky enough to make a biomedical discovery of lasting value, I would affect the knowledge of many physicians and through them I might help to save the lives of thousands as yet unborn. So I went into my career with my eyes open.”303 Keeler’s “discovery of lasting value,” though, came not from medicine but from turning his attention to two characteristically “Bussey” subjects, domestication and coat color inheritance.

His concept that coat color genes had multiple and related, or pleiotropic, effects on behavior, gross morphology, and physiology originated as an attempt to explain the

Gordian knot of domestication. He took to the problem with zeal and, like Castle, tried to establish its legitimacy on physiological grounds, using multiple animals as models.

Unlike Castle, he openly took up many other forms of evidence, including some baldly anecdotal or derived from broad cultural wisdom. Keeler’s concept of pigment gene pleiotropy and the social implications that sprang from it were informed by existing cultural ideas about the relevance of external color for explaining inward character.

Clyde Keeler- Education and Early Career

Before matriculating to Harvard for graduate work in 1923, Keeler had taken his bachelor’s and master’s degrees from Denison University in Granville, Ohio, where

303 Keeler, C.E. The Gene Hunter: Excerpts from a Life in Bio-science. Milledgeville, Georgia: Philip Giles, 1984. Quoted from Howard, I.K. Clyde Edgar Keeler (1900-1994). p.489.

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William Castle had been an undergraduate years before.304 The son of a watchmaker and educator, Keeler did not grow up on a farm like his future mentor. His autobiography, however, recounted his early love of nature and fond memories of visits to the farm of a childhood friend and the surrounding woods.305 His interest in nature, especially living things, combined with the examples set by his great uncle, uncle, and cousin, all physicians, to convince Keeler to pursue medicine. In 1920, while still a student at

Denison, he began working for the Eugenics Record Office at Cold Spring Harbor under the direction of Charles Davenport. Perhaps the promise of social betterment, explicit in the agency’s goals, appealed to him as it did many prospective physicians. Expecting much of Davenport’s memory two years later, Keeler wrote to him, “You will probably remember me as the chap who helped Dr. [George W.] Corner by going to the city slaughter house for embryos and to inspect uteri and corpora lutea of the hog.” 306 Some favorable impression must have remained, for Keeler was invited back to Cold Spring

Harbor as a member of the eugenics training corps for the next three summers. In the spring of 1923, as it became clear that Keeler would not be able to afford a medical education, Davenport suggested Castle and the Bussey Institution. He also provided the letter of introduction:

I want to call your special attention to Mr. Clyde Keeler who is coming to work with you as a graduate student this autumn. Mr. Clyde Keeler has been with us during the summers for about three years. He is a graduate

304 Keeler was actually awarded his Master’s of Science from Denison in 1925 after completing his thesis, “Does Unilateral Extrication of the Eye Affect Cranial Symmetry in Mus musculus?” at the Bussey Institute. He had, though, finished all coursework for the degree by 1923. See Keeler, C.E. The Gene Hunter. p.57. 305 Ibid. p.12. 306 Keeler to Davenport June 9, 1922. Davenport Papers B.D27.2, Folder “Keeler, Clyde E. #2,” Series IIB, APS.

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of your Alma Mater, Denison University. Clyde Keeler is very active physically and mentally. He has a gift for drawing which he hopes to use to help him thru [sic.] college. It is possible that he may be a little too versatile for an ideal scientific career. I believe that if he sticks to one topic at a time, as a good scholar should, that he has a career before him. I have talked with him a good deal about you and he is looking forward to his new work with a great deal of enthusiasm.307

Keeler’s reputation as unfocused, thus, preceded him. A parcel he brought with him to Cambridge would help him establish a standing as a keen observer. He was given a pair of discarded albino mice from his former colleague, Emelia Vicari, at Cold Spring

Harbor as he visited along his route to Massachusetts. Samuel Detwiler of Harvard’s zoology department assigned Keeler the task of preparing sections of the eyes of rare birds that Detwiler had retrieved from China. Keeler practiced his skill on progeny from his Cold Spring Harbor mice and discovered that the retina lacked rods, one of the key photosensitive structures of the eye. Detwiler blamed Keeler’s “poor technique for losing half the retina.”308 Later analysis demonstrated that Keeler was correct and not merely an inept technician; the strain of mice he had been given were mutants, completely devoid of rods. He later identified this condition as the result of a single recessive mutation. His work on “rodless retina” mice also gave him a brief introduction to the field of comparative psychology when he developed maze tests with the aid of Castle’s former student and colleague, Robert Yerkes.309

307 Davenport to Castle September 4, 1923. Davenport Papers B.D27, Folder “Castle, William E.,” Series I, APS. 308 Pittler, S.J., Howard, I.K., and Sidman, R.L In Memoriam: Clyde E. Keeler (1900-1994). Experimental Eye Research. 1995 Vo.60:3 309 Keeler, C.E. Rodless Retina, an Opthamalic Mutation in the House Mouse, Mus musculus. Journal of Experimental Zoology. 1927 Vol.46:355-407.

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At this early point in his graduate career, Keeler was still ambivalent about joining Castle at the provincial Bussey Institution. “It is possible I shall make up my mind to remain at the [Museum of Comparative Zoology] and take what work I see fit at

Bussey,” Keeler confided to Davenport late in his first semester at Harvard.310 Keeler was concerned not only by the relative few courses offered at the Bussey but also that the work ongoing there might not help him reach his goals of human benefit. He learned, however, upon greater contact with members of the institution, that many Bussey men had humankind in mind even as they bred mice, rabbits, or cattle. Dunn, one of Keeler’s predecessors as caretaker of the Bussey mice, admitted late in his career, “I don’t think that human heredity was ever very far out of my mind while I was doing… things with the mouse.”311

Keeler ultimately completed his graduate work at the Bussey and under Castle. It seems that once the decision had been made Keeler realized the advantages of his arrangement at the Institute. He was able to carry over his work on his “rodless retina” mice to the Bussey, thanks to Castle’s encouragement of his students’ own research questions. Like many of its “inmates” Keeler formed a genuine attachment to the

Bussey. His brief history of the institute, rife with anecdotes, recalled his close association with the Bussey staff including Mr. Patch, the head caretaker, Mrs. Kelley, the custodian, and Billy Reardon, the animal caretaker, and the involvement of Bussey students in local fancy shows.312 Keeler recalled the condescending tone of “medical

310 Keeler to Davenport December 13, 1923. Davenport Papers B.D27.2, Folder “Keeler, Clyde E. #2,” Series IIB, APS. 311 Quoted from Rader, K.A. “The Mouse People” p.337-338. 312 Keeler, C.E. How it Began. p.182, 185.

150 men” and the regular Harvard faculty when they referred to the Institute as the “Cowpath

College.”313 Rather than becoming a medical man himself, as had been his initial goal,

Keeler became immersed in the culture of the Bussey Institute, whose intellectual commerce drew it closer to breeders than to traditional morphologists or physicians.

During his tenure at the Bussey Keeler became a champion for the house mouse as a model for human heredity and disease. Following his characteristic breadth of interest, he published a book entitled, The Laboratory Mouse: Its Origin, Heredity, and

Culture in 1931. The book demonstrates well Keeler’s belief in the unity of knowledge, that most knowledge had value in relation to wisdom from other sources. Now employed by Harvard’s Howe Medical Laboratory as the institution’s first medical geneticist,

Keeler convinced his employers to send him to the Aegean island of Tenedos, to the site of the Temple of Apollo where an ancient cult had associated the god especially with mice, to search for the origins of albino mice. In the book Keeler wove various historical accounts, ancient mythologies, and contemporary laboratory reports into an explanation of the origin and diversity of the modern laboratory mouse. He even provided a Rosetta

Stone of sorts, which offered translations between fanciers’ terms, geneticists’ terms, and genetic notations for the known mutants of the house mouse.314 There was, to him, no reason that the earlier cultural importance of mice should not inform their modern social relevance and usage. According to historian of science Karen Rader, Keeler’s book

“self-consciously used the metaphor of domestication- especially the idea that laboratory domestication represented a process continuous with [Mus musculus’] evolution as a

313 Keeler, C.E. Who Killed the Bussey?- unpublished manuscript. 1982. p.1. 314 Keeler, C.E. The Laboratory Mouse: Its Origin, Heredity, and Culture. Cambridge, Massachusetts: Harvard University Press, 1931. p. 34.

151 human symbiont- to argue that there is no important boundary between past and present practices.”315 The mouse, thus laden with historical and cultural meaning, became

Keeler’s chosen medium with which to build a pioneering career in medical genetics.

Keeler began his work in this new field in a familiar setting. He was hired shortly after earning his Sc.D. as a Research Fellow in the nascent Howe Laboratory of

Ophthalmology at the Harvard Medical School. His duties included research and teaching in the area of medical genetics for which Harvard Medical was poorly equipped, thus he was allowed to work almost daily at the Bussey rather than at Cambridge until the institute’s closing in 1936. Keeler was fortunate to remain under the Bussey’s aegis.

Many Bussey alumni found their opportunities limited outside of Forest Hills. Despite the promise of genetics, few academic jobs had been set aside for the field even in the middle 1920s when Keeler graduated.316 The constant struggle for funding at the Bussey was only a prelude to the difficulties awaiting those who left its grounds, and intellectual freedom was surely scarcer elsewhere. Keeler ultimately learned these lessons when the

Bussey Institute was shuttered in 1936, the result of elite prejudice and petty jealousy in his interpretation. In Keeler’s later reflections on his own career, bitterness surrounds the closing of the Bussey. From that point on he viewed his career largely as a struggle, against envious personalities or sometimes overwhelming and largely irrational forces.

Keeler had already been fired once when his boss Frederick Verhoeff convinced

Lucien Howe that there was no future in medical genetics. Keeler’s departing report, which detailed his studies on heritable health conditions, evidently forced Howe to

315 Rader, K.A. The Metaphor of Domestication in Genetics. In Where the Wild Things Are Now: Domestication Reconsidered. R. Cassidy and M.H. Mullin (Eds.) Oxford, United Kingdom: Berg Publishers, 2007. p.190. 316 Rader, K.A. “The Mouse People” p. 346-347.

152 reconsider and Keeler was restored to the payroll. While at the Howe Laboratory, he discovered the first genetic heart defect and a hair structure mutant, wavy2, and studied several health problems of importance to humans.317 However, “[w]ith the disbanding of the Bussey Institution,” as Keeler later recalled, “I lost my mouse raising laboratory. I took a few cages to the top floor of building C at the Medical School, but the room was quite unsuitable for muriculture and genetic experiments. This and the unfavorable attitude of the Medical Staff in general made my situation untenable.”318 He managed to carry out a few studies on Dalmatian dogs at the Howe Laboratory after the closing of the

Bussey, but it became clear to him that if he wished for his career to have a broad and lasting social impact, he would need to leave Harvard.

Life Beyond the Bussey

While Keeler struggled to establish medical genetics at Harvard, his efforts did not go unnoticed outside of Cambridge. He had established some notoriety as an early expert in a young and socially relevant field. By the middle 1930s Keeler had earned a

Guggenheim Fellowship for his efforts and soon thereafter attracted the attention of Alan

Gregg, Director of the Rockefeller Foundation’s Medical Sciences Division. Gregg used his influence over the Rockefeller purse strings to shape the future of medical knowledge and practice in the United States. He diverted the bulk of Medical Sciences Division funding from programs in medical education to basic research in “psychobiology.” This

317 Keeler, C.E. How it Began. p. 188. 318 Keeler, C.E. Who Killed the Bussey? p. 3.

153 decision was predicated on the fundamental eugenic assumption that many of the nation’s health and social problems were rooted in the indomitable inheritance of the diseased mind. Given Gregg’s emphasis on the inheritance of mental characters, his efforts to develop “psychobiology” fostered growth in medical and behavior genetics, two fields that historian of science Diane Paul suggests are logically inseparable: “There was, in fact, no real distinction between medical and what would later be called behavior genetics. The emphasis on mentality, in turn, reflects the fields’ eugenic origins.”319

Clyde Keeler’s career, begun in one of these fields and ending in the other, is an uncommonly good illustration of Paul’s argument.

Keeler traveled a considerable epistemic distance in his seventy years as a geneticist, but many of his goals and assumptions remained the same throughout. He was committed to improving human health and assumed that many health problems (and their solutions) lie in the germ. He also assumed a relationship between mental and physical traits, that the health of the mind was tied to the body’s constitution. He shared this belief with many other eugenicists, including Frederick Osborn, Director of the American

Eugenics Society, who maintained that if the mentally unsound were prevented from reproducing their kind, “that would take care of their physical characteristics at the same time… Our practical program of eugenics needs then to be concerned only with mental qualities.”320

Keeler’s own position on eugenics is somewhat difficult to determine from the spare mentions of his name in the body of scholarly and popular history devoted to

319 Paul, D.B. The Politics of Heredity: Essays on Eugenics, Biomedicine, and the Nature-Nurture Debate. Albany, New York: State University of New York Press, 1998. p.63. 320 Quoted from ibid. p.64.

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American eugenics. He is most often named in connection with his role on the American

Neurological Association Committee for the Investigation of Eugenical Sterilization, which severely undercut philosophical foundations and early evidentiary support for practical eugenical policy. The report of the ANA Committee, issued in 1936, cautioned against the widespread implementation of coerced sterilization for the infirm, feebleminded, or otherwise disabled. The Committee, headed by the Boston psychiatrist and early critic of eugenic policy Abraham Myerson, was less circumspect about the motivations and epistemic support for sterilization; they declared them illusory and sophistic. Myerson’s group critiqued social degeneracy theory from its origins in psychology to its wider adoption among the biological and social sciences and the culture at large. The report also read: “A great deal of the work which has been done is entirely invalid and has only historical significance… [This includes] the earlier work which comes from Davenport, [Aaron] Rosanoff, and the American Eugenic school with its headquarters at Cold Spring Harbor, New York.”321 Even the section entitled “Genetics and its Relations to Eugenics,” with which Keeler was most involved, began: “On the whole, eugenics receives scant support on any scientific basis from genetics.”322 Keeler believed, however, that his voice had been minimized in the report. When he presented a copy to his former mentor Charles Davenport, he offered by way of self-defense:

321 The Committee of the American Neurological Association for the Investigation of Eugenical Sterilization (Myerson, A., Ayer, J.B., Putnam, T.J., Keeler, C.E., and Alexander, L.) Eugenical Sterilization: A Reorientation of the Problem. New York: MacMillan Company, 1936. p.88. The ANA Committee was assembled from researchers at Harvard Medical School. 322 Ibid. p.69.

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“…although this is ostensibly the analysis of a Committee, the report bears undeniably the individual flavor of the Chairman.”323

Conversely, in Edwin Black’s War Against the Weak, Keeler is suggested to have willfully ignored Nazi anti-Semitism and notions of racial purity in his positive review of the eugenical work being done in Otmar Freiherr von Verschuer’s laboratory at the

Kaiser Wilhelm Institute of Anthropology, Human Heredity, and Eugenics. However, in a widely published review of Verschuer’s Erbpathologie, Keeler sharply criticized the notion that any such thing as racial purity fell along the lines of contemporary political subdivisions: “It is probably quite as difficult for Dr. von Verschuer to close his eyes to the many diverse strains of blood that mingle within Germany’s political boundaries as it would have been for Galton to have denied the Angle, Saxon, Jute, Norwegian, Dane,

Pict, Scot, Irish, Norman, Roman, Phoenician, and other bloods that flow in the English.”

Keeler was, indeed, impressed by the work done in Verschuer’s laboratory and recommended it as a model for American medical genetics. He held, however, that any knowledge thereby gained could and should first be purged of the “phantasy [sic.] of racial purity.”324 Throughout the late 1930s Keeler exhibited skepticism, incredulity, and mockery toward eugenical positions that he held to be untenable, especially those associated with ideas of racial purity and superiority. His evaluation of eugenical

323 Keeler to Davenport January 28, 1936. Davenport B.D27, Folder “Keeler, Clyde E.” APS. Keeler was likely added to the Committee because of his association with Myerson during their shared tenure at Harvard Medical School. Myerson had introduced Keeler to the field of Mental Health and the two maintained a correspondence for a number of years despite the differences in their interpretations of the evidence for inheritance of mental traits. See also: Keeler, C.E. The Gene Hunter. p.200. 324 Keeler, C.E. “National” Eugenics As an Element of Race-National Religion. Journal of Heredity. 1938. Vol.29:71-72. See also: Keeler, C.E. Articles of Eugenic Faith. Journal of Heredity. 1937. Vol.28:19-22.

156 research was neither as uncritical as Black indicates, nor as disapproving as the report of the ANA Committee suggests.

In the years following the report of the ANA Committee, Keeler took multiple opportunities to clarify his stance on eugenics. He acknowledged the sins of eugenics past, including the indiscriminate application of the label (and stigma) “hereditary taint.”

He was convinced that the apparently quantitative nature of many diseases rendered it impossible to expect “a physical or mental ‘Utopia’ to spring at once from our efforts.”325

Keeler was, however, convinced that intrinsic and transmissible disease, especially of the mind, created unknown personal tragedy and an insufferable social burden. There was nothing ethereal or epiphenomenal in Keeler’s eugenical view of the mind. Mental diseases were rooted squarely within the physical body. It followed that their consequent social costs should have a physical solution, lying in the genes.

The Wistar Institute: Site of Discovery and Controversy

At the time that Keeler was feeling stifled in his post-Bussey days at Harvard, the

Wistar Institute of Anatomy and Biology in Philadelphia was struggling to interpret the volumes of data left behind by the late Henry H. Donaldson. Keeler’s name was raised among the staff and administration of the Wistar Institute, and for good reason. Not only did Keeler’s expertise in genetic interpretation suggest him, but also the distance between

Cambridge and Philadelphia was not so great as geography would suggest. The Wistar and Bussey Institutes had established ties shortly after the latter opened in 1908, as both

325 Ibid. p.78.

157 institutes sought to establish rat colonies for research purposes. The laboratories maintained regular contact until the Bussey closed, though their most extensive collaboration began in the fall of 1933 when William Castle and Helen Dean King began their joint work on linkage in the rat.326 At the same time Castle was collaborating with his former student, Keeler, on a series of studies concerning the inheritance and incompatibility of blood groups in rabbits and humans. It is likely that Castle provided introductions between Keeler and the staff at the Wistar Institute at this time, for Keeler was named as a collaborator on a project to study special traits in one of King’s rat strains in the Director’s Report for 1935.327 With the demise of the Bussey and the unsympathetic attitude of his colleagues at Harvard Medical, Keeler resigned from the

Howe Laboratory in 1939 and was hired later that year to assist in King’s Department of

Genetics at the Wistar Institute.328

Keeler’s interest and expertise in medical genetics fit well with the research program at the Wistar Institute, whose investigators had pioneered the use of animal models for studying human health and evolution. By the middle 1930s when Keeler became associated with the institute, the research ongoing there was, in some cases, advertised as explicitly eugenical. Nearly all of the work in the Department of Genetics was centered on analysis of the institute’s rat populations, particularly the strains whose

326 King, H.D. Report to the Director in Greenman, M.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1933. p.28, Wistar Institute Library. 327 King, H.D. Report to the Director in Greenman, M.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1935. p.10, Wistar Institute Library. 328 Greenman, M.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1939. p.3, Wistar Institute Library. Keeler followed another Bussey alumnus, John A. Detlefsen, to the Wistar Institute. Detlefsen had arrived at the Wistar in the early 1920s after several years and numerous difficulties trying to establish genetics in the College of Agriculture at the University of Illinois under Eugene Davenport. Detlefsen is first mentioned in Greenman, M.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1922. p.13, Wistar Institute Library.

158 ancestry could be traced to a handful of gray Norways caught in the alleys, stables, and various unsavory corners of Philadelphia in 1919. From the earliest use of rats at the institute, under Henry H. Donaldson, the albino variety was held to be an exceptionally good experimental model for modern humans. Donaldson, who died the year before

Keeler arrived at the Wistar, had been collaborating with Helen Dean King to describe the changes consequent to bringing the wild gray rat into captivity. He contributed the bulk of the anatomical data for the study, culminating in the pair’s first report in 1929.

King, meanwhile, analyzed various “life processes” of the strain of rats under domestication, including fertility, the reproductive period, sex ratio, behavior, mortality, and mutations. She published her results in the 1929 report and again in 1939 and noted in the latter account that “Dr. Donaldson’s records for organ changes in these generations will be published later.”329 When Keeler arrived at the institute later that year he was given access to Donaldson’s loose anatomical records in the hopes that he would parse out their medical and, more broadly, human relevance.

In the following year Edmond Farris, the new Director of the Wistar Institute, mentioned in his report that “Dr. Keeler… is endeavoring to correlate the findings on the rat with medico-genetic problems of man.”330 Keeler had inherited a wealth of data from

Donaldson’s decades-long effort to uncover the effects of domestication. He went quickly to work on this material and was able to contribute to the institute’s showing at the 1940 American Association for the Advancement of Science meeting in Philadelphia, for which the Wistar Institute played host. Keeler presented a reevaluation of

329 King, H.D. Life Processes… During Fourteen Years in Captivity. American Anatomical Memoirs. 1939. No.17:5. 330 Farris, E.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1940. p.13, Wistar Institute Library.

159

Donaldson’s earlier comparison of the foramens magnum of albino and gray Norway rats. He became fixated, however, on the subject of Helen Dean King’s exhibit entitled,

“Captive Gray Norway Rats and Their Mutant Derivatives.”

King had bred her strains of captive gray rats for more than forty generations to this point and derived from them seven varieties: black, ruby-eyed dilute, albino, albino- waltzer, curly-coated, cinnamon, and cinnamon-stub-tailed. Sports of this kind were the stock-in-trade for geneticists, and King’s mutants were especially appealing for they were of known provenance and manifested their mutations on a nearly uniform hereditary background. The Director’s Report for that year recorded that “[t]he exhibit of Doctor

King attracted unusual attention, and several requests have been granted to reproduce certain of her records in books and articles.”331 In the exhibit generations of King’s cage- bred gray rats were shown descending in a single line, like beads on a string, punctuated by branching events that showed in which generation each coat-color, morphological, and behavioral mutant appeared. The diagram and accompanying text gave the impression that time and evolution had been condensed through the breeding of this strain of rats. In her report of a year earlier King suggested that this abbreviated evolution was the result of the more mutable germ from older and younger individuals that would not have the chance to breed except under domestication’s auspices.332 This solution was, of course, speculative: she had no evidence that very young and very old rats gave rise to more sports than rats in their prime, but this explanation conformed to her assumption domestication had increased the variability of her rats.

331 Ibid. p.16. 332 King, H.D. Life Processes… During Fourteen Years in Captivity. p.62-63. King followed C.C. Little in this interpretation of the nature of domestication’s extreme variability.

160

Even before the AAAS meeting in December 1940, when King’s presentation overshadowed Keeler’s own, he had questioned the validity of many of the findings and interpretations King made concerning her mutant strains. According to Keeler, the captive gray rats in the latter generations had not relinquished the “obnoxious traits” of the early captives, or even of the original feral animals. They hissed, gnashed teeth, and huddled to avoid handling. “They were still thrashing demons,” Keeler insisted, this, after dozens of generations in captivity.333 The albino mutants derived from the captive strain were hardly tamer than their gray cousins and displayed an entirely different temperament than the docile albinos favored in laboratories. The tendency that King saw toward tame behavior in the recent generations of her rats was little more than wishful thinking Keeler concluded. Rather, he trusted the impressions of Ruth Meeser, a breeder and caretaker at the Wistar with daily experience handling King’s rats. According to

Meeser, whom Keeler had inherited as an assistant from the late Henry Donaldson, unless the captive gray rats were gentled before they opened their eyes they were nearly impossible to handle as adults. Even if this care was taken, the rats reacted violently to all but their familiar handler.

Keeler offered a radically new interpretation of King’s experimental results. The attempt at experimental domestication had worked, he claimed. There were domesticated rats within King’s captive strains, though they were not where she assumed them to be. King, and Donaldson before her, had anticipated a gradual change within their colony of gray rats toward the morphological and behavioral condition of the laboratory albino. Keeler offered an alternative explanation in a letter to Robert Yerkes

333 Keeler, C.E. Coat Color Gene Synthesis of Tame Behavior in the Rat, Mink, and Fox. Mind Over Matter. 1964. Vol.9:19.

161 in late September: “[T]he laboratory rat was not tamed by many patient generations of selection by man, but rather at one fell swoop by the introduction of three [M]endelian genes: Albinism, Piebald, and Non-agouti Black.”334 Captivity had not produced King’s mutations; it merely revealed them. Domestication was, thus, the result rather than the cause of single gene mutations, the epiphenomenon rather than the phenomenon. The

Non-agouti allele, Keeler claimed, was responsible for the bulk of the behavioral and anatomical modifications, including diminished cranial capacity, found in the laboratory albino. The reduced adrenal glands, typically found in tame albinos, were also present in nearly the same condition in rats homozygous for non-agouti. Upon noting this correlation, Keeler said of the non-agouti animals, “Those rats couldn’t get mad if they wanted to.”335 Albinism masked the black pelage by which the mutation was known in all of the most commonly used laboratory rat lines, including the stock strain at the

Wistar Institute. Keeler distinguished his “genetic approach” from the Lamarckian and orthogenetic views that he claimed were still guiding interpretation at the Wistar at the time of his arrival.

A product of the Bussey Institute, Keeler was undoubtedly prepared for genetic analysis and was inclined to investigate the coat color mutants that emerged within

King’s captive strain. The staff members of the Wistar Institute were not, however, ignorant of genetic explanations and had tried to implement them in their studies of the rat. Shinkishi Hatai had determined that black and piebald were latent in the normal

Wistar albino stock in 1914. King had taken an interest in her coat color mutants as they

334 Keeler to Yerkes September 25, 1941. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. Manuscripts and Archives, Yale University Library (hereafter YUL). 335 Quoted from: St. John, W.F. Research Scientist at Milledgeville Learns about Redheads from Mice. Atlanta Constitution Magazine. October 17, 1948.

162 appeared and endeavored in the latter half of the 1910s to correlate coat color with

“physiological characters such as growth, fertility, longevity, etc.”336 She had also noticed the peculiar nature of the black (Non-agouti) mutant rats in her experimental stocks in 1929: “Rats of this black race exhibited none of the vicious traits of the wild

Norways; they are as tame and as easily handled as are the Albinos…In none of their characteristics, apparently, do the two races differ, except in coat color.”337 Donaldson, too, had looked for a relationship between coat color and the weight of the hypophysis with the help of Ruth Meeser. In turn, Meeser noted the importance of the latent black allele for the diminution of brain size in 1929: “Where black is evident in the color, brain weight is low. Important: I find where black is latent W.I. albinos brain weight low- where black factor is absent mutant albinos brain weight is high.”338 By Keeler’s own admission it was the caretaker, Meeser, who had brought the idea of correlated coat color and docility to his attention.339

In his several accounts of the discovery, Keeler recalled Meeser asking him at the dissecting table, “Do you hear that rat squealing?” When he replied that he had,

Meeser added, “It’s a black one.” A search in the Colony Office confirmed Meeser’s suspicions. She later identified an albino by its signature call and pointed out differences in muscle tonus among coat color mutants. What clinched the argument, though, was her demonstration of altered behavior among coat color varieties of the Norway rat. With the

336 King, H.D. Report to the Director in Greenman, M.J. The Wistar Institute of Anatomy and Biology Director’s Report for the Year Ending December 31, 1916. p.8 and 1918. p.11., Wistar Institute Library. 337 Quoted from Burks, B.S. Temperament and Single Gene Substitutions in Wistar Rats. Folder “Rat Articles,” Wistar Institute Library. Originally received by C.E. Keeler October 4, 1943 from Robert Cook. 338 Ibid. p.3. 339 Keeler, C.E. The Gene Hunter. p.160-163.

163 help of heavy gloves and forceps Meeser ushered a mixed lot of coat color varieties, “one by one, the vicious shrieking, thrashing, jumping, urinating, defecating monsters,” into a new cage. Keeler recalled, “When she had transferred all but three or four, she removed the gloves, laid down the forceps and lifted out a quiet, unperturbed [black] rat on the palm of her hand… From that moment on, the principle of pigment gene pleiotropy, with effects on morphology, physiolgy [sic.] and psychosomatic behavior became the principal object of my research.”340

Still Keeler felt that by late 1941 the discovery was his to claim and there was little reason for him to think otherwise. He acknowledged Meeser’s role in observing and even in formulating the idea of correlated color, morphology, and behavior. He seemed, however, to be unaware of her depth of understanding about coat color genetics. Rather, he treated her observations like those of other breeders, fanciers, and caretakers he had known- as valuable source material for new genetic analyses. King’s observations relating to the peculiar character of rats with the non-agouti allele were similarly unknown to the public and were probably not well known within the walls of the Wistar

Institute. Communication and institutional memory were sometimes lacking at the

Wistar. Thus, different researchers recapitulated numerous matings and experiments over the years of the domestication program apparently without knowledge of the preceding trials.

Keeler set to work performing crosses of the various Wistar strains, including mating wild gray to stock Wistar albino rats (previously carried out by Hatai and again by

King), and divined the presence or absence of Non-agouti, Piebald, and Albino alleles,

340 Keeler, C.E. The Gene Hunter. p.160-163.

164 latent or manifest, in nearly all of King’s mutant strains. He then correlated this information with Donaldson’s rich anatomical records and communicated his discovery and his enthusiasm at least to Robert Yerkes by late September. A month later he presented his thesis before a Zoological Seminar at the University of Pennsylvania. The press immediately heralded the significance of Keeler’s findings. Thomas Henry of the

North American Newspaper Alliance reported that Keeler’s “discovery of a ‘civilization gene bundle’… may prove to be the most significant in biology since the original discovery of the basic laws of heredity.” Henry imagined that Keeler’s work might

“eventually point the way to developing greater control of man’s emotions- in effect breeding more civilization into him- and thus reduce or eliminate war or crime in the world.”341 The talk was subsequently featured in an editorial entitled “Domestication

Genes” in the Journal of Heredity. The journal’s editor, Robert Cook, suggested that

Keeler’s findings “may herald a far-reaching discovery in the genetics of intellect and emotion.”342 Cook’s desire to understand the significance of the discovery was shared by many others, including Yerkes, other early recipients of Keeler’s report, and Keeler himself. To this point, however, no official report of the discovery had been subjected to professional scrutiny, despite its widening notoriety. As winter approached, Keeler wondered aloud to Cook about the impact of his theory and how it would hold up to scrutiny at the impending AAAS meeting in Dallas: “The darned thing has such wide

341 Henry, Thomas R. Discover Temperament to be an Inherited Character in Animals. The Calgary Herald. October 31, 1941. p.21. 342 Cook, R.C. “Domestication Genes.” Journal of Heredity. 1941. Vol.32(11):400-401.

165 implications in evolution and psychology that I would like to anticipate some of the questions that will undoubtedly arise when it is presented.”343

The implications were, indeed, wide. Keeler’s reinterpretation of domestication was a dramatic break with almost all previous views, including his own. Instead of the timeless march of slow progress and human ingenuity or conquest over nature, domestication was made to seem like a fortunate allelic accident, the providence of random mutation. Of those who learned of Keeler’s discovery, few were satisfied to limit its interpretation to animal domestication. The importance he attached to a select few coat color mutations countered the view that evolution proceeded by the slow accumulation of relatively small effect gene mutations, which was then gaining traction among a number of prominent biologists. His deterministic suggestions about the heritability of complex personality traits threatened to fuel longstanding debates contrasting the physical and epiphenomenal views on the nature of the mind.

Contemporary American schools of functional, social, and behavioristic psychology denied that behaviors were so strongly dependent upon physical heredity. Each of these schools maintained the importance of the environment, whether a particular social interaction, adaptation, or recognized stimulus, for determining animal behavior. Robert

Yerkes, who was well aware of the discourse ongoing in American psychology, warned

Keeler ahead of the Dallas meeting, “[Y]ou can well afford at present to be extremely cautious and conservative, to avoid generalizations, and to minimize your commitment to interpretations. I think you have leads that are almost certain to prove important, and their development can proceed most advantageously if you avoid arousing controversy at

343 Keeler to Cook December 5, 1941. Robert C. Cook Papers 1882-1992. Library of Congress.

166 this point.”344 Such a bold thesis, it seemed, was likely to arouse incredulity or even hostility from the adherents of vastly different views of evolution, psychology, and inheritance.

Keeler had other reasons for concern. He knew that the report was premature, lacking especially evidence on the psychological side of his putative genetic, morphological, and behavioral relationship. Aside from a few measures of vocal tone and frequency made with an oscillograph, the crucial behavioral evidence came from anecdotal experience, that is, from the experience of handling each mutant strain in no systematic way. He also had difficulty demonstrating that the relationship between coat color genes, behavior, and morphology was a generalizable phenomenon rather than a peculiarity of the strains investigated at the Wistar Institute. Keeler faced problems of another nature as well. Just weeks before the Dallas meeting, he complained to his friend

Robert Cook that his co-presenter, Helen Dean King, seemed to be angling for more than her due share of credit.345 Buoyed by his own enthusiasm and the excitement that surrounded his preliminary reports, Keeler nevertheless delivered, alongside King, a talk on the “Multiple Effects of Coat Color Genes in the Norway Rat, with Special Reference to the ‘Marks of Domestication.’”

Keeler and King’s presentation sought to mitigate, to some degree, the impressionistic nature of their behavioral work. Even as they felt obligated to admit that

“…our measures of wildness and tameness are of the impressionistic sort,” their use of bar graphs masked their rough descriptions of emotionality (e.g. savageness and

344 Yerkes to Keeler December 6, 1941. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. Manuscripts and Archives, YUL. 345 Keeler to Cook December 5, 1941. Robert C. Cook Papers 1882-1992. Library of Congress.

167 wildness) by displaying them alongside more easily quantifiable morphological metrics.

Still, they expressed extreme confidence in their early perceptions: “We believe that the

[behavioral] differences between strains are so distinct that their validity is certain.”346

Going against Yerkes’s advice, the pair felt warranted in explaining the broad implications of their discovery. On the subject of domestication they suggested that “coat color genes may have played an even more important role than has man in the domestication of farm animals by providing attractive coat-color correlates for certain preferred domesticative behavior complexes that man desired to select,” an idea reminiscent of breeders’ notions about the manifold importance of color.347 In a section entitled “Human Applications,” they ventured:

[P]ersonality may be synthesized by combining various behavior- modifying genes… The implications are in line with certain data in the fields of psychiatry and psychology which tend to indicate that this mechanism applies also to man, and our demonstration in the case of the Norway rat seems to necessitate a consideration of hereditary factors in all future experiments dealing with behavior.348 Despite the talk’s shortcomings and overstatements, it garnered a good deal of enthusiasm for the discovery and its implications. Associated Press coverage of the

A.A.A.S. meeting reported that “A number of scientists said the discovery is one of the most significant in years. For it goes far to show why color, shape and texture in people can be also an inseparable part of their characters.”349 Barbara Burks, a psychologist at

346 Keeler, C.E. and King, H.D. Multiple Effects of Coat Color Genes. p.246. 347 Ibid. p. 249. 348 Ibid. p. 249-250. 349 Blakeslee, H.W. “Science Links Hair Tints, Temperament: Similar Genes Control Rats, Human Traits.” The Daily Times Herald (Dallas, Texas) December 30, 1941.

168

Columbia and Robert Cook’s fiancée, reported that a “‘Spokesman’ has said … that the paper was a high runner-up for the annual A.A.A.S. prize.”350

Keeler had hoped for, and perhaps anticipated, the recognition the Dallas paper had received. He was motivated to present the discovery in its preliminary state, not for lack of any critical faculty, but because he feared being squeezed out of a crucial breakthrough that he believed was rightly his. He wrote to Cook that Helen Dean King had started making bolder claims about her priority, perhaps because she perceived

Keeler as a threat to her own, justifiable claims on the discovery. Keeler was most upset, however at being left out of the narrative at all. King, he claimed, had told a Philadelphia

Inquirer reporter that “she wrote the Dallas paper, which is the story now being put out by the administration. I was merely the assistant!... The stormy petrel did not know what was in my paper until the day before I gave it at the Zool. Soc. Seminar! Of all the crust… [T]he reporter might be sligh [sic] enough to see that there is something queer about the set up… And yet the lady is cagey.”351 In another letter to Cook he recounted facetiously, “I had the pleasure (?) as anticipated of hearing H.D.K. relate how she conceived the problem of the mutant rats… I had copies of the Dallas paper on hand, so the [Board of Managers] got copies (much to Farris’ dismay) when they visited my room.

I guess King can’t take a part of the credit from me now, but I hate like hell to see her rob

350 Burks, B.S. Temperament and Single Gene Substitutions in Wistar Rats. Unpublished, p.2- Originally received by Clyde E. Keeler from Robert C. Cook, October 4, 1943. Folder: Rat Articles, Wistar Institute Library. 351 Keeler to Cook April 9, 1942. Robert C. Cook Papers 1882-1992. Library of Congress.

169 the grave even if Donaldson can’t cash in on the glory. The old man had some wise ideas, but did not live to see them materialize.”352

In the months that followed, Keeler gave the impression that the Wistar Institute under Farris was lurching toward its demise, sustained at this point only by selfish and political motivations and personal vendettas. He argued that King and Farris had complementary agendas, which would profit from the selective removal of their political enemies within the Wistar. King began to assert her influence by having Louise Duhring, a fixture of the Wistar under Milton Greenman and King’s “life-long checkmate” in

Keeler’s description, fired. Keeler further claimed that King and Farris conspired to undermine the Advisory Board by canceling its annual meeting in order to dismiss

Armondo Augulo Y Gonzalez, Richard Weissenberg, and Richard Hugh McCoy without objection and to enact new policy without oversight. For these insults, George

Washington Corner and George Wislocki resigned their long held posts on the Advisory

Board. The same tactic was to be used again in 1942 when it was announced that

Keeler’s contract would not be renewed.353 King herself had been threatened with

352 Keeler to Cook June 3, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. Reprints of the published version of the Dallas talk (Journal of Comparative Psychology. 1942. Vol.34:241-250.) arrived with an addendum by Keeler, in which he disavowed the version of events leading to the discovery, as it did not recognize Donaldson’s contributions. He wrote, “At the time of the [University of Pennsylvania] Zoological Seminar and thereafter the credit to Dr. Donaldson went unquestioned until I rewrote the manuscript for presentation at the Dallas meetings, at which time credit for these items was altered by powers beyond my control to read as given in the reprint.” See “Contributions- Keeler” No.7, Clyde E. Keeler Papers, Series 8, Box 6. Special Collections, Georgia State College and State University, Milledgeville, Georgia (Hereafter GCSU). 353 Keeler to Cook April 9, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. In his autobiography, Keeler did not mention much of the controversy with King directly and suggested instead that economic concerns rooted in the war forced Wistar to shed several of its research fellows and assistants. Finances, though, had been used in conjunction with other reasons for firing employees at the Wistar in years past, see chapter 3 this volume. See also Keeler, C.E. The Gene Hunter. p.159.

170 dismissal under similarly ambiguous circumstances 20 years earlier.354 In the intervening years George E. Coghill fell to the same fate. In both cases the Advisory Board had been circumvented and the Board of Managers had complied with the wishes of the Director.

In line with Keeler’s version of events, perhaps King had drawn from her own example the means for removing her political enemies.

To complicate matters further Keeler and King’s priority dispute coincided with the United States’ entry into World War II. Keeler, a reserve infantryman, was warned that he would soon be called into service. The Army kept Keeler in a sort of limbo for months, which he worried would force him to forfeit his idea to King and her co- conspirator, Farris. At bright points in this period, Keeler was able to turn back to the work on mutant rats: “It looks now that if the war can wait for my talent a few days longer, I shall be able to tie up the behavior due to unit coat color gene action with its intermediary steps 1) brain 2) glands 3) sense organs.”355 He hoped to produce a more complete paper on the subject, which would include new genetic interpretations and more rigorous analysis of the behavior side of the phenomenon. Most importantly, he wished to ensure that he would not be denied credit for its discovery. Thus, he prefaced the first draft of the paper with an historical review that “[recounted] the Donaldson and Hatai work leading up to the discovery of the effect of the black gene on behavior and Miss

Meeser’s role in it,” in addition to his own, primary, role. “The whole story that I have on this would make a large book,” he relayed to Cook, “so I’m only trying to rush enough to patent the behavior and a few other things that Farris is now planning to take over for

354 See the preceding chapter in this volume for more on King’s near dismissal from the Wistar Institute. 355 Keeler to Cook March 28, 1942. Robert C. Cook Papers 1882-1992. Library of Congress.

171 his own field as soon as I leave... This is exactly what my wife said would happen about

9 months ago- when I had to decide whether to throw up the white flag or fight for it, and go down fighting if need be.” As time passed Keeler grew more despondent about his chances for securing priority. He was without work, without a laboratory, and still marooned by his indefinite status with the Army. In his own words: “Those who are crushed by the gods are crushed indeed.”356

Of course, he realized that his claims to priority would be all the more convincing if he could fill in the gaps in evidence and interpretation from the preliminary report given at Dallas. He tried to establish the universality of the docilizing effect of the non- agouti allele by finding it latent in strains of domesticated laboratory rats used across the country. He also made new crosses between tame albinos and wild gray rats to see if extracted black rats retained most of the docile temperament from their tame parents. He sought out new black mutants that appeared among strains of gray Norways to determine if their behavior, and not simply coat color, was strikingly different from that of their parents. To provide measures of temperamental differences, Keeler used a battery of simple behavioral tests, including handling, emergence, and annoyance tests. “I do want the thing to be technically without flaw, if and when it sees print,” he wrote Cook.357

Keeler’s tests were simple and he generally neglected social interactions between rats, favoring instead their emotional response to human contact, which Keeler regarded as more relevant to the phenomenon of domestication. His tests, though, did not differ significantly from other contemporary evaluations of emotionality.

356 Keeler to Cook August 24, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. 357 Ibid.

172

With this paper Keeler also hewed closer to Yerkes’ advice about cautious and minimal interpretation.358 His experiments on rats yielded discussion of rats, not of humans and the potential of behavior synthesis. The exuberances of the Dallas paper had almost certainly been Keeler’s own, as they are congruent with the “flights of imagination” that he shared with Cook and Yerkes.359 In the new manuscript, however, they were conspicuously absent. His friends and early reviewers persuaded him that a conservative presentation would lend him more credulity, especially as he was embroiled in a controversy over the ownership of an idea. Cook forwarded the draft Keeler had sent him to Barbara Burks and from there it reached L. C. Dunn, Bussey alumnus and her

Columbia colleague.

Dunn largely approved of Keeler’s evidence for the association of coat color genes and general behavior, but challenged several of his interpretations. Of the phenomenon itself, Dunn questioned whether the association of such distinct traits was not the result of linkage, or close proximity of two genes on the same chromosome, rather than pleiotropy, or the multiple actions of a single gene. Although the concept of a single factor with multiple effects dates back to Mendel’s work on peas, linkage was, by the early 20th century, considered the more likely and certainly the more demonstrable explanation for associated traits. Keeler was “downcast” upon hearing this criticism. He felt that the co-occurrence of black coat color and docile behavior in the original non- agouti mutant strain and its recurrence in multiple crosses without ever segregating was evidence enough to suggest pleiotropy over mere linkage. “You may say that this is

358 Keeler, C.E. The Association of the Black (non-agouti) Gene with Behavior. Journal of Heredity. 1942. Vol.33(11):371-384. 359 Keeler to Yerkes October 8, 1941. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. Manuscripts and Archives, YUL.

173 reducing it to a molecule-splitting controversy,”360 Keeler wrote to Cook, but linkage did not reflect what he had known from the experience of breeding and handling these animals, nor did it allow for such rich implications about the associated nature of the body and mind. The association was invariable, he argued, and he would postpone publication until he could demonstrate that fact. Cook convinced him, however, to publish with the caveat that linkage could not be ruled out, reminding him with an editor’s grace: “I can’t see that either the common sense of the situation, or the position you find yourself in at the moment justify you leaning sort of over-cautiously backwards… Like you, I am betting on pleiotropy. Whatever it is, I want it to be the

‘Keeler-effect’ and not the ‘Farris-effect.’ So get going, big boy!”361

Keeler was more cooperative with the other recommendations from Dunn and

Burks. They cautioned against making grandiose claims or reaching too far in his interpretations. No such overstatements appear in the finished article. Burks, in agreement with Dunn, commented that the introduction was “too discursive” and contained too much history, which in light of Keeler’s priority dispute might be viewed with suspicion. She reminded him of an article she had in preparation that would cover the historical development of the idea from the perspective of an outside, and ostensibly objective observer. She assuaged, “We can certainly see to it that a reprint of my historical article is sent to every geneticist who might be interested.”362

Burks meant for her article to relieve some of the anxiety of the priority dispute both for Keeler and for Cook, who had taken on some risk in advocating for, even

360 Keeler to Cook August 24, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. 361 Cook to Keeler September 17, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. 362 Burks to Keeler July 9, 1942. Robert C. Cook Papers 1882-1992. Library of Congress.

174 privately, one side of an unsettled controversy. As Burks had promised, the article depicted not only the technical history of the discovery but also its far-reaching implications. Her own enthusiasm for the idea was clear and compensated for the dry, restrained delivery she advised Keeler to affect.

Burks had established her career in psychology by evaluating the influence of nature and nurture in determining personality and intelligence, usually concluding in favor of nature. She, thus, remarked, “The implications for this 1941-42 announcement from Wistar are therefore more novel, more pregnant, for psychology than for genetics.”

Despite her association with Keeler, her historical account placed the formation of the

“nutshell” of the idea in the minds of Donaldson, King, and Meeser before Keeler had become involved with the domestication project. In a letter to Cook, she pointed to “data that Keeler hadn’t known about, which to some extent modify his claims as to the firstness of ideas.”363 She reserved special consideration for Miss Meeser, the technician and breeder whose careful observations were the seed of Keeler’s pigment gene pleiotropy. “Laboratory assistants,” Burks wrote, “who live with the animals which are the subject of research procedure and who learn to know them not as abstract building blocks in the construction of an hypothesis, but as pulsating individuals who bite or do not bite under a given set of circumstances are an essential part of the scientific process of wresting facts from a reluctant universe.”364 In preparation for the article and in the spirit of firsthand experience, Burks paid a visit, or “pilgrimage,” to the Wistar Institute in February 1942. At that point she noted, “[a] beginning has been made in the

363 Burks to Cook July 2, 1942. Robert C. Cook Papers 1882-1992. Library of Congress. 364 Burks, B.S. Temperament and Single Gene Substitutions in Wistar Rats. p.7.

175 application of rigorous laboratory tests, but the differences in these strains are so gross that merely to see is to be convinced.” Keeler showed her the various coat color strains of rats housed at the Institute: “Now going from cage to cage, watching Keeler take out rats at random, the writer acted as reporter, and the rat personality pictures took vivid form.” At one of the cages, an albino mutant, untempered by the non-agouti and piebald alleles found in the tame Wistar albino stock, bit into Keeler’s finger. “Keeler carried his wound with pride during the remainder of the tour- it was a symbolic wound!”365 It was evidence literally in the flesh, the very kind of experiential confirmation that Keeler so valued.

Although King was given nearly equal credit for the discovery, Keeler was pleased that this secondary account recognized his contributions as well as those of

Donaldson and Meeser. Keeler belatedly thanked Burks in late May 1943 for her courage and efforts on his behalf: “Most persons shrink from anyone upon whom the hierarchical finger of wrath had been placed lest they get mud splashed on their own raiment. But you and Cook have fought staunchly beside me even when Castle and my wife repeatedly urged me to abandon the field.”366 He was hopeful that the “crusade” for recognition had been successful, even as he lamented about the lasting damage the controversy would bring to his reputation.

The history of pigment gene pleiotropy at the Wistar, which Keeler had forfeited to Burks in the interest of academic propriety, had not, however, seen print. Nor would it. Burks was found dead near the piers of the Hudson Bridge on May 25, hours after

365 Ibid. p.6. 366 Keeler to Burks May 20, 1943. Robert C. Cook Papers 1882-1992. Library of Congress.

176 absconding from the Payne-Whitney Psychiatric Clinic where she had been receiving treatment for the last month. In one of her last letters to Cook she wrote: “I have been thinking a lot about nature and nurture lately, and it is a puzzling business.”367 Cook, in turn, framed her death in these terms. She had faced considerable pressure with her research and funding, but these had relented while her melancholy remained. More to the point, Cook thought, she had sensed a change within herself that “was not merely part of the depression but something more fundamental.” She described competing with a secondary personality, part of a “psychothymic temperament she inherited from her father,” who had died the previous November. The sense of her deepening illness, the gradual loss of her identity, and the prospect that this was all inevitable drove her further to despair Cook surmised, “and she definitely chose not to risk reaching a point where she could not destroy this emerging antithesis… [I]t seems to me that her death was no more ‘unnatural’ than [if] she had had a ruptured appendix or thrombus. Some part of her body was sick, and it happened to be an obscure sickness of neural tissue.”368

Keeler’s relation to this tragedy seems slight and coincidental. His late letter gave little indication that he knew the severity of Burks’ illness. Her disease was, however, precisely the kind of ailment that had brought Keeler to study medical genetics in the first place, a mental disease ostensibly rooted in the body and passed on through the germ.

Burks, Cook, and Keeler had been thrown into association over their shared interest in

Keeler’s deterministic explanation of personality inheritance. The implications for them were clear: gene-determined personality pointed to the physical basis of mental health or

367 Quoted from Cook to Yerkes November 12, 1943. Robert Mearns Yerkes Papers. Series I, Box 11, Folder 195. Manuscripts and Archives, YUL. 368 Ibid.

177 disease and perhaps to the tools by which they could be controlled. The same thinking, though, may have convinced Burks that she was doomed by heredity. “Nature and nurture are a puzzling business,” Cook echoed his lost fiancée, “and we are only on the threshold of knowledge. The time will come I hope when so gifted and brave and gentile

[sic.] a mind and personality would have avoided that sad fate.”369 This was the promise and peril of the scientific work done on this dichotomy, the real gravity and urgency of the work to find solutions to the critical problem of mental health. Keeler had lost an advocate, a relatively minor loss in the scope of this tragedy. It had, however, the practical consequence of removing the historical account of the discovery from wider circulation. Keeler himself did not receive a copy of Burks’ draft report until the fall of

1943 as Cook uncovered it among her personal effects. His priority, thus, hinged on his follow-up report, which had been purged of his version of events.

Keeler’s last-ditch effort was evidently enough. The sword, represented by King and Farris’ imminent publication, never fell. Farris did publish, with Eleanor Yeakel of the Wistar, results similar to Keeler’s, but not until 1945 when the controversy was hardly fresh and he could only cite and confirm the findings of Keeler (1942).370 King had by then turned her attention to the rich field of cancer genetics. Keeler had won.

Pigment gene pleiotropy, for better or worse, was his legacy. The conflict between

Keeler and King was, of course, about priority, but it was as often described in terms of the difficult and combative personalities involved. In the race to patent the means of

369 Ibid. 370 Farris, E.J. and Yeakel, E.H. Emotional Behavior of Gray Norway and Wistar Albino Rats. Journal of Comparative Psychology. 1945. Vol.38:109-118.

178 genetic behavior analysis and synthesis, Keeler exacerbated his reputation as an insufferable personality.

Research in Exile: Pigment Gene Pleiotropy After Wistar

Meanwhile, Keeler’s career troubles continued. He was still unsure of the Army’s designs for him. Prospective jobs at the Lankenau Institute for Medical Research, Clark

University, and Brooklyn College never materialized. He reluctantly registered with a teacher’s agency, which placed him at The Edgewood School in Greenwich, Connecticut where he was assigned courses in biology as well as nutrition and first aid. Between

1943 and 1945 Keeler migrated south, following a trail of one-year appointments in

North Carolina and Georgia. With each leg of his journey, he drifted further from the centers of biological research, funding, and esteem where he had begun his career nearly twenty years earlier. The institutions where he found work were as remote nominally as they were geographically: The Women’s College, University of North Carolina

(Greensboro, North Carolina); Wesleyan College (Macon, Georgia); and Georgia State

College for Women (Milledgeville, Georgia).371 By the middle 1940s genetics had reached the provinces, including women’s colleges, and women had earned distinction in the field (e.g. Keeler’s arch-nemesis, Helen Dean King, who was graduated from Bryn

Mawr, though before it or any school offered training in genetics). In most cases, however, women’s colleges lacked the resources- equipment, funding, and personnel- of

371 Keeler may have chosen to leave the first two appointments, where he reported conflict with a anti-evolution preacher (Women’s College) and a feminist cabal who threatened to stifle his career (Wesleyan). He remained in Milledgeville for the remainder of his career, despite other controversies. See Keeler, C.E. The Gene Hunter. p. 159.

179 the universities of the Northeast, or even the state universities further west, especially if they resided in the South. Keeler could not have regarded these moves as career advancements, doubly marginalized as he was. He remained blithely optimistic, however, about the potential for the concept of pigment gene pleiotropy. In the midst of his search for long-term employment, he imagined leading a research group comprised of the lions of neurology and psychology to tackle the heritable complex of associated body and mind: “I want Augulo to cooperate on the neurology and either Stone, or Hall, or

Lashley or all three or some of their students to do the testing of the animals I synthesize.

That would put the results beyond criticism… [T]he proposition could be used to build a whole Institute of Genetic-Psychology or Psycho-genetics.”372

Keeler, of course, understood his circumstances following his academic exodus.

Without adequate and semi-permanent laboratory space he could not hope to perform any meaningful research involving multiple breeding strains of rats. From 1943 to 1947 his published papers included few data that had not been collected prior to his leaving the

Wistar Institute in 1942. He reasoned, however, that he could still gather evidence to demonstrate that coat color gene-determined personality was a general phenomenon. He had begun this effort by surveying strains of tame albino rats in use in laboratories across the country, other than the Wistar strain with which he had made the initial observation.

As early as 1942 he sent letters and questionnaires to mink and raccoon breeders, asking them to evaluate the temperament of coat color varieties in these animals recently brought into captivity. “Our first duty is to collect and eventually to sift all available isolated reports in order to select the materials most suited for further adequately

372 Keeler to Cook June 15, 1942. Robert C. Cook Papers 1882-1992. Library of Congress.

180 controlled inquiry,” he wrote later.373 In other words, he actively sought anecdotal support for the idea of associated color and behavior with the knowledge that experimental confirmation of these reports would be postponed indefinitely.

There was a practicality to this decision. Keeler had no means of conducting controlled studies in this period. His status necessitated this approach, if he was not to abandon the field. As late as December 1944, he struggled with whether to continue publishing on the “rat material which is well controlled mathematically before we launch out into the general field where there are many observations, but where the observations are not backed up by mathematically checked analyses.”374 Keeler, despite his candor about interacting with breeders, delayed publication of these observations for fear of seeming uncritical. As his exile wore on, however, he worried more about being forgotten. In each of his stations during these years, he gathered impressions from local experts and more distant correspondents. His case for pigment gene pleiotropy increasingly relied on the impressions of men and women whose experience had never included a laboratory. Keeler introduced his unorthodox approach in a letter to the editor of the Journal of Heredity (Cook). He relayed that “this part of the Deep South offers exceptional material” for the study of associated coat color and temperament in certain stock animals. He quoted a “Negro preacher” in a stereotyped drawl, it seems in order to preserve the authenticity of the account:

A gray mule is the lastines’ mule dey is. Ah knows dat fo a fack, an ah bin a stock man all mah life. Ah guess it’s cause de way he lives. Now a brown Texas mule, he jes’ weahs hisself out, rushin’ round an’ always

373 Keeler, C.E. Coat Color, Physique, and Temperament: Materials for the Synthesis of Hereditary Behavior Trends in the Lower Mammals and Man. Journal of Heredity. 1947. Vol.38(9):272. 374 Keeler to Cook December 30, 1944. Robert C. Cook Papers 1882-1992. Library of Congress.

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havin’ a fit. But a gray mule jes takes it easy, an’ you caint nevah make him hurry and deys jes no tellin’ how long he’ll live.375

What began as informal queries in 1942 became the substance of Keeler’s publications by 1947. His use of breeders’ accounts was not motivated solely by the pressing need for evidence and material for publication. He placed a high value on the evidence of experience. Thus, he wrote, “One has only to attend a Dog Show in order to find ample evidence to prove that breeds inherit ‘behavior trends’ with reference to particular situations. That is, individuals of two contrasted breeds tend to solve the same problem or situation in different but characteristic fashions, which behavior patterns are inherited somehow in the breed.” Keeler also began to use more reports from men and women experienced in animal breeding and training to support the concept of pigment gene pleiotropy. In a 1947 paper, he gave titles to his “unimpeachable sources,” but distinguished them only by their expertise with a particular animal. His sources included

Charles Darwin (and by association, Darwin’s contacts), professors, zookeepers, trainers, dealers, cowboys, Indians, colonists’ accounts, skinners at meat packinghouses, breeders, and fanciers. Keeler argued that the commonality among them, besides their ability to furnish an example of associated color and behavior or morphology, was their discernment: “it is not the imaginative person who is intrigued by the idea, but rather the careful, keen observer, who can describe particular cases.”376 Others, however, wondered if it was Keeler who was not being shrewd enough while amassing his evidence. An unnamed, “noted mammalogist, suggested that ‘perhaps these reputations are in the

375 Keeler, C.E. Coat Color and Temperament in Mules. Journal of Heredity. 1945. Vol.36(8):243. 376 Keeler, C.E. Coat Color, Physique, and Temperament. p.272.

182 category of old wives’ tales.’”377 Thus, the link was made between Keeler’s evidence for genetically related color and temperament and broad, traditional, and frequently unreliable cultural knowledge. Keeler admitted, “Much of the following list of cases must by necessity be anecdotal,” but reminded his readers that “here fruitful lines of scientific investigation often start.”378

Keeler’s appreciation for experiential knowledge of animal breeding and care resonated with his proselytizing efforts in eugenics. The arbiters of American eugenics and the breeders of the nation expressed mutual sympathies that extended beyond the notion that artificial selection could be applied to humans as it had been with animals and plants. As Charles Davenport relayed to his former student, Robert Yerkes:

…at the present moment [I] think that the Eugenics work finds a more sympathetic association with students of heredity than with students of medicine; with breeders of corn than with state sanitarians… I find that mothers of families and farmers are more interested in Eugenics than medical men. This is because the more active medical men, who have been brought up in cities and have gained their training largely in municipal hospitals, have really never come in contact with the facts of heredity; whereas every mother of a large family and every breeder of fancy stock and of pedigreed plants, knows already a lot about heredity.379

Breeders and eugenicists were at the same time linked by shared political and social concerns. It was not coincidence then, argues historian of science Barbara

Kimmelman, that American eugenics found its first nationwide representation within the

American Breeders’ Association. Kimmelman demonstrates that Nativist political agendas, cultural norms, and pastoralism underlay both the popular agriculturalists’

377 Ibid. 378 Ibid. 379 Davenport to Yerkes, September 18, 1911. Robert Mearns Yerkes Papers. Series I, Box 13, Folder 230. YUL.

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Country Life movement and the budding eugenics movement in the early years of the

20th century.380 American eugenics, thus, grew in the company and culture of breeders.

Both agriculturalists and eugenicists respected the rigor and apparent objectivity of scientific methods, but for the matter of becoming convinced of the explanatory power of heredity, for both groups, seeing was believing. The USDA Yearbook of Agriculture for

1936 emphasized the primacy of firsthand experience in the breeding of plants and animals: “…observation and judgment, of course, is supplemented by scientific tests and precise measurements, but the feeling for the material must come first.”381

Keeler showed a similar high regard for the judgment of experienced hands, in his genetic research and his support for the eugenic cause. Just as work with animals had convinced many breeders of the potential of eugenical policies, he argued that the best strategy for making local leaders in the movement was by “encouraging experience relative to eugenic problems. I have induced several persons, with a casual interest in eugenics to study some particular trait in a defective family of their own community, and to make as complete and detailed a survey as possible. The investigator is never the same after such an experience.”

Keeler’s pedagogical preferences seemed to reinforce this conviction of the value of sensory experience. In his avocation as an artist, Keeler made many renderings and sculptures for the purpose of illustrating biological ideas, including flipbooks for explaining fertilization and mitosis, terra cotta figurines to demonstrate coat color inheritance at the Harvard Tercentenary, and numerous drawings for articles in the

380 Kimmelman, B.A. The American Breeders’ Association. p.183-189. 381 Hambidge, G. and Bressman, E.N. Better Plants and Animals: A Survey of Superior Germ Plasm in Yearbook of Agriculture. Washington, D.C.: United States Department of Agriculture, 1936. p.126.

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Journal of Heredity. When it came time for him to explain the concept and implications for pigment gene pleiotropy, he tried to provide as rich a sensory experience as possible.

In 1947 he “represented diagrammatically the effects of a single gene upon various developmental reactions by means of the branching of a tree… [where the] tips of the twigs symbolize the completion of physical development.”382 The tree, he explained, was situated in a dense jungle of trees of reaction, where only the canopy was visible to the eye. At the base of each tree, however, lay a single mutant allele that gave rise to these reactions. He also referred to the basal mutant allele as a “handle.” The meaning of this sobriquet he explained in a letter to Robert Yerkes: “The greatest advantage that I see to my approach is the ‘coat-character handle’ by which we can lift out or insert at will each gene for behavior trend, making possible synthesis as well as analysis.”383 The effect the coat color handle concept was to lay a hand on the problem of complex hereditary relations, to make it tangible, workable. The animal became medium in his hands by the power of this new conception, as eugenicists had always desired and as breeders had always known. The gene, too, became less an abstract entity and more of a physical tool to be put to work synthesizing new combinations of traits. Sensory understanding and experience, thus, appealed not only to the artisan in Keeler, but also the eugenicist.

Keeler used the concept of an allelic handle again later in the year when he coined the term “gene therapy.” The idea was essentially an elaboration of his concept of

“lifting out” or “inserting” cascades of related reactions by the use of a coat color handle.

He proposed permanently eliminating hereditary diseases “by a homozygous

382 Keeler, C.E. Coat Color, Physique, and Temperament. p.271. 383 Keeler to Yerkes December 17, 1946. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. YUL.

185 modification of the genetic formula.”384 There was no real discussion of what such modification would entail, but given the state of induced gene modification in 1947,

Keeler would have been limited to selective mating. The idea of combining and substituting whole trees of reactions could result in the synthesis of entirely new trait complexes. This, he thought, constituted real manipulation. While he cautioned that “at present we may not envisage the application of such gene therapy to medico-genetic diseases in man,” “gene therapy” was clearly an attempt to bring the idea of trait synthesis to the broader field of medicine and closer to human health.

Alan Gregg had been keenly interested in the human implications of Keeler’s researches since he read his 1942 monograph on the taming and morphological effects of the non-agouti allele. Upon reading the report, Gregg wrote to C.C. Little of the Jackson

Memorial Laboratory, “I have been watching for some time to see the emergence of some one who will work as a geneticist on temperament or behavior characteristics in animals and especially in animals whose environments can be more accurately controlled and more reliably recorded than can be the case for human beings.”385 At this time Gregg envisioned a definitive experiment that he hoped would demonstrate that intelligence was primarily heritably determined, rather than fostered by education. He described the project to Robert Yerkes: “If it were possible to develop a line of small dogs of conspicuous intelligence and if the laboratory methods of obtaining intelligence were to prove more reliable than the empirical fumbling of dog breeders, who are usually more interested in shape, color, and other morphological characteristics, we should have a

384 Keeler, C.E. Gene Therapy. Journal of Heredity. 1947. Vol.38(10): 294. 385 Gregg to Little February 9, 1943. RG 1.2, Series 200A, box 127, Folder1126. RAC.

186 demonstration of the rule of the inheritance of intelligence, which would carry interest and possibly conviction to a very large number of persons.”386 It is likely that Gregg had

Keeler in mind for such a project when he began vetting Keeler in 1943. However, mixed reviews from Frank Hanson and C.C. Little, who both mentioned the Wistar controversy, kept Gregg at bay for a number of years.387 A key concern for both men seemed to be Keeler’s interpersonal relations. Wrote Little: “I know Keeler well. He is an able, energetic, and rather difficult person…I had considered Keeler for a place at the

[Jackson] Laboratory here, but was a little afraid as to how he would work with others.”388 He recommended instead the more affable John Paul Scott, whom Gregg ultimately selected to lead the study on dogs at Bar Harbor.389

At the commencement of the intelligence study Scott organized a Conference on

Genetics and Social Behavior at the Jackson Memorial Laboratory, which brought

386 Gregg to Yerkes February 24, 1944. Robert Mearns Yerkes Papers. Series I, Box 22, Folder 399. YUL. 387 Hanson to Gregg February 24, 1943. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. Also Little to Gregg February 17, 1943. Same folder. 388 Ibid. 389 Gregg was disappointed with the findings that Scott and his colleague John Fuller produced from their Rockefeller-funded study on the inheritance of intelligence in dogs. Scott and Fuller found significant environmental effects, even among breed differences, and little evidence of heritable intelligence patterns. See Paul, D.B. The Politics of Heredity. p.71-75.and Dewsbury, D.A. Origins of Behavior Genetics: The Role of the Jackson Laboratory. Behavior Genetics. 2009. Vol.39:3. Scott and Keeler were some of the very few graduate trained geneticists in the country with an interest in behavior in the late 1930s and early 1940s. Scott urged Robert Cook to expedite the publication of his paper “Genetic Differences in the Social Behavior of Inbred Strains of Mice” for the reason that “Dr. Keeler is now doing some work along the same lines with rats.” See Scott to Cook January 12, 1941 Robert C. Cook Papers 1882-1992. Box 13, Folder 3, Library of Congress. When he was installed at Bar Harbor, however, Scott did offer Keeler lab space to continue his studies on pigment gene pleiotropy: Keeler to Yerkes December 17, 1946. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. YUL. Interestingly, Scott’s former mentor, Sewall Wright, instructed his pupil to take his investigation of social behavior further: “You do not refer to other characters- color, morphology, etc. I think that experiments should be planned so as to bring to light clearly any possible correlations… There is a possibility that correlations of great interest [could] be established which would be of great assistance in the analysis of behavior.” Wright to Scott November 11, 1946. Sewall Wright Papers MS.Coll. 60, Folder “Scott, J. Paul 1935-1965,” Series I, APS.

187 together the small but growing community of researchers at the nexus of heredity and behavior. Keeler was invited to present new analyses of the rat work. Gregg and Robert

Morrison of the Rockefeller Foundation, which bankrolled the conference, were also in attendance.

Upon their long delayed meeting, Gregg (apparently assuaged of Keeler’s character) impressed upon Keeler the likelihood that the Rockefeller Foundation would fund his research should he apply. As was the etiquette for soliciting Foundation money,

Keeler had a colleague, James Stokes, Chairman of the Division of Natural Sciences and

Mathematics at Georgia State College for Women, sponsor his application. Stokes’ plea came, not merely on behalf of Keeler or the college, but the entire South, emphasizing the challenges facing the region: “I have never known intimately a man of greater energies, creative imagination, and scientific ability than Dr. Keeler. The South does not have any talent that it can afford to waste and in not utilizing Dr. Keeler’s talent for research in

Medical Genetics we are wasting valuable human resources.”390 Gregg responded with a positive impression of Keeler, but delayed a decision on funding until he could assess

Keeler’s situation in Georgia. The College was in some ways a disappointment.

Reported Gregg, “The Georgia State College for Women has no funds for research work, and both [Keeler] and [Stokes] showed wry delight when I told them that I would ask

President Guy H. Wells how much was the college budget for research work.”391 The nearby Central State Hospital, the state’s only dedicated mental health facility, was understaffed and reflected the racial disparities of the region. However, with a stable

390 Stokes to Gregg November 13, 1946. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. 391 Gregg, A. Memorandum of Interview with Clyde Keeler and James Stokes at Milledgeville, Georgia. February 19, 1947. RG 1.2, Series 200, Box 127, Folder 1126, RAC.

188 population of 9,000 patients it offered the human material that both Keeler and Gregg assumed would be the future application of Keeler’s work. Gregg reported to the

Rockefeller Foundation: “the impression received from Milledgeville was somewhat less depressing than I had expected,”392 before deciding to award Keeler a modest grant of

$10,000 over five years.

With funding from Rockefeller and the implicit support of the Georgia State

College for Women, Keeler was prepared to commence with original studies to demonstrate the association of coat color genes with behavior and various morphological traits. He had only to choose which approach would best demonstrate the putative phenomenon. While still at the Wistar Institute, Keeler had planned to continue to study coat color mutants of the Norway rat. He wished to perform more outcrosses on the coat color varieties derived from the second generation of the Wistar albino to wild gray cross.

He also planned to continue his survey of tame albino strains in use around the country for latent non-agouti and piebald and to await the appearance of more tame, black

“mutants” from King’s nearly inbred captive gray population. Theodosius Dobzhansky wrote in praise of this plan, “I believe that your approach to the problem is absolutely right. If a mutation taking place in a genetically uniform strain produces multiple effects, this is evidence that these effects are due to a single gene. If we could observe the same mutation to arise in the same strain repeatedly and find that every time it produces the same complex of effects, the evidence would be conclusive.”393 Keeler, though, did not have the luxury of carrying out his planned investigations at the Wistar. After his

392 Gregg, A. Report to Rockefeller Foundation February 19, 1947. RG 1.2, Series 200, Box 127, Folder 1126, RAC. 393 Quoted from Keeler, C.E. Behavior Synthesis. p.374.

189 untimely departure, Keeler published only one paper bearing the results of new experiments on rats, and those experiments were begun at the Lankenau Institute prior to

Keeler leaving the Wistar.394 He offered several new correlations of coat color and morphology based on analyses of Donaldson’s leftover data, but no new experimental results. It is difficult to determine whether or not he had access to the strains of rats upon which he had formulated his theory. Keeler continued to keep rats for much of his remaining career for use in research and pedagogy, but it is unlikely that he would have been able to acquire examples of King’s captive grays, with their homogeneous residual hereditary backgrounds, or the coat color mutants derived from them.

Rather than proceed with numerous rounds of outcrossing or “extraction” in the rat, Keeler decided to build the case for pigment gene pleiotropy by demonstrating its generality among mammals. This decision was, of course, predicated on the survey of breeders, fanciers, and trainers he had conducted after being released by the Wistar

Institute. Now with some financial and institutional security, Keeler chose a new subject, the American mink, for the next phase of his research.

In some ways the mink suggested itself for the analysis of pigment gene pleiotropy. It belonged to an entirely separate order of mammals than the rats Keeler had studied previously. Moreover, mink were known to occur in multiple coat colors on commercial ranches and were famously savage if taken from the wild. They had been brought into captivity relatively recently (after 1900) to ensure a steady supply of pelts for the luxury apparel market. In this time numerous hitherto unknown mutations had appeared and were preserved by vigilant and enterprising breeders.

394 Shopbach, R.R., Keeler, C.E., and Greenberg, H.A. Some Variations in Basal Metabolic Levels of Rats. Growth. 1943. Vol.7:83-93.

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Keeler believed that if the common assumption was true and domestication involved the slow (and often unconscious) selection for minute differences in behavior, then, less than 50 years after being confined, the temperament of mink would remain essentially unchanged. If distinct differences in behavior were found among coat color mutants, then he would have confirmation that pigment and temperament were generally related in mammals. In other ways this was a decision of convenience. Keeler had been in contact with a number of mink breeders for the previous five years, including Larry

Moore of Saumico, Wisconsin, who co-authored a paper on mink pelage mutations with

William Castle in 1946. By the late 1940s, the scale and sophistication of some mink operations, like the Moore Mink Ranch, created an enticing opportunity for geneticists like Castle and Keeler. Large ranches such as these usually maintained a number of coat color varieties with sufficient numbers of each to allow for statistically meaningful analyses of their heredity. They also maintained meticulous breeding records and pedigrees.

For Keeler, the prospect of housing, maintaining, and breeding thousands of animals and carefully documenting their genealogies was an impossibility, with or without Rockefeller support. By relenting some control over the operations of his experiment to professional breeders, he could reap the benefits of their size and investment while incurring almost no expense of his own. The risk, and perhaps the cost, of this scheme came in terms of credibility. Breeders were not, of course, trained either in genetics or behavioral analysis. Keeler, though, relied to a considerable extent on their judgment as well as their material and records.

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At least by 1946 Keeler had begun work on “mutation mink” in cooperation with the Moore Mink Ranch, perhaps facilitated again by his former mentor, Castle. Their arrangement was thus: the Moore Ranch would operate according to its predetermined breeding and business plans; Keeler requested varieties of mink, whose carcasses would be sent to him via refrigerated storage after being pelted (the carcass was marked to identify the coat color of the living animal); five experienced breeders at the ranch then provided Keeler with descriptions of the mink varieties as to their “nervousness” or

“gentleness”; finally Keeler dissected the carcasses and weighed the glands and organs he thought relevant to the study, including the brain, adrenal glands, ovaries, and testes.

Keeler had initially planned for a more quantitative measure of behavior. Photoelectric cells, installed in the cages of various coat color mutants, measured the instances of shadows produced by moving mink passing between a light source and the cell. This record of activity was taken as a gauge of the degree of viciousness (according to Keeler, the two were correlated). The “electronic eyes,” however, “required constant supervision at the Wisconsin Ranch,” and workers there could not be expected to devote themselves to these machines.395 Keeler instead relied on the judgment of breeders on the ranch, emphasizing that each breeder had at least ten years experience with the mink varieties in question.396

The Moore Mink Ranch maintained a policy of breeding all new mutant varieties to the standard Dark mink (essentially indistinguishable from wild type mink) to ensure vigor and then extracting the mutants again in the F2 generation. The Dark mink was

395 Keeler, C.E. Research Report to the Rockefeller Foundation June 28, 1954. p.7. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. 396 Keeler, C.E. and Moore, L. Synthesis of Psychosomatic behavior Trends in the Taming of Mink. Bulletin of the Georgia Academy of Science. 1961. Vol.19(3):67.

192 used in Keeler’s studies as a “control,” or the presumed progenitor, against the mutant varieties Silverblu (maltese blue), Pastel (chocolate), Kohinur (also Black Cross- dilution with spotting), Blufrost (also Breath of Spring- pale bluish dilution), Ambergold,

Palomino, Aleutian, Black-eyed White, and Moyle Buff. Although the Moore Ranch housed at least 20 mutant varieties of mink in the dozen years that Keeler worked in association with them, he was primarily limited to these 8 color mutants because of their marketability. Keeler first looked for correlations between color mutations and differences in body size or weight. He found that “[in] the records all genes producing larger than normal size continued to do so (Moyle Buff, Silverblu, and Pastel). All genes having a small size effect continued to produce a small size effect (Breath of Spring,

Aleutian, Black Cross, Palomino, and Ambergold).”397 Again, Keeler’s experimental design was compromised by the need of the ranch to produce ever-larger mink. A general selection for larger size was applied to all marketable varieties, though Keeler maintained that the positive and negative size trends of the color alleles persisted despite this pressure. These trends, which either increased or reduced the length and weight of the mink, were presented as opposed vectors, which gave the impression that the effects of coat color genes on morphology were additive as in Keeler’s description of Norway rats. Data from coat color hybrids showed the combinatorial effect of these genes, though, the extent of their combined effects was not always in line with expectations.

Keeler had the idea that mink coat color genes had quantitative effects on morphology, but was ill equipped to demonstrate this fact beyond simple two-factor statistical analyses.

397 Keeler. C.E. Detection and Interaction of Body Size Factors Among Ranch-bred Mink. Bulletin of the Georgia Academy of Science. 1961. Vol.19(3):38.

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The work on mink, however, did not primarily concern morphology. Rather,

Keeler was interested in showing that the once wild mink had been rendered tame in little more than a generation’s time and that the appearance of certain coat color alleles was to blame. Recalling the source of his discovery, Keeler described his work on the mink: “A twelve year study involving 108,717 ranch bred mink was undertaken to discover the factors determining domestication, because mink are in the process of being domesticated, a term that includes tameness and docility. Domestication implies a radical alteration, on a genetic basis, of that machinery the end-product of which is behavior.”398

Keeler’s survey of breeders and caretakers on the Moore Ranch yielded the experienced opinions that Pastel and Silverblu mink were consistently “gentle,” while

Aleutians and Sapphires were “nervous.”399 He also reported that the mutation Silverblu platinum had appeared on two separate occasions on mink ranches. In both instances, the animals and their progeny were renown for their docility.400 Black-eyed White mink were a relatively recent addition to the Moore Ranch. As was customary, the new mutant was bred to Dark and the F1 progeny were then backcrossed to the Black-eyed White parent. The result was a near 1:1 ratio of Black-eyed White to Dark Hybrid. Keeler, as he was accustomed, relayed his firsthand experience with the progeny of the backcross:

Everywhere in the animal shed Black-eyed Whites were sleeping in the cages. No Darks were observed, so the question was asked, ‘Where are the Darks?’ The reply was that the odor of strangers had awakened them and they were hiding in the nest boxes. The hundred-odd missing Dark Hybrid mink were found when all the nest boxes were opened, but not a

398 Keeler and Moore Synthesis of Psychosomatic Behavior Trends. p.66. 399 Keeler explained that Sapphire was breeding type that was homozygous for both Aleutian and Sliverblu, the Aleutian locus evidently having an epistatic effect on behavior over Silverblu. Ibid. p.68. 400 Keeler, C.E. The Minute Structure of Hair and Correlations with Morphology, Physiology, and Behavior. Journal of the Tennessee Academy of Science. 1951. Vol.26(2):108.

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single Black-eyed White that had taken refuge in the nest boxes was discovered.401

He hoped also to show the intermediate physiological links between coat color and behavior. To accomplish this he dissected and weighed the adrenal glands and gonads of 1,125 mink of various coat colors. What he found agreed with the data Henry

Donaldson had mined from coat color strains of rat more than 30 years earlier. More docile mink tended to have adrenal glands and gonads that were smaller relative to their body weight than did Dark, wild mink. Given the different nature of mink coat color mutations, these findings were not directly transferable to rats, but Keeler felt that he had proven his point. Pigment gene pleiotropy applied as well to carnivores as it had to rodents.

His emphasis on the adrenal gland was mediated by a new hypothesis about the physiology underlying pigment gene pleiotropy. As Dunn had suggested, pleiotropy was notoriously difficult to establish, especially when lacking a clear mechanism for the multiple effects of a single gene. Keeler had the nebulous idea that coat color was somehow related to the complex interaction of nervous tissue, sensory organs, and the ductless glands. Donaldson’s data revealed that all of these structures were modified by domestication, which, in Keeler’s interpretation, meant that they were modified by coat color genes. In a letter to the Journal of Heredity, A.J. Lea, Surgeon Lieutenant of the

British Royal Navy, offered that the relatively recent proposal that melanin and epinephrine both originate from 3,4- dihydroxyphenylalanine (DOPA) suggested a

401 Keeler and Moore Synthesis of Psychosomatic Behavior Trends. p.68.

195 biochemical explanation for Keeler’s pigment gene pleiotropy.402 Although Lea suggested that this explanation precluded pleiotropy on the grounds that the gene product was not itself causing both the color and behavioral traits, the concept of a cascade of related biochemical reactions was perfectly in line with Keeler’s definition of pleiotropy.

Shortly after his exchange with Lea, Keeler’s explanations of pigment gene pleiotropy focused more closely on the adrenal and the metabolism from tyrosine to melanin and the catecholamines: epinephrine and norepinephrine.403

With financial backing from the Rockefeller Foundation and a minor travel grant from the Genetic Society of America, Keeler was able to present some of his preliminary findings in the mink as well as his more extensive data on the Norway rat at both the 8th

International Congress of Genetics in Stockholm and the 13th International Zoological

Conference in Paris in 1948. Keeler’s enormous exhibit, 15 feet long and festooned with ceramic rat models fashioned by the presenter, drew considerable attention at the

Stockholm meeting. It was the sole exhibit described by Miloslav Demerec in his synopsis of the Congress in Science.404 Keeler reported back to the Rockefeller

Foundation that at the Congresses he had met with at least five other researchers who claimed to have evidence of pigment gene pleiotropy in their own experimental

402 Lea, J.A. Biochemistry and Temperament in the Agouti Rat. Journal of Heredity. 1943. Vol.34(7):212. 403 Keeler was induced to study the adrenal more closely by his own results as well. His 1943 paper with Shopbach and Greenberg ruled out hypothyroidism as the cause of the black mutant’s docility. See Shopbach, R.R., Keeler, C.E., and Greenberg, H.A. Some Variations in Basal Metabolic Levels. p.83- 93. 404 Demerec, M. Eighth International Congress of Genetics. Science. 1948 Vol.108(2801):250.

196 materials.405 He also faced criticism at the meeting, relating to the difficulty of demonstrating pleiotropy. In his recollection:

Adhering to the ‘one gene- one function’ idea, [my critics] denied that the physical modifications of glands, sense organs, and brain could possibly be pleiotropic manifestations of the black pigment-producing gene itself, but contended that these alterations must be due to a second mutant gene closely linked on the same chromosome with the black-producing gene. They did concede, however, that if one caught an adult, black sport among wild Gray Norway rats, and if that sport was found to be tame, then this would be a case of pleiotropy.406

Just over a year after this challenge was leveled at Keeler, his network of practical informants provided him with what he considered definitive evidence of pigment gene pleiotropy. Harvey Clarke, an exterminator who worked in central Georgia, brought to

Keeler a black Norway rat he had captured near a barn on the sprawling Central State

Hospital campus. The mutant was found amid dozens of common gray Norway rats while Clarke was fumigating burrows beside the barn. This rat, he noticed, failed to put on the vicious display typical of gray rats that have been captured while trying to escape the fumes. Clarke remembered his earlier contact with Keeler and delivered the rat to him that evening. Some handling was required to situate the rat to a new cage, and it displayed nervous behaviors, but never attempted to bite Keeler. Several days later, when Keeler felt the effects of the poison had passed out of the rat’s system, he ventured to stroke the captive rodent. Again, it showed fear but relented to contact without biting.

Inside of a week it permitted extensive handling. Keeler and Clarke returned to the barn and captured a few dozen rats, among them were two adult black mutants, both reportedly tame.

405 Keeler, C.E. Report of Researches July 14, 1949. p.1. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. 406 Keeler, C.E. The Minute Structure of Hair. p.107.

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Keeler considered these mutant rats to be “predomesticated,” prepared by their genetic constitutions to bear the stress of captivity and human contact. Furthermore, he argued, they highlighted the unlikelihood that two distinct, but linked genes would mutate simultaneously with consistent results not only in King’s inbred grays, but also in the wild in central Georgia. Two years later, Kurt Becker of the Robert Koch Institute reported a similar tendency to tame behavior among the 146 black mutant Norways he captured in the greater Berlin area.407

Keeler also carried on a number of minor studies and observations related to his survey of pigment gene effects on behavior, morphology and physiology at the same time as his work on mink. These included a number of studies on birds, which generally recalled the impressions of Charles Otis Whitman on “degenerate” features in color mutations in pigeons. In 1948 Keeler coauthored a paper with a turkey breeder and a professor of Poultry Husbandry at the University of Georgia concerning observations that a “faded-bronze” mutation in turkeys induced dramatically smaller size, lesser viability of poults, fragility of bones, and rough plumage. The study was limited to a prohibitively small number of observations, in part, because shipping faded-bronze poults from the ranch in Pennsylvania to the University of Georgia exacerbated their already low chances of survival.408 Keeler later included these observations with others collected on rheas, parakeets, ducks, rice birds, and mockingbirds from the ornithological literature.409 The bulk of these were impressions of mutants in the albino series, which were portrayed,

407 Becker, K. Über das Vorkomen Schwarzer Wanderratten. Der Zoologische Garten. 1952. Vol.19:223-233. 408 Keeler, C.E., Hoffman, E., and Shearer, R.K. Faded Bronze Plumage: An Autosomal Mutant in the Turkey. Poultry Science. 1949. Vol.28(5):633-635. 409 Lee, F. and Keeler, C.E. Pigment Variations and Their Correlates in Birds. Auk. 1951. Vol.68:80-85.

198 almost invariably, as tame, small, and feeble. At the suggestion of Alan Gregg, Keeler also evaluated the association of plumage color and behavior in bisque and albino laughing doves and reported a clear correlation between the two traits.410 Keeler even reported finding evidence of pigment gene associated behavior in albino paradise fish in some preliminary trials. The effort to establish generality, however, was not strictly about broadening the reach of the phenomenon. It had at the same time a pointedly specific goal and telos. “When it became evident that we were dealing with a general vertebrate phenomenon,” Keeler wrote, “it was decided to search for possible correlation in man as well as in mink.”411

Before committing himself solely to the human subject, Keeler carried out another nearly decade-long study on mammals. Following his work on mink, he looked for an

“animal of high intelligence possessing more ways of expressing behavior than in rats and mink.”412 In his later reflections, Keeler portrayed this strategy as a stepwise approach toward the discovery of pigment gene pleiotropy in humans.413 He thought first of dogs, echoing Alan Gregg that these intensively selected and thoroughly inbred animals displayed marked differences in morphology and behavior. Dogs, though, were already domesticated and would not show the dramatic alteration of temperament

410 Gregg mentioned differences in behavior between varieties of pigeons known as “archangel” and “carneux,” especially the irritability and defensiveness of archangels. See Gregg to Keeler December 20, 1948. RG 1.2, Series 200A, Box 127, Folder 1126, RAC. Keeler then relayed his experience with laughing doves to Gregg: Keeler to Gregg February 20, 1950 Same folder. 411 Keeler, C.E. Research Report to the Rockefeller Foundation June 28, 1954. p.3. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. It was, significantly, the psychiatrist and sociologist Jacob L. Moreno who convinced Keeler to develop the case for generality rather than becoming more technical in studies of the Norway rat. See Keeler to Cook December 30, 1944 Robert C. Cook Papers 1882-1992. Box 13, Folder 3, Library of Congress. 412 Keeler, C.E. Coat Color Gene Synthesis of Tame Behavior. p.20 413 Keeler, C.E. The Gene Hunter. p.7.

199 associated with the appearance of a relative few single gene substitutions. Instead he chose farm foxes. These animals, like mink, had been brought into captivity only recently, displayed several mutations of the coat, and were raised in large numbers by fur breeders, who would absorb most of the costs of the study.

Keeler again arranged with a commercial breeder, Edward Fromm of Hamburg,

Wisconsin, who had responded to Keeler’s inquiry that he was aware of behavioral differences in the various mutant variants on his farm.414 Shortly thereafter Keeler was invited to the farm to evaluate the animals. Fromm instructed Keeler to enter the open air priming range alone, still unknown to the foxes, and to approach their feeding trough.

Keeler later recounted:

When I arrived with my entourage of foxes at the feeding station I made myself small by getting down on one knee, and this seemed to relieve somewhat the tension of my hosts. I held my wrist to my mouth and made a series of mousy squeaks. At this an interested crowd began to close in on me from every direction. All the foxes near me were Ambers with an occasional Glacier. Farther away I could recognize a Pearl or a Silver and in the far distance a few Platinums. Red foxes never show themselves like this, although they sometimes find their way into the range.415

Keeler was impressed by the arrangement of these foxes, a gradient of coat colors according to their aversion to him, a stranger. “This must be another case of behavior synthesis leading to tame behavior,” he wrote, “We must investigate it.”416 His cooperation with Fromm was similar to the relationship he maintained with Larry Moore

414 Fromm to Keeler December 17, 1962. Clyde E. Keeler Papers Series 1, Box 1, Folder, FF21. GCSU. It is unclear how Keeler became acquainted with Fromm. It may have simply been reputation, as Fromm was one of the largest fox breeders in the United States at the time. Keeler, though, claimed to remember being positioned in the same exhibition space as the Fromm Brothers Fox Farm at the 1933 World’s Fair in Chicago, where Keeler displayed rabbit pelts in connection with his work at the Howe Laboratory and Bussey Institution. See Keeler, C.E. The Gene Hunter. p.99. 415 Keeler, C.E. Behavior Synthesis p.205. 416 Ibid.

200 during the mink project: Fromm provided the pelted carcasses of farm foxes, from which

Keeler would remove the adrenal glands and pituitaries. Again adrenal weight relative to body weight was highest in the wild type animal, the red fox, which bore no coat color mutations. Relative adrenal weight declined progressively for Silver (non-agouti black with silver modifiers), Pearl (non-agouti black and blue), Amber (black, blue, and chocolate), and Glacier (black, blue, chocolate, and white) foxes, that is, in inverse relation to the number of mutations. Relative pituitary weight declined in a similar pattern, though the effect was generally not statistically significant.417 These results agreed with those Keeler had found more than twenty years earlier in rats and a decade prior in mink.

With Fromm’s foxes, though, Keeler was able to go further in his experimental manipulations than he had with either the rat or the mink, despite the fox’s relatively slow reproduction cycle and development. Thanks to Fromm’s willingness (and perhaps curiosity), Keeler was able to perform at least one crucial test cross, which mated a tame

Amber to a wild red fox. The hybrid was then mated back to the Amber parent. This backcross yielded 8 color morphs as Keeler had predicted (4 were previously unknown) and allowed him to conduct behavioral tests on animals with known genetic constitutions, at least of the selected coat color genes. He measured their “startle distance,” or the least distance allowed between a researcher and the fox before the animal fled, and “run distance” or the distance the fox covered after being startled before turning back to view the researcher. These distances were generally less in animals homozygous for recessive mutant alleles at two or more loci than for those with fewer homozygous recessive loci.

417 Keeler, C., Ridgway, S., Lipscomb, L., and Fromm, E. The Genetics of Adrenal Size and Tameness in Colorphase Foxes. Journal of Heredity. 1968. Vol.59(1):82-84.

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These results were similar but not perfectly congruent with Keeler’s earlier observations of the non-experimental farm foxes. He ascribed these differences to the fact that all of the foxes from this experimental backcross were at least heterozygously recessive at each coat color locus. Thus, the foxes homozygous at one locus or none might still have displayed the mitigating effects of recessive coat color mutations in simplex (i.e. incomplete dominance). Protein bound iodine in the blood also increased in a stepwise fashion for animals of 1, 2, and 3 coat color mutations, respectively. Keeler took this to mean that the thyroids of foxes with heavier mutation loads were less active or efficient in processing the protein bound iodine to thyroxin. Mutant foxes also carried higher loads of catecholamine derivatives in their urine, which Keeler explained as a tendency to metabolize and excrete the “fight-or-flight” hormones epinephrine and norepinephrine.418

As Keeler transitioned from mink to foxes, he met with challenges from the administration of Georgia State College for Women, newly named in 1961, The

Women’s College of Georgia. According to Keeler the College President, Robert E. Lee, had opposed faculty research since his election to office in 1956. The following year

Keeler was denied a promotion to the Chair of the Biology Department. In 1961, after 13 years at the College, Keeler was offered a partial contract for three months, to teach chemistry rather than biology.419 Following this insult, Keeler sought refuge at the nearby Central State Hospital where he was named Director of the Research Division.

His new surroundings, at the nation’s largest mental health institution, suggested to him new ways of investigating pigment gene pleiotropy, including the use of psychomimetic

418 Keeler, C., Mellinger, T., Fromm, E., and Wade, L. Melanin, Adrenalin, and the Legacy of Fear. Journal of Heredity. 1970. Vol.61(2):81-88. 419 Keeler, C.E. The Gene Hunter. p.200.

202 drugs to mitigate the anxiety and “wildness” of red foxes. This turn in his research also suggests the facility with which Keeler analogized his experimental animals to his intended subject, humans. Keeler soon described the wild red fox as an ideal model for semi-specific human mental diseases, including schizophrenia and other psychoses.420

Keeler and his associates successfully reduced the red fox’s excessive fear in proportion to the dosage of fear reducing drugs, like Oxazepam. This treatment brought the behavior of the red fox in line with that of its tame coat color derivatives, a parallel that

Keeler emphasized to suggest the applicability of pigment gene pleiotropy to the field of mental health: “The gradational removal of fear by means of increased dosages of psychomimetic drugs is one of the triumphs of modern medicine, but a stepwise reduction in fear and anxiety response associated with the adding together of Mendelian genes has been little explored.”421 By uncovering the principle at work in domesticated animals that made them less fearful in captivity, Keeler believed he also had a means for the permanent mitigation of the anxieties and other mental diseases afflicting modern humans.

Applying Pigment Gene Pleiotropy to the Human Species

Human mental health, for Keeler, was primarily a social problem. This belief, in conjunction with his desire to make a medical genetic discovery of broad and lasting impact, drove his research on pigment gene pleiotropy in domesticated and semi- domesticated animals. Thus, he felt comfortable following a more than half-century-old

420 Keeler, C.E and Mellinger, T. Wild Foxes for Testing Psychoses Before and After Tranquilizer Treatment. Excerpta Medica. 1966. Vol.117:252. 421 Keeler, C., Mellinger, T., Fromm, E., and Wade, L. Melanin, Adrenalin, and the Legacy of Fear. p.81.

203 notion from anthropology that suggested that civilized and social humans were the counterparts of domesticated animals. Robert Yerkes, who later urged caution upon

Keeler, may have stoked Keeler’s imagination in this direction in their early correspondence: “…[A]ssuming your conclusions apply to man, you are pointing a way to the modification of human nature, whose chief shortcomings are at present, many of us suspect, temperamental.”422 Keeler showed a strong willingness to speculate along this line in a later letter to Yerkes, beginning with the bald assumption:

Most of the genes in primitive man are dominant to those in more civilized men, although we have no real wild tribes with which to compare. It may be then that civilization rides forward on recessive mutant genes.

But if one may bear with this fantastic notion, we may even go behind civilization itself. Civilization requires that a species become level headed and sluggish. It must quit jittering and scampering away in fear, but it must take time to secure its food supply, its armament, its organization of society, its division of labor. Just this sort of thing seems to have happened to the Cinnamon rat. Apparently it is a bigger and better rat in development, and it does not suffer from fear so much as the Gray Norway from which it is derived.423

Keeler revisited these themes in a paper published in the journal Sociometry during his sojourn at Wesleyan College, but though he was willing to put these “fantastic notions” in print, he knew that the leap from domesticated animal to civilized human was more easily made in the mind than demonstrated in the laboratory.424 However likely the correlation between pigment and anatomy, physiology, emotionality, or intelligence,

Keeler knew he would have to reveal it against the formidable heterogeneous hereditary background of modern human groups.

422 Yerkes to Keeler September 27, 1941. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. Manuscripts and Archives, YUL. 423 Keeler to Yerkes October 8, 1941. Ibid. 424 Keeler, C.E. Gene Determined Physical Modifications. p.30-41.

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When his nomadic trip south ended with stable employment in Milledgeville, he began planning for a project to demonstrate pigment gene pleiotropy in human populations. In a December 1946 letter to Yerkes, Keeler suggested using a paired control design on siblings or an inbred population to reduce the residual hereditary backgrounds of his subjects. He prepared his proposal in earnest just months after Alan

Gregg solicited his application for a Rockefeller Foundation grant. As he considered using mink to further his experimental work on pigment gene pleiotropy, he also began thinking of the ideal human groups with which to demonstrate the correlation of color, morphology, and behavior. Keeler detailed his plans to study pigment gene pleiotropy in humans as part of another grant proposal (coauthored with the anthropologist David B.

Stout), this time for Axel Wenner-Gren’s Viking Fund in 1948. The most likely candidates, he determined, were an albino sub-group of the Cuna Indians of Panama’s

San Blas Islands and red heads from a somewhat inbred population in Denmark.

The suggestion to investigate red heads for correlations between their characteristic hair tint and other physical and mental traits recalled for many the diffuse cultural wisdom that redheads are extraordinarily passionate and short-tempered. The title of an Atlanta Journal article on Keeler’s proposed work assumed familiarity with this notion: “Are Redheads Really Hotheads? Georgia Scientist Studies Hair of Mink and

Man for Answer.”425 Of course, Keeler had selected red heads for exactly this reputation.

The study was, from the first, an evaluation of widespread cultural wisdom as well as an attempt to demonstrate pigment gene pleiotropy within a human group. Keeler began by first surveying the impressions of men and women whose experience included numerous

425 Fanning, Oden “Are Redheads Really Hotheads? Georgia Scientist Studies Hair of Mink and Man for Answer.” Atlanta Journal. May 7, 1948.

205 observations of red heads, much as he had done prior to the work on mink. Among the dimensions that Keeler hoped to study were red heads’ famously irascible temperament, their reaction to anesthetics, propensity for tuberculosis infection, and excessive bleeding.

Keeler consulted psychological and physiological reports, physicians, anesthesiologists, prison wardens, military officers, mothers of red heads, and red heads themselves. These sources, for the most part, confirmed the impression that red heads exhibit extreme passions and problems associated with the blood, as might have been expected from the familiarity of these ideas in the popular consciousness. Keeler was willing to admit that the common reputation about red heads had inspired him to study their behavior, but he maintained that these banal ideas could not yet be given from the status of “scientific” knowledge:

Curiously enough, [the] probable associations of capillary phenomena with physiology and behavior are in line with the age-old, popular conception of red heads, which may turn out to have some basis in fact. It is desirable that observations be made in this direction, but they must be made on a strictly scientific basis, and this is a rather difficult thing to do. However, such scientific researches are not impossible, and I believe that in the near future they will reveal true associations of morphology, physiology, and behavior associated with hair colors in man.426

Keeler’s description made it seem that the connection between the cultural wisdom regarding red heads and his motivation to study them was coincidental rather than fundamental. He knew well the risk of imbibing too many notions uncritically, against which his mentor William Castle had often warned. Ironically Keeler reasserted the distinction between scientific and practical, traditional, lay, or other ways of knowing at a point in his career when he relied to a substantial degree on the impressions of men

426 Keeler, C.E. The Minute Structure of Hair. p.110.

206 and women who did not belong to the community of scientists. Practical and scientific challenges ultimately forced Keeler to abandon the project. As late as 1964 he included a request to readers of the Milledgeville (Central) State Hospital Bulletin of Current

Research that they send him any data they might have from autopsies performed on redheads. He hypothesized that their adrenal glands would be larger on average than those of blond, brown, and black haired individuals. The reason for the request, he explained, was that “we rarely have a redheaded autopsy and we do not have any reference to any statistics on the subject in the literature.”427 Even if he had procured data himself, perhaps from the Danish populations he once considered, he felt the residual hereditary backgrounds of these groups were still too varied despite their relative stability. Keeler terminated his red head study having conducted only preliminary survey work and publishing only proposed plans.

His research on the incidence and correlated effects of albinism among the Cuna

Indians of San Blas was much more extensive and deliberate. Families and populations with high rates of albinism had become well known to geneticists at least since the pedigree analyses of Gertrude and Charles Davenport in 1910 revealed stable inheritance of the trait.428 That Keeler was the first geneticist to plan a substantial study of the Cuna is not surprising in light of the relatively few anthropological surveys completed on them prior to 1940 and the difficulties in reaching the San Blas. Keeler initially planned to carry out his genetic studies in conjunction with David Stout who had completed his

Ph.D. dissertation on the ethnology of the Cuna in 1946. This collaboration fell through,

427 Keeler, C.E. Request for Autopsy Records. Milledgeville State Hospital Bulletin of Current Research Experimental and Clinical. 1964. Vol.3(1):16. 428 Davenport, G.C. and Davenport, C.B. Heredity of Skin Pigment in Man II. American Naturalist. 1910. Vol.44(528):705-731.

207 however, after their failure to secure support from the Viking Fund in 1949.429 Keeler instead planned for a solo trip to the San Blas the following summer, drawing from his

Rockefeller grant. He ultimately made 22 trips to the islands to collect data, genetic and otherwise, on the Cuna.

One of the first challenges Keeler faced was determining the rate of albinism among the Cuna. Previous estimates by Stout, Harris, and others varied significantly, though all showed an unusually high incidence of albinism. These rates were difficult to determine accurately as Cuna populations were divided into many villages, some nearly inaccessible, some notoriously hostile to outsiders. Moon Children, as albinos were known among the Cuna (according to the belief that their mothers, while pregnant, gazed at the moon), had also inspired a number of cultural customs and practices, ranging from ostracism to reverence, which may have frustrated efforts at estimating their numbers.

Keeler ultimately estimated that Moon Children occurred at a general rate of 0.67% of the Cuna population versus an estimated rate of 0.0059% albinos in the general human population, a more than 100 fold increase. This, he declared, was the highest rate of albinism in the world.430

Keeler assumed that inbreeding was responsible for the high incidence of albinism in the Cuna. It also made them ideal candidates for genetic study, given their ostensibly homogeneous hereditary backgrounds. Of course, Keeler could not be sure of the amount of genetic variation in the Cuna population. They had been visited by

429 Keeler thought that Stout would submit a grant application with the Viking Fund in late 1948 or early 1949, but he later learned that Stout had not submitted the application nor had he informed Keeler that he was no longer on the faculty at Syracuse University. This event may have soured relations between the men. See Paul Frejas to Keeler September 13, 1949. Clyde E. Keeler Papers Series 9, Box 10, in Scrapbook 2. GCSU. 430 Keeler, C.E. The Incidence of Cuna Moon-Child Albinos. Journal of Heredity. 1964. Vol.55(3):115.

208 numerous interlopers in the past, from 16th century Spaniards to post-Canal Panamanians.

Keeler, though, was assuaged by their persistent suspicion of outsiders. He heard tales of past occupations, which ended in the systematic slaughter of the foreigners and their

“hybrid” children.431 Keeler ingratiated himself to the Cuna by helping to formulate a phonetically based written language for them. This side project was the beginning of

Keeler’s deep immersion in Cuna culture, which was as much for his personal interest as it was to facilitate his genetic studies.432

The first of Keeler’s reports on the Cuna relied primarily on impressions from his brief visit in the summer of 1950. Moon Child height and weight was generally below that of normally pigmented Cuna of the same age. As might have been expected from prior knowledge about albinos, Moon Children suffered disproportionately from premature wrinkling, skin lesions, and cancers of the skin. Villagers reported that Moon

Children were slow to develop sexually, remained physically weak throughout life, and were chronically ill. They also carried a reputation for being stubborn, diligent though physically weak, and occasionally explosive in temper. Their faces appeared incapable of complex expression. Keeler noted no difference in their intellectual abilities versus pigmented Cuna. He also confirmed that Moon Child albinism, like other forms studied, was most likely caused by a single recessive allele. Keeler followed up these impressions a decade later with a much more sophisticated and intensive study of Moon Child

431 Keeler, C.E. The Caribe Cuna Moon-Child and its Heredity. Journal of Heredity. 1953. Vol.44(5):163. 432 Keeler’s interest in Cuna culture grew to include their art, mythology, folklore, and history. His amateur anthropological efforts on the Cuna were spent in the largely discredited field of epigraphy. He hypothesized that pre-Columbian explorers from the Old World had seeded the Cuna’s ancestors and those of other Native American groups with basic religious and cultural elements. See Keeler, C.E. Secrets of the Cuna Earthmother: A Comparative Study of Ancient Religions. New York: Exposition Press, 1960.

209 heredity. The latter project involved twenty-two principal researchers, including physicians, psychiatrists, a psychologist, an optometrist, a dentist, a biochemist, and an oncologists drawn mostly from the staff of the Central State Hospital where Keeler had been named Director of Research in 1961. Six albino males, paired with normally pigmented males of the same age, were brought to Central State Hospital in Milledgeville for testing. Keeler and Donald Moyer, a dermatologist, selected the individuals because their family histories were relatively complete, and presumably something could be said of their heredity. Extensive anthropometric measures confirmed Keeler’s earlier impressions about the smaller size of the albinos. Moon Child bones were more gracile, and their facial structure differed in a number of slight ways. The new tests leant some credibility to Keeler’s observations, but failed to yield many new distinctions between pigmented and albino Cuna. Blood tests showed few differences, save for a high level of protein bound iodine, as was found in tame foxes. Moon Child teeth were in all respects indistinguishable from those of their pigmented tribesmen. Psychological testing revealed no likely differences in intelligence, as Keeler had noted. The mental health professionals on the research team agreed that the Moon Children behaved differently in a number of ways than pigmented Cuna, but they could not distinguish between the effects of heredity and those of culture bound beliefs, customs, myths, and practices concerning the albinos among the Cuna, so intimately were these factors bound.

Keeler’s interest in the Cuna was never limited solely to their heredity. As he had done with mice more than two decades earlier, he found relevance in the history, mythology, and other cultural meaning surrounding the Moon Children. His work, on the border of human physical and mental health and heredity, to some degree demanded a

210 broader look at the importance of color inheritance. With the Cuna Moon Children,

Keeler thought that he had not only found evidence of pigment gene pleiotropy among humans, but also a model for how a society dealt with a hereditary anomaly, alternative to the Western democratic model.

In an article for Eugenical News, Keeler provided a brief and possibly apocryphal history of the Moon Children of the Cuna. One of the Cuna’s great cultural heroes and perhaps its first chief, Nele Sipu, decreed that “all men are brothers,” which Keeler took to mean that Moon Children were to be equal members in Cuna society. Sipu was reportedly a Moon Child himself. Keeler, though, seemed to doubt that equity had ever been fully realized, despite the Moon Children’s privileged place in some Cuna ceremonies. Moon Child infanticide was still common at the time of Keeler’s visit, accompanied by the explanation: “The midwives felt sorry for Sipu.” The Cuna’s aversion to foreigners was hardened by their brutal treatment at Spanish hands, dating possibly back to Columbus’ landings. In small, secluded, and, indeed, inbred populations, albinism flourished despite the customs of infanticide and restrictions on

Moon Child marriage. “Enlightenment,” nevertheless came, and the practice of infanticide was discouraged, especially in the larger villages. Keeler seemed to lament the demise of this eugenic safeguard: “The indications are that unless the present tendencies are reversed the Moon-Child incidence will continue to increase among the

Caribe-Cuna Indians.”433

The Cuna illustrated well the cross-influence of history, culture, and heredity.

Keeler seemed to appreciate the depth of this interaction, though, his experience with the

433 Keeler, C.E. An Attempt to Eliminate a Genetic Syndrome in Man. Eugenical News. 1950. 35(3):44.

211

“primitive” Cuna brought to the fore the frustrations he had with norms and trends in his own culture, ones he shared with a number of latter day eugenicists. In his later years,

Keeler worried openly about the effects (especially eugenical) of the government welfare programs enacted in the middle 20th century.434 In a letter to Robert Cook, Keeler expressed, through the contrast of primitive and civilized, animal and man, his discomfort with the dysgenic course of modern civilization, leading to the tacit question of whether modern mores and practices were an improvement over the primitive:

Remember my demonstration on rats, mink, and foxes that temperament and fear also involve genes that can be directed by pleiotropic tags to such simple traits as coat color. This means that we can select genetically for calmness in a maddening world…

The ancients exposed the defective neonates on the mountain tops, naively expecting the gods to make the decision. The Cunas left it to the father to tell the midwives to bury in the sand the newborn if it turned out to be an albino. And with the Panamanians considering infanticide a crime today the already highest albino rate in San Blas is increasing enormously.

Genetics has made enormous strides in Agriculture and Animal Husbandry because here we can play dictator… If and when we ever change our social system and our thinking to permit these practices with human genetics, we shall improve the human race… But whenever gonadectomy has been suggested in the legislative halls for inferior individuals, the legislators begin to feel pain in their groins. The price of species Homo improvement is too great for the average individual to make voluntarily.435

As Keeler moved to establish pigment gene pleiotropy in a human group he was made to understand that the social implications of his findings were not strictly under his control. Alan Gregg, upon reading Keeler’s plans for work on the Cuna, warned, “There are plenty of people who would jump at the conclusion that your findings scientifically

434 Keeler, C.E. undated. “Notes on History” Clyde E. Keeler Papers Series 7, Box 5. GCSU. 435 Keeler to Cook February 28, 1978. Robert C. Cook Papers 1882-1992. Library of Congress.

212 supported race distinctions on the basis of color, and, thanks to the present and recent political and social tensions, a misinterpretation of that kind may prove to be a hornet’s nest for you.”436 Keeler replied that he had been concerned with this possibility from his earliest findings with rats, and his anxiety had only grown since moving to the South. He was not possessed of the notion that skin color in humans defined any discrete types. He agreed with Gregg that modern human groups were too heterogeneous to display sharp behavioral and physiological distinctions in correlation to the color of their skin, regardless of whether pigment genes carried pleiotropic effects or not.437 He further clarified in published papers that his studies on albinism in an inbred population could not be extrapolated to substantiate or explain race differences.

Nevertheless, his admonitions had to contend with deep-seated cultural notions about skin color and race distinctions, many of which had been granted scientific legitimacy over the past century. That pigmentation translated to real race differences was, in some biological circles, axiomatic around the turn of the 20th century. The author of Colour in Nature: A Study in Biology was, thus, able to write: “…[E]ven slight differences in intensity of pigmentation may correspond to relatively vast constitutional differences… [For instance] the difference in hair-colour corresponds to all those profound mental and moral differences which separate Celt from Saxon. That the mental and moral differences are associated with physical ones hardly needs proof to the biologist.”438 Throughout the 1920s and 1930s knowledge of quantitative traits, those determined by the combined effects of a number of genes, complicated scientific support

436 Gregg to Keeler August 16, 1950. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. 437 Keeler to Gregg September 12, 1950. Ibid. 438 Newbigin, M.I. Colour in Nature: A Study in Biology. London: J. Murray Company, 1898. p.289-290.

213 for such basic sociological assumptions. Keeler’s pigment gene pleiotropy, however, seemed a viable alternative to this interpretation. By invoking the multiple effects of a single, simply inherited, recessive allele, culturally familiar associations of color, physiognomy, and behavior (i.e. stereotyped race distinctions) could be maintained with support extending to the biochemical level. Barbara Burks’ history of pigment gene pleiotropy included the tacit suggestion that race was never far out of mind even as “the association of color of coat and color of personality” was being evaluated in the humble rat. “Most picturesque…” she recorded among her descriptions of rat personalities, “was the contribution of a Negro attendant, Edward Tyler: re the blacks, ‘Those were the laziest things I ever saw in all my life. They just didn’t care.’”439 No geneticist, psychologist, or anthropologist, however, used Keeler’s ideas or data to attempt to legitimate race distinctions. Keeler was, as he had promised Gregg, extraordinarily cautious to prevent the misunderstanding that his research supported such conclusions.

Sensitivity to issues of race was heightened in the years following Keeler’s announcement of pigment gene pleiotropy by the revelation of ethnically targeted atrocities in foreign theaters of war and greater acknowledgment of racial tensions domestically (Keeler relayed his own experience with a fear campaign perpetrated by the

Ku Klux Klan in Milledgeville in a letter to Gregg in the fall of 1948).440 These factors contributed to the rise of reformist eugenics, and a consequent departure from racially tinged mainline eugenics. The shift of focus and leadership in the American eugenics movement may be associated with the lack of racial commentary on Keeler’s theory.

439 Burks, B.S. Temperament and Single Gene Substitutions. p.6. 440 Keeler to Gregg December 6, 1948. RG 1.2, Series 200A, Box 127, Folder 1126, RAC.

214

Although Keeler entertained the idea that tendencies in personality were associated with red hair and albinism, he was not willing to use pigment gene pleiotropy to compare or define racial types. Nor, it seemed, was anyone else.

Conclusion

Clyde Keeler remained convinced, in over forty years of proselytizing, that someday pigment gene pleiotropy would be rediscovered and finally recognized for its import to the study of evolution, psychology, and sociology.441 What had happened, however, to the idea that had inspired so much interest and speculation early in the 1940s, among psychologists and sociologists as well as biologists, by the 1980s, when Keeler seemed instead like the last priest of a forgotten mythology? Surely Keeler’s ardor had not waned. He rarely missed the opportunity to mention his theory in articles, lectures, books, and letters, even where its pertinence was questionable. His enthusiasm, however, failed to inspire other biologists or psychologists to investigate the putative phenomenon.

This failure had less to do with Keeler’s enthusiasm than his credibility amid shifting priorities in the sciences and socio-political movements to which he believed his theory pertained.

In the first years after Keeler presented his findings on mutant rats, they were greeted with substantial interest, publicity, and praise. Press reports buoyed his initial enthusiasm for the idea of pigment gene pleiotropy. These were able to simultaneously emphasize the novelty as well as the vague familiarity of coat color associated form and

441 Keeler made some effort to convince historian of science Gar Allen of the historical and present importance of pigment gene pleiotropy to sociological, psychological, and evolutionary theory. Keeler to Allen December 15, 1981 and January 8, 1982. Personal contact with the recipient.

215 behavior to the at-large public. “The old belief that redheads and blondes are different because of the color of their hair received an astonishing scientific explanation today…” wrote the Associated Press’ Science Editor Howard W. Blakeslee following Keeler and

King’s presentation at the Dallas meeting of the A.A.A.S. in 1941, “It is a shock to find that [coat color] genes actually govern characters in rats, and by implication may do something of the same sort in men and women.”442 Later reports covering Keeler’s work on mink, foxes, and humans repeated the same impressions, suggesting that his interpretation of color’s influence on temperament never gained currency among the public but was consistently recognized for its relation to popular wisdom. Scientists of many stripes joined the press in the initial fillip for Keeler’s findings.

Keeler had considerable early support from prominent figures in the psychological and social sciences, fields that he imagined would benefit more from his discovery than his native biology. Karl Lashley announced to Keeler, “It seems to me that you have one of the most important discoveries for psychology that has turned up in a decade,” echoing the slightly more qualified praise of his antecedent at the primate laboratory at Orange Park, Robert Yerkes. Similar approbation came from Barbara Burks and Earnest Hooton. The majority of Keeler’s supporters in the social sciences found common cause in the eugenical implications of pigment gene pleiotropy and a strongly hereditarian view of social problems and their solutions.

Keeler noted, in retirement, that in the course of building the case for his thesis a shift had occurred within the psychological community such that: “my psychological proponents were no longer in power and nobody in the psychological field paid any

442 Blakeslee, H.W. “Science Links Hair Tints, Temperament”

216 attention.” His thesis faced mounting criticism from a younger generation of psychologists, many who reasserted the importance of the environment and social interaction in determining personality and temperament. These criticisms were joined by results that apparently contradicted Keeler’s own from others under the umbrella of

“psychobiology.”

J.P. Scott published a paper on social behavior in black and agouti strains of mice less than a month after Keeler and King’s presentation in Dallas. The paper designated the black and agouti strains as “pacifists” and “aggressors,” respectively, in accordance with Keeler’s findings. Scott, though, was interested in social behaviors rather than the reactions of the mice to humans as Keeler had emphasized (Keeler’s preoccupation with explaining the “marks of domestication” caused him to focus more on human-animal interactions than reactions to conspecifics). In social encounters, the pacifist black strain appeared more curious, but was not aggressive unless attacked, then it was usually dominant in fights.443 Keeler had also noted that coat color strains of rat differed in aggressiveness when engaged with other rats versus being held by a caretaker. Benson

Ginsburg and W.C. Allee corroborated many of Scott’s findings (using the same strains of mice, C57 black and C-3H agouti), particularly that black mice were dominant to agouti mice in combat. They also noted, however, that in nonsystematic experiences in handling these mice, the black strain was more pacific in human hands.444 The point that aggressiveness or social dominance may differ between conspecific and heterospecific interactions was not elaborated in either article and may have left the impression that

443 Scott, J.P. Genetic Differences in the Social Behavior of Inbred Strains of Mice. Journal of Heredity. 1942. Vol.33(1):11-15. 444 Ginsburg, B. and Allee, W.C. Some Effects of Conditioning on Social Dominance and Subordination in Inbred Strains of Mice. Physiological Zoology. 1942. Vol.15(4):501.

217

Keeler had simply mischaracterized the effect of the nonagouti allele. Ginsburg also suggested in another paper that the tame behavior that Keeler noted in his agouti rats could be the result of handling rather than genetics.445 In 1953 William R. Thompson, studying behavior in some of the same mouse strains as Scott, Ginsburg, and Allee, noted that in open-field tests of food drive, emotionality (measured by rate of defecation), and exploratory drive, “[c]ontrary to the observations of Keeler… there was little indication that coat-colour genes might usefully serve as markers for functional traits.”446 Peter

Broadhurst, a psychologist at the University of London, found that open-field defecation rates did not support the view that black and albino rats were more “sluggish” than their agouti counterparts. Although Keeler himself noted,447 and Calvin Stone corroborated,448 that defecation rate was not a suitable criterion for distinguishing domesticated from wild strains, these findings helped to undermine Keeler’s thesis, especially in the field of comparative psychology.

As Keeler complained about the attitude of the newer psychologists toward his thesis, he also grew frustrated with the critiques from his colleagues in genetics. He railed particularly against the assumption that correlated but disparate traits must be the result of linkage rather than pleiotropy. In a synopsis of his life’s work, written late in his career, Keeler moved to “sum up quod erat demonstrandum: in rats, mink, foxes, and man anatomical, physiological and behavioristic alterations accompany hair color gene

445 Ginsburg, B.E. All Mice are not Created Equal: Recent Findings on Genes and Behavior. The Social Service Review. 1966. Vol.40(2):126. 446 Thompson, W.R. The Inheritance of Behaviour: Behavioural Differences in Fifteen Mouse Strains. Canadian Journal of Psychology. 1953. Vol.7(4):151.

447 Keeler, C.E. The Association of the Black (non-agouti) Gene with Behavior. p.383. 448 Stone, C.P. Wildness and Savageness in Rats of Different Strains. In K.S. Lashley Studies in the Dynamics of Behavior. Chicago: University of Chicago Press, 1932.

218 mutations. It is not ‘one gene-one effect’ as many believe. The initial step may be ‘one gene-one enzyme’ that initiates a complex tree of chemical reactions that affect anatomy, physiology and behavior through alterations in glands, sense organs and brain.” His approach hinged on the unlikelihood of docile behavior and anatomical and physiological marks of domestication co-segregating with coat color alleles in rats, mink, and foxes if these traits were in fact only linked and not derived from the multiple actions of a single gene. Other geneticists suggested that a better approach would involve repeated outcrossing to dispel the linkage hypothesis, or locating and evaluating new black sports from the same inbred, captive gray population. Both of these plans required Keeler to continue using captive gray rats with minimal variation in the hereditary backgrounds upon which coat color differences were displayed, that is, King’s rats. After Keeler’s ouster from the Wistar Institute, this was hardly practicable.

Keeler’s effort to generalize pigment gene pleiotropy in mink and foxes (while consciously ascending a scale of mental complexity with humans at the zenith), though, was not simply a matter of necessity. By the late 1940s, he had a long history of interaction with breeders and fanciers, beginning with his exposure as a student at the

Bussey Institute. It is difficult to determine precisely the influence that these interactions had on Keeler’s thinking and the course of his career. It is clear, however, that Keeler often found diverse strains of knowledge relevant to the understanding of genetics, such as the deep historical and cultural emphases woven into his 1931 book The Laboratory

Mouse. He was candid about his relation to the fancy, more so than many of his fellow

Bussey alumni. Even before he had formulated the idea of pigment gene pleiotropy (with the help of Wistar breeder and caretaker Ruth Meeser), his contacts with the breeders’

219 world helped him contribute to the establishment of two cat breeds, the Burmese and

Himalayan, and eventually resulted in no less than 9 papers on the genetics and breeding of cats, most coauthored with members of the fancy. Keeler ultimately published 13 academic papers with breeders listed among the investigators. Employment and funding troubles only drew him into a closer exchange with breeders, as he plumbed their experience and wisdom for further examples of color associated morphology and behavior. He reported one successful contact in a letter to his old friend and colleague

Emilia Vicari: “’An old fancier’ who trained mice for show purposes, declared, ‘white, lavender, and fawn mice are easiest to train, then piebalds, and lastly black, but these are very dull in learning.’ Sometimes, the old boys had something.”449

In 1954 Keeler was at work on The New Laboratory Mouse, a revision of his earlier work. This was more than a new edition, however. Keeler conceded that other books already covered the latest scientific knowledge on mice. The new book, though, was not intended for biologists, instead he declared, “We have slanted it toward the fancier who raises mice for the pleasure of it.”450 In the book, Keeler underscored the complementary interests of breeders and geneticists and suggested how these could lead to their mutual benefit: “To the researcher, I would say, ‘Don’t throw away a strain when you are through with it, that could possibly interest the fancier’ To the fancier, conversely, ‘Don’t discard an animal because it is a freak. A freak may be a mutant.

449 Keeler to Vicari no date. Clyde E. Keeler Papers Series 9, Box 10, in Scarpbook 1. GCSU. 450 Keeler, C.E. The New Laboratory Mouse (typescript). Fan Du Lac, Wisconsin: All-Pets Books, Inc., 1954. p.3. (never published). In Clyde E. Keeler Papers Series 9, Box 10, in Scrapbook 1. GCSU.

220

And mutants are materials upon which embryology, physiology, genetics and other specialties of biomedical sciences feed.”451

Keeler’s concept of pigment gene pleiotropy bore a resemblance to breeders’ beliefs about the importance of coat color as a marker for latent and seemingly unrelated traits and broader popular wisdom concerning the relation of color and temperament, a familiarity that Keeler’s listeners consistently recognized. His elaborations on the idea, including the concepts of the “coat color handle” and “gene therapy” made coat color genes appear like tangible entities, which medical geneticists could insert or extract at will for the benefit of a patient or, more accurately, of the society at large. The Swedish human geneticist Carl A. Larsen recognized the selective power suggested by Keeler’s view of the coat color gene: “The possibility to attack behavior, normal, near normal, and outright abnormal, with the tools of genetic models, becomes indeed more promising when behavior variants have, as in the Norway rats and Moon-child albinos, a color tag.”452 Eugenicists had used the analogy to the breeder’s practice to justify a selectionist social agenda for decades; the utility of coat color markers for mental traits was not lost on them.

As Keeler assumed closer ties with breeders the reform movement within

American eugenics drew closer to the core of medical genetics work including genealogical analysis of specific pathologies, utilizing animal models for human diseases, and working with actual patients in hospitals and mental health clinics to understand the interaction of hereditary factors of disease with environmental, social, and other factors.

451 Ibid. Section: “Roll Call of the Mutations” (not paginated). 452 Quoted from Keeler, C.E. The Gene Hunter. p.247.

221

Outside of this trend and with multiple studies ongoing, Keeler’s pigment gene pleiotropy appeared insufficiently nuanced to reveal anything of value about the association of behavior and constitution. Near the end of Keeler’s Rockefeller Foundation grant, one of the Foundation’s administrators, Robert Morrison, judged:

I must confess that against the background of the genetics conference which I am currently attending at Cold Spring Harbor, K’s remarks about a correlation of temperament with various hereditary characteristics such as skin color seemed rather loose and uncritical. The trip to Panama this summer is for the purpose of studying the temperament of the many albinos among the San Blas Indians but it does not seem to me that K. will have the equipment or the time to make any but very superficial observations. I would guess that the recent developments in the field of human genetics will increasingly make K. look rather less outstanding than he did ten years ago.453

Keeler, too, returned to the core of medical genetics late in his career, conducting studies on Klinefelter’s and Down syndromes and finally in an attempt to demonstrate that the rodless retina mouse he had discovered in 1924 was identical to the strain that came into use as a model for retinal degeneration three decades later. He was ultimately successful in this last goal, publishing with a team of geneticists and ophthalmologists almost 70 years after his first publication in the same journal, the Proceedings of the

National Academy of Sciences.454 Through the demands of this work and his amateur anthropological studies, however, Keeler remained convinced that his greatest discovery, pigment gene pleiotropy, had simply been neglected or misunderstood by the scientific

453 Morrison, R.S. Interview with Dr. Clyde Keeler June 14, 1950. RG 1.2, Series 200A, Box 127, Folder 1126. RAC. 454 Pittler, S.J., Keeler, C.E., Sidman, R.L., and Baehr, W. PCR Analysis of 70 Year-old Sections of Rodless Retina Demonstrates Identity with Mouse RD Defect. Proceedings of the National Academy of Sciences. 1993. Vol.90(20):9616-9619.

222 community. He imagined that someone would rediscover the phenomenon and validate his work on the subject, as had happened with his investigations of rodless retina mice.

Pigmentation has become increasingly important to the understanding of animal behavior since Keeler retired in 1975, and the view of its field of possible effects has become increasingly complex. The contemporary field of color-associated behavior is beyond the scope of this historical account, though some of these recent studies have touched directly on the subject of pigment gene pleiotropy, often without reference to

Keeler’s work on the subject. Celia Cottle and Edward Price set out to evaluate the conflicting reports of Keeler and his detractor Broadhurst on agouti and non-agouti

Norway rats. Their findings supported both investigators, that is, “[d]ifferences between agouti and black rats in ease of handling support the findings of Keeler (1942) and the lack of significant differences in open-field ambulation and defecation are in agreement with Broadhurst (1958, 1960).”455 Cottle and Price supported the view that color was relevant when the rats were interacting with humans, but left the question of pigmentation’s effects on conspecific or other interactions with nonhuman animals open.

Virginia Hayssen and her associates extended the studies of Cottle and Price to deer mice

(Peromyscus maniculatus). They determined that non-agouti deer mice were less aggressive, less active, and easier to handle than their agouti counterparts. They attributed these temperamental distinctions to differential regulation of melanocyte- stimulating hormones (MSH). Specifically, they suggested that MSH binding to melanocortin receptors on neurons might have been inhibited by the antagonistic action

455 Cottle, C.A. and Price, E.O. Effects of the Nonagouti Pelage-color Allele on the Behavior of Captive Wild Norway Rats (Rattus norvegicus). Journal of Comparative Psychology. 1987. Vol.101(4):393.

223 of the agouti protein product.456 In this hypothesis α-MSH, which mediates melanin production, could also be connected to adrenal function through the action of its progenitor protein, adrenocorticotropic hormone, which stimulates cortisol production as well as the pathways that produce both epinephrine and melanin. Hayssen referenced

Keeler’s original joint paper with King, but failed to cite his later studies or development of a neuro-endocrine mechanism for the pleiotropic action of coat color genes, albeit nebulous by comparison to her own. What seems indelible in this narrow line of work, however, is the recognition of likeness to the traditional ideas of breeders, the past arbiters of hereditary knowledge. Hayssen wrote in 1997, apparently without fear of seeming naïve or uncritical, “Animal breeders have long understood that selection on the basis of coat color is often associated with subsequent differences in temperament and behavior. The behavioral differences between agouti and nonagouti deer mice and their similarity to the differences between agouti and nonagouti rats may help explain decades of anecdotal wisdom.”457

Keeler, by the middle 1940s, had already suffered somewhat from the impression that he uncritically imbibed “old wives’ tales.” His own ideas were undoubtedly influenced by his sustained interaction with breeders and fanciers and sometimes from the wider field of cultural wisdom. He had been prepared for this relationship by his training at the Bussey Institute, where, alongside Castle’s insistence on physiological

456 Hayssen, V. Effects of Nonagouti Coat-color Allele on Behavior of Deer Mice (Peromyscus maniculatus): a Comparison with Norway Rats (Rattus norvegicus). Journal of Comparative Psychology. 1997. Vol.111(4):419-423. See also Hayssen, V., Gunawardhana, S., and Meyer, J. The Agouti Locus May Influence Brain Catecholamines: Regional Differences in Norepinephrine and Dopamine in the Brains of Two Colormorphs of Deer Mice (Peromyscus maniculatus). Comparative Biochemistry and Physiology. 1994. Vol.107C:51-55. 457 Hayssen, V. Effects of Nonagouti Coat-color Allele. p.423.

224 explanations of genetic phenomena and epistemic purity in science, he also trained in the breeding and care of whole colonies of small mammals and participated alongside breeders in fancy shows. Practical needs, for employment, research material, and funding drew Keeler into closer association with breeders, especially between the early to middle

1940s. The war years were uncertain for many scientists, but doubly so for Keeler who also had to contend with the reputation of a difficult temperament after his bitter conflict with Helen Dean King. Breeders, fanciers, farriers, cattlemen, and others with daily experience selecting, mating, and caring for animals gave Keeler the opportunity to continue to amass evidence, albeit anecdotal, in support of his theory of associated pigment and personality. When he achieved some stability, in employment and funding, he chose to pursue some of the leads he had gathered during these lean years, though with an eye to answering his scientific critics who doubted that his phenomenon was truly pleiotropy. Keeler was pursuing genetic problems, not practical or aesthetic ones. The breeders with whom he interacted probably would not have envisaged the research program that he designed, but their materials, records, participation, and knowledge were ultimately integral to his case for pigment gene pleiotropy. This reliance in combination with Keeler’s high regard for experiential knowledge fostered the impression, sometimes correct, that he could be an uncritical consumer of ideas. Keeler’s damaged credibility, in turn, reflected changing standards of evidence in genetics and growing distinctions between the science and its early associate, breeding. Around the turn of the century it was a maxim for many students of heredity that temperament, including all manner of mental and moral qualities, was co-inherited with color. Even as late as 1954, J. A. Weir reported, “There is a prevalent view, not without evidence, that color and temperament

225 are associated,” though he cautioned, “Study of morphology is important but offers no shortcut to success for the breeder.”458 Keeler contemporaneously hoped that pigment gene pleiotropy would provide exactly such a shortcut, especially for the amelioration of heritable mental diseases among humans. The declining stock of such simple and strongly hereditarian eugenical ideas among geneticists in the 1950s and 1960s, supplanted by more quantitative approaches and a greater appreciation for the role of the environment, made Keeler’s idea seem increasingly irrelevant. Shifting social preoccupations, standards of evidence, research emphases, and techniques have contributed to the striking distinction between modern genetics and the experiential and traditional knowledge of breeders. Such is the distinction that modern geneticists, like

Hayssen, can remark about the surprising validation that the geneticist provides for breeders’ anecdotal ideas about color and temperament with little to no sense of the importance or continuity of their interaction.

458 Weir, J.A. On the Concept of Inherited Constitution. Journal of the Kansas Academy of Science. 1954. Vol.57(2):118.

226

Beasts and the Burden of Metaphor: Domestication in American Psychobiology to the Middle 20th Century.

In the early 1950s Helen Spurway began an extended critique of the thinking about domestication in ethology with a question: “Can wild animals be kept in captivity?”459 She was, however, more interested in the question implied: Does a wild animal in captivity cease to be wild? This question had very live methodological implications for the study of animal behavior, which had struggled to find the proper balance between representing nature and maintaining control since becoming an experimental science around the turn of the 20th century. Spurway argued, though, that the implications extended well beyond method. Domestication, she pointed out, was an ineluctably human activity and any deductions derived from it “about the animal without us should be made in the consciousness that we cannot avoid being actors, however much we would prefer to be an audience.”460 Just as domesticated creatures owed their identities in large part to their interactions with humans, a substantial volume of social

(and indeed scientific) thought contended that the civilized self could not be fully understood without reference to its closest animal companions. With this analogical thinking in mind, Spurway declared, “It is impossible to be objective about domestication

459 Spurway, H. Can Wild Animals be Kept in Captivity? New Biology. 1952. Vol.13:11. I also include as part of this extended review and critique: Idem. Behold, My Child, the Nordic Dog: Review of King Solomon’s Ring by Konrad Lorenz. British Journal for the Philosophy of Science. 1952. Vol.3:265-272; Idem. The Causes of Domestication: An Attempt to Integrate Some Ideas of Konrad Lorenz with Evolutionary Theory. Journal of Genetics. 1955. Vol.53:325-363. Spurway’s husband, J.B.S. Haldane, also commented on the subject in his 1956 Huxley Memorial Lecture: see Haldane, J.B.S. The Argument from Animals to Men: An Examination of its Validity for Anthropology. Journal of the Royal Anthropological Institute of Great Britain and Ireland. 1956. Vol.86(2):1-14, esp.5. That domestication and its implications for man and civilization were a frequent topic in the Haldane/Spurway laboratory may also be seen in the poem “The Insemination of the British Bulldog or the Disadvantages of Domestication by A. David Blest, part of a collection presented to Haldane by his students on Guy Fawkes Day, 1952. In Smith, J.M. Cautionary Tales for Aspiring Species or the Beast’s Book of Blunders. Trends in Ecology and Evolution. 2001. Vol.16(12):719. 460 Ibid. p.28.

227 ecotypes. Darwin (1868) called them deteriorated, Delacour (1951) coarse. These adjectives reflect the Greek contempt for usefulness, the Romantic admiration for the noble savage, and the embarrassment of a Herrenvolk in the contemplation of its

Quislings.”461

Spurway’s claim is striking because it suggests that this spirit of vicarious judgment was so basic to the thinking about domesticated animals that it precluded the modern ideal of scientific objectivity, especially in the evaluation of their behavior. This was not a problem to be solved by greater vigilance; it was a fact to be accepted and acknowledged. It was also a damning indictment for a field that, perhaps more than any other at the dawn of the experimental age, had been self consciously consumed with questions about its claims to objectivity and scientific legitimacy. The study of the mind and its expressions seemed always to be plagued by self-awareness and the normative considerations that accompanied it. Spurway hinted as much in her critique: “the study of behavior, which, as it relies on the observation of inappropriate actions… may be described as the contemplation of sin.”462

The use of animals under tightly controlled conditions seemed to provide the appropriate distance, while evolutionary theory suggested that what was learned about the animal mind could help crack the citadel of human mentality. In the mainstream of

American psychology these animals ultimately came to be seen as model organisms, a means of generalization about intellect and behavior and a bulwark against the

461 Ibid. p.26. Spurway had in mind a Quisling of her own acquaintance, namely the lion of ethology, Konrad Lorenz. Her review, which came in the wake of the Second World War and an extramarital affair with Lorenz, attempted perhaps naively to distinguish between Lorenz’s biological ideas that she deemed “important” and those she regarded as ideologically motivated and “retrograde.” 462 Spurway, H. Behold, My Child. p.272.

228 speculative philosophy that marked psychology’s past. The behaviorist viewpoint, which dominated American animal behavior study for much of the 20th century, identified experimental control with objectivity. It also encouraged generalizations from model organisms, especially the domesticated albino rat, to other animals, principally humans, by deemphasizing the diversity between and within species.

An alternative viewpoint existed, however, from the earliest years of experimental animal behavior study in the United States. Historian of science Cheryl Logan, in the context of the selection and development of the albino rat as a model organism in 20th century psychology, neurology, and physiology, has described this approach as one that

“sought generality from the empirical assessment of diversity,” rather than assume that such generality existed among vertebrates or could be created by the production of a

“standard” organism.463 The late Philip J. Pauly described an allied distinction between engineering and progressive evolutionary ideals in early 20th century biology.464

Following this historical tradition, I identify a “psychobiological” alternative to the behaviorist approach, which at the height of the latter’s hegemony of animal behavior research in the United States was reduced to a minority, albeit significant role. I connect this not to the effort to establish standard animals but to a related problem with similarly pressing practical and philosophical consequences: how researchers judged the impact of

463 Logan, C.A. “[A]re Norway Rats… Things?” p.291. For more on the transition between the emphasis on diversity to the emphasis on generality, see: Idem. The Altered Rationale for the Choice of a Standard Animal. p.3-24; Before There Were Standards: The Role of Test Animals in the Production of Empirical Generality in Physiology. Journal of the History of Biology. 2002. Vol.35(2):329-363; The Legacy of Adolf Meyer’s Comparative Approach: Worcester Rats and the Strange Birth of the Animal Model. Integrated Physiological and Behavioral Science. 2005. Vol.40(4):169-181. 464 Pauly, P.J. Controlling Life.

229 domestication on experimental animals and how they extrapolated their findings to humanity.

There is a danger in generalizing about psychobiology, as the field does not have a precise definition (or, more accurately, it has too many definitions). Robert Yerkes highlighted this ambiguity in a brief exchange with Knight Dunlap in 1934 that focused upon the origin and usage of the term “psychobiology.” In his last letter on the subject,

Yerkes declared with some exasperation, “[T]he combination is so natural that one would suppose that hundreds of persons might have thought of it.”465 The commonalities in psychobiology were not in methodology or disciplinary arrangement, but in point-of- view.

Psychobiologists in the early 20th century viewed the mind, including its functions and expressions, as a product of countless adaptive changes in the phylogenetic history of the species. They emphasized what they believed to be innate behaviors as examples of the adaptive value and evolutionary significance of behavior. These behaviors, in turn, could be used to answer basic biological questions concerning variation, development, heredity, and evolution. They maintained that the mind was inextricably linked to the body and their functional correlates, physiology and behavior, were likewise united.

Theirs was an integrative approach, rejecting strict reductionism as it failed to recognize

465 Yerkes to Dunlap November 2, 1934. Robert Yerkes Papers Series II Subject Files Group No.569, Box 71, Folder 1363, YUL. On this ambiguity, see also, Dewsbury, D.A. “Psychobiology.” American Psychologist. 1991. Vol.46(3):198-205; Joel Elkes described the mercurial nature of psychobiology thus, “It seems that, living thing that the term was, it would change hue and substance as soon as it sensed human presence.” Quoted from Elkes, J. W. Horsley Gantt: Moments from Memory in Critical Issues in Psychology, Psychiatry, and Physiology. Thomas Ban and Frank J. McGuigan (eds.) Monographs in Psychobiology: An Integrative Approach. Vol.2. New York: Gordon and Breach, 1987. p.72.

230 the roles of the whole organism, its context, and history in the formation of behavior.466

They also suggested that the animal should not be subordinated to method or to the question under investigation. Many of these features fall under the slightly less problematic concept of “functionalism” (popularized by the psychologist James Angell and others), and the two terms are related.467 “Psychobiology” is used here in the recognition that the term’s compound nature signals an approach that occupied a middle space between biology and psychology and resists pigeonholing investigators into one or the other discipline.

These psychobiologists found the behaviorist emphases on control and on generality, problematic. The problem was essentially that raised by Spurway’s question: did captivity, control, and dominion- in a word, domestication- fundamentally change the animal, such that generalization about the nature of its behavior became impossible?

While psychobiologists raised important questions about behaviorist assumptions regarding the controlled environment of the laboratory and generalizations about animal subjects, their own attempts to answer these questions revealed a number of assumptions, cultural biases, and normative commitments related to the understanding of domestication. Absent the generality of the behaviorists, psychobiologists frequently relied on analogy, especially that between domestication and civilization, to make the precarious leap from the tightly controlled conditions of the laboratory to the human social world.

466 Dewsbury, D.A. “Psychobiology.” p.203. 467 Indeed, historian of science Ruth Leys used them interchangeably in reference to Adolf Meyer’s programmatic designs for psychology and psychiatry. See Leys, R. Meyer, Watson, and the Dangers of Behaviorism. Journal of the History of the Behavioral Sciences. 1984. Vol.20:130.

231

As historian of science Gregg Mitman has explained, metaphor in science, as elsewhere, is “embedded in cultural experience, and by uncovering metaphor we unveil the social and political context that helped shape its construction. Through metaphor and language generally, scientists introduce terms into their science that are both value laden and context specific.”468 As a proxy for civilization, domestication suggested itself to several psychobiologists in the early 20th century as a means to study the connections between organic and social evolution. As it highlighted the distinction between what was artificial and what natural, often conceived in the normative terms of what is and what ought to be, domestication also raised questions about the appropriateness of human agency- ambition, progress, and intervention- in the world. The analogy thus aroused special questions for an American animal behavior study dominated by the use of a single domesticated animal, the albino Norway rat. Around mid-century, psychobiologists used the construct of domesticates-as-degenerates as more of an internal critique of the conventions of experimental psychology than any wider social commentary. This criticism centered on the ubiquity of the Norway rat and the diminished role of instinct in behaviorist psychology. Throughout the 20th century in American animal behavior study, the conception of the domesticated animal, particularly the Norway rat of the laboratory, occupied a changing but always allegorical role: first as a model of civilization and its discontents and, later, as the embodiment of the excesses of the field.

468 Mitman, G. Evolution as Gospel: William Patten, the Language of Democracy, and the Great War. Isis. 1990. Vol.83(3):448. On the role of metaphor and analogy in science, see: Hesse, M.B. Models and Analogies in Science. Notre Dame, Indiana: University of Notre Dame Press, 1966; Harraway, D. Animal Sociology and Natural Economy of the Body Politic, Part 1: A Political Physiology of Dominance. Signs. 1978. Vol.4:21-36; Stepan, N.L. Race and Gender: The Role of Analogy in Science. Isis. 1986. Vol.77(2):261-277.

232

The Domestication Analogy and Notions of Progress in Pre- 20th Century Social and Scientific Thought

Frederick Cuvier asserted in the opening line of his essay on the study of behavior in domesticated animals that “[t]he strangest prejudices have been found regarding the state of animals in captivity, and the most singular judgment passed upon the works to which their actions have given rise.” He continued, “The origin of these ideas is easily discovered. They proceed from the same source as most of the errors which have been entertained with respect to the nature of animals; the ideas to which the study of man gave rise were applied to these beings.”469 But what were these values and prejudices that lay behind the analogy from domesticated animals to man and how had they found use in social and scientific discourse in the decades prior to the rise of experimental psychology?

By the time it came into use by scientists in the 19th century, this analogy and its normative consequences were, as Nancy Leys Stepan wrote with respect to another scientific trope, already “long-standing, long-familiar, [and] culturally endorsed.”470

Some of the most potent moral judgments arose from the recognition that domesticated animals were distinct from their wild forbears and, thus, might be considered unnatural or artificial. Domesticated animals offered a window into the natural condition of man, if

469 Cuvier, F. Essay on the Domestication of Mammiferous Animals, with Some Introductory Considerations on the Various States in which We May Study their Actions. Edinburgh New Philosophy Journal. 1827. Vol.3:303, a translation of Cuvier’s “Essai sur la domesticité des mammiféres, précédé de considerations sur les divers états des animaux, dans lesquels il nous est possible d’étudier leurs actions,” Memoires du Muséum d’Histoire Naturelle. 1825. Vol.13. Richard W. Burkhardt Jr. explains that the younger Cuvier’s role as garde de la menagerie of the recently established menagerie of the Muséum National d’Histoire Naturelle shaped his opinion of the utility of domesticated animals for illustrating natural phenomena. See Burkhardt Jr., R.W. Constructing the Zoo: Science, Society, and Animal Nature at the Paris Menagerie, 1794-1838, in Animals in Human Histories: The Mirror of Nature and Culture. Mary J. Henninger-Voss (ed.). Rochester: University of Rochester Press, 2002. p.243-244. 470 Stepan, N.L. Race and Gender. p.265.

233 only by contrast to their wild counterparts, as well as an index of the change that was wrought upon his body and mind since leaving the state of nature. The captivity inherent in domestication brought to mind issues of liberty and free will, while the provisions that were made under domestication recalled the rise of the welfare state and the debilitating effects of luxury. Others, including Cuvier, viewed domestication as a model of the material and moral perfectibility of human society. In these years, anthropologists, zoologists, anatomists, psychiatrists, and other emerging scientific experts concerned with demonstrating the social relevance of their science found domestication a superbly adaptable metaphor, in part because it spoke to a central 19th century tension between progress and peril, advance and decline.

The concept of the domesticate-as-degenerate had a relatively deep cultural history, prior to its adoption by would-be social experts in the 19th century. That domestication was somehow connected to the degeneration of type was familiar to practical breeders, who, prior to the 18th century had little reason to expect improvements among their stock. Historian Nicholas Russell notes that “[s]ince deterioration was a such a common experience, one of the most frequently stated breeding objectives was to hold the status quo and prevent degeneration of the stock.” Bitter experience, Russell argues, bolstered Platonic and Christian assertions that bygone generations were nobler and more perfect than their present day descendents. As with the introduction of sin, bringing the mufflon, the aurochs, and the boar out of the wild and into captivity destined their lines to diverge further and further from their original perfection. This dogma persisted even as new breeding strategies made stock improvement a realistic goal. Well into the 20th century the geneticist Jay Lush complained that when “selected parents

234 generally had offspring not as good as themselves [it] was generally regarded as one of the mysteries of heredity or, more discouragingly still, as proof of some kind of automatic and powerful tendency to degenerate and the breeder trying to improve his stock was in some way fighting against an important law of nature.”471

In the culture surrounding animal breeding, degeneration was not merely a phenomenon affecting domestic animals, but was a common motif in the modern agricultural literature. The flowering of Romantic agrarianism around the turn of the 19th century coincided with widespread interest in the question “whence the natural man?”

Together, these conveyed a sense of longing for a pastoral way of life most often idealized and, thus, irrevocable. Nevertheless, popularizers of this view, which included agriculturalists as well as philosophers, physiciens, and physiocrats, claimed that examples of the natural man could still be found, beyond the confines of the city. One such idealist, Zurich physician Hans Caspar Hirzel, argued, “in the country, humanity presents itself to our view, in a state of innocent simplicity, resembling in some degree the state of nature.”472 Romantic agrarianism, though, was defined as much by condemning the city as by exalting the countryside. Its authors self-consciously revived ancient dichotomies between rural virtue and urban vice.473 It was the perceived

471 Lush, J.L. Genetics and Animal Breeding in Genetics in the 20th Century: Essays on the Progress of Genetics During its First 50 Years. L.C Dunn (ed.). New York: MacMillan Company, 1951. p.497. 472 Quoted from Johnstone, P.H. The Rural Socrates. Journal of the History of Ideas. 1944. Vol.5(2):167. Hirzel found his own example of unspoiled humanity in Jacob Gujer (Kleinjogg as he was known to Hirzel’s readers), a paragon of simple virtue. Consistent with Romantic agrarianism more broadly, the lesson and tenor of Hirzel’s Die Wirthschaft eines Philosophischen Bauers were decidedly moral rather than agricultural. As with the wild men and feral children, Klienjogg became something of a cultural phenomenon among elite scholars and statesmen throughout western Europe, including Goethe, the elder Mirabeau, Quensay, Arthur Young, and Prince Louis Eugene of Würtemberg. 473 This contrast was made plain in the writings of Cato, Cicero, Varro, Virgil, Palladius, and perhaps nowhere more forcefully than in the preface to Columella’s De Re Rustica: “[We] have quit the sickle

235 wickedness of the city that inspired fears of urban degeneracy and, in response, the back- to-nature cult of the late 18th century.474 Even for the city’s harshest critics, however, a return to the wilderness was unthinkable. Instead the authors of agrarianism emphasized the moral and emotional benefits of farming because they perceived a lack of these qualities in an increasingly urban-industrial world of which they were reluctantly part.

According to historian David B. Danbom, it was the spiritual fulfillment of agriculture and rural life, its transcendental value, that made pastoralism “an ideology for all seasons, attractive to great varieties of people who [were] called upon to share no more than a degree of alienation from modern society.”475

The Country Life Movement, one of the efforts to organize and act upon this broad sentiment in the first decades of the 20th century, shared with the popular American

Eugenics Movement a nativist concern about urban degeneracy, inspired by declining birthrates among native families and the apparent fertility of the immigrants settling in urban slums. As historian of science Barbara Kimmelman has shown, the ties between these movements were institutional as well as ideological. Leadership of the American

Breeders’ Association (the nation’s first significant body to promote eugenical research)

and the plough and have crept within the city-walls; and we ply our hands in the circuses and theatres rather than in the grainfields and vineyards; and we gaze in astonished admiration at the posturings of effeminate males, because they counterfeit by their womanish motions a sex which nature has denied to man, and deceive the eyes of the spectators. And presently, then, that we may come to our gluttonous feasts in proper fettle, we steam out our daily indigestion in sweat baths, and by drying out the moisture of our bodies we arouse a thirst; we spend our nights in licentiousness and drunkenness, our days in gaming or sleeping, and account ourselves blessed by fortune in that ‘we behold neither the rising of the sun nor its setting.’ The consequence of this is that ill health attends so slothful a manner of living; for the bodies of our young men are so flabby and enervated that death seems likely to make no change in them.” Vol. 1 English translation by H.B. Ash. Loeb Classical Library. Cambridge, Massachusetts: Harvard University Press, 1954. p.14-15. 474 Johnstone, P.H. In Praise of Husbandry. Agricultural History. 1937. Vol.11(2):80-95. 475 Danbom, D.B. Romantic Agrarianism in Twentieth-Century America. Agricultural History. 1991. Vol.65(4):7.

236 overlapped with prominent Country Life supporters, including the leading agricultural scientists Willet M. Hays and Liberty Hyde Bailey.476 Hays expressed the consonance of the hereditary and the transcendental benefits of rural life in his 1912 speech to the

National Farmer’s Congress, ‘The Farm, the Home of the Race’: “We have come to look to the open country as the source of fresh blood with which to keep up the vitality of the race and of the nation’s moral and religious life.”477 Following in the agrarian tradition,

Bailey, elsewhere, heaped scorn on the “parasitic… elaborate and artificial” city.478 The praise of husbandry (and of rural life in general) and its attendant critique of the city provided a familiar backdrop upon which eugencists could launch their own critique of the modern urban-industrial world that seemed so unfamiliar and threatening. With the union of the American Breeder’s Association and the Country Life Movement, agrarianism brought together the romantic vision of the farm with the concrete lessons of heredity provided by the animals raised there. American agrarianism was, thus, drawn more closely into the nexus of scientific theory and social advocacy.

The identification of man as a domesticated animal inspired a raft of commentary on the validity of the comparison and its social implications by the middle 19th century.

“The exuberance of the subject is such that we can give but slight indication of it here,” explained physician Henry Holland in the Quarterly Review, confident that even in his cursory description the argument would “be understood by everyone.”479 Indeed, familiarity was one of the heuristic advantages of the domestication analogy that

476 Kimmelman, B.A. The American Breeders’ Association. p.163-204. See also, Cooke, K.J. The Limits of Heredity. p.269-274. 477 Hays, W.M. The Farm, the Home of the Race. American Breeders’ Magazine. 1913. Vol.4(1):16. 478 Quoted from Danbom, D.B. Romantic Agrarianism. p.3. 479 Holland, H. ‘Natural History of Man.’ Quarterly Review. 1849-50. Vol.86:26.

237 appealed to scientists in the 19th century. It seemed to make plain more complex and controversial arguments about human origins and evolution.

The association of domestication and degeneration appeared, as did the seed of many subsequent ideas in natural history, in the works of Georges Louis Leclerc comte de Buffon. Degeneration, in Buffon’s usage, was nearly synonymous with variation: “… with regard to Nature, improvement and degeneration are the same thing, for they both imply an alteration of original constitution.”480 It described the change wrought upon a plant or animal by incongruities between the organism’s “internal molding force,” developed under one environmental regime, and its newly encountered conditions of life.

Domestication, for Buffon, offered an extraordinary example of this modification:

“If to [the] natural causes of alteration in free animals, we add that of the empire of man over those which he has reduced to slavery, we will be astonished at the degree to which tyranny can degrade and disfigure Nature; we will perceive the marks of slavery, and the imprints of her chains; and we will find that these wounds are deeper, and more incurable in proportion to their antiquity; and that, in the present condition of domestic animals, it is perhaps impossible to restore their primitive form, and those attributes of nature which we have taken from them.”481 The identification of domestication as slavery reflected the opposition to human bondage common among philosophes. The congenital and likely permanent degeneracy seen in domestic animals was akin to the moral debasement of man stripped on his independence and unable to exercise his will.

480 Buffon, G.L.L. Natural History: General and Particular Translated by William Smellie. London: W. Strahan and T. Codell, 1781. Vol.7 p.398; Ibid. p.399. 481 Ibid. p.396.

238

As the historians of science Adrian Desmond and James Moore have sedulously detailed, the moral justification for slavery remained an important subtext for the scientific use of the domestication analogy through the first half of the 19th century. At this time, they argue, “The big human question, Unity or Plurality [of the races], was being made to hang on fancy and farm breeds.”482 The particulars of this debate extend beyond the scope of the present chapter, but it did encourage the reciprocal understanding of domesticated animals and man in 19th century natural history and anthropology, such that “it remained true that one of the best places to pick up contemporary chicken lore was a human-race book.”483 It also helped to establish the moral consequences of the use of the domestication analogy within the sciences and their instantiation in the image of the domesticate-as-degenerate.

Johann Friedrich Blumenbach had been one of the earliest naturalists to argue from the common origin of domesticated varieties to the genealogical unity of the human races: “These instances of diversity… [may] be taken as clear and safe examples of the variations which may be expected to arise in the descendants of one stock… the whole difference between the cranium of the Negro and that of an European, is by no means greater than that equally striking difference which exists between the cranium of the wild boar and that of the domestic swine.”484

But why were domestic animals especially suited to this argument? To make the case for common ancestry, Blumenbach also required mechanisms by which to explain

482 Desmond, A. and Moore, J. Darwin’s Sacred Cause: How a Hatred of Slavery Shaped Darwin’s Views on Human Evolution. New York: Houghton Mifflin Harcourt, 2009. p.225. See also p.199-227. 483 Ibid. p.212. 484 Quoted from Prichard, J.C. Researches into the Physical History of Man. London: John and Arthur Arch, 1813. p.353.

239 the degeneration of the natural or original man into the diversity of modern races. Thus, he made clear that the analogy between domestic animals and man was also an analogy between the processes, domestication and civilization, that produced them. He went further to declare that these processes were one and the same, that domestication, one of man’s earliest arts, was in fact his nature:

The difference between [man] and other domestic animals is only this, that they are not so completely born to domestication as he is, having been created by nature immediately a domestic animal. The exact original wild condition of most of the domestic animals is known. But no one knows the exact wild condition of man. There is none, for nature has so limited him in no wise, but has created him for every mode of life, for every climate, and every sort of aliment… But the consequence of this is that there is no second animal besides him in the creation upon whose solidum vivum so endless a quantity of various stimuli, and therefore so endless a quantity of concurring causes of degeneration, must needs operate.485

This discussion of humans, domestication, and degeneration bore the unmistakable influence of contemporary social debate, in this instance on the nature (and artifice) of man and the consequences of his entry into civilization. Blumenbach derided the fascination and credulity with which prominent Enlightenment figures treated reports of feral children discovered in forests throughout Europe. He illustrated with the example of Wild Peter, a feral boy discovered near Hameln in 1724 who subsequently became the subject of intense scientific scrutiny. Linneaus gave Peter a unique taxonomy, Juvenis hanoveranus; “Buffon, De Pauw, and J.J. Rousseau… exalted him as a specimen of the true natural man.”486 For Rousseau especially, Peter and other feral children exemplified the nobility of man’s unspoiled nature and suggested the

485 Blumenbach, J.F. Beyträge zur Naturgeschichte I in The Anthropological Treatises of Johann Friedrich Blumenbach and The Inaugural Dissertation of John Hunter, M.D., on the Varieties of Man. Translated by Thomas Bendyshe. London: Longman, Green, Longman, Roberts, & Green, 1865. p.294. 486 Ibid. p.331.

240 adulterating influence of civilization, which was antithetical to that nature. Blumenbach doubted both that these children were truly primitive and that they represented any past human condition. He borrowed a counterexample from Voltaire, one of the most vehement critics of Rousseau’s concept of the noble savage: “If one meets with a wandering bee, ought one to conclude that the bee is in a state of pure nature and that those who work in company in the hive have degenerated?”487 According to

Blumenbach, man had never understood his domesticated nature and so wrongly assumed his present condition was unnatural, degraded, and false. He affirmed, “[Man] is born naked and weaponless, furnished with no instinct, entirely dependent on society and education.” Man’s own domestication was never a choice, but his nature.

Blumenbach’s disciples in the argument for unity ranged from the pious and unassuming James Cowles Prichard to the polemical William Lawrence, whose main exposition on the subject was banned in Britain for its materialistic heresies and dangerous Continental influences. As distant as these men were by reputation, both modified portions of Blumenbach’s domestication analogy and expanded its applications to contemporary social concerns.

For instance, where Blumenbach had left a wedge between man, imbued with reason, and domestic beast, beholden to unchanging instinct, Prichard saw continuity in the human and animal mind. He variously described the instincts of domesticated animals as “greatly altered,” “obliterated,” or more often “improved,” and from the presumed likeness of domestication and civilization he predicted that “[a] priori we might expect to discover in the psychological characters of human races changes similar

487 Ibid. p.339 (footnote)

241 in kind [to those of domestic beasts], but infinitely greater in degree.”488 In his Treatise on Insanity, he also asserted that “[t]he brain receives a different development in the progeny of cultivated races,” and while this usually resulted in greater perfection, it also increased the number and variety of mental disorders in civilized nations.489 These opinions arrived as mental illness was becoming an increasingly public concern (by midcentury Britain had dramatically increased its number of public institutions for the insane) and from the pen of one of the early contributors to an emerging field of psychiatry in Britain.

Lawrence, who slavishly reproduced many aspects of Blumenbach’s human domestication argument, disagreed with his Göttingen mentor’s position that man never enjoyed a truly wild and natural condition and had always been in some measure civilized. Instead, Lawrence seized on the artificialities of civilization and used the domestication analogy to suggest the consequences of entering this mode of life. He noted that animals “in a wild state have very few and simple diseases, if any: domesticated animals have many; and they are more numerous in proportion as the

488 Prichard, J.C. The Natural History of Man: Comprising Inquiries into the Modifying Influence of Physical and Moral Agencies on the Different Tribes of the Human Family. London: H. Bailliere, 1843. p.75. Prichard became more convinced of the continued importance of instinct in humans after his longtime friend Thomas Hancock defended instinct against the charge that it was the preserve of brute animals alone and claimed it was actually the more virtuous of the two Providential determinants of behavior, instinct and reason, as it was the more natural and less perverted. As evidence Hancock also relied on an analogy between animal domestication and human civilization: “as man is compressed in society, their instincts like those of the brute, give place to the culture of Reason: while, as man takes his range in the wilds of uncultivated nature, and seeks no other guide for his direction, his senses acquire a perfection which sometimes far exceeds the laboured deductions of reason.” Hancock, T. Essay on Instinct and its Physical and Moral Relations. London: William Phillips, George Yard, 1824. p.54. Hancock seems to have been influenced in this thinking by the moralist John Gregory’s A Comparative View of the State and Faculties of Man with Those of the Animal World (1765). For more on Hancock’s influence on Prichard’s thinking about instinct see: Augstein, H. F. JC Prichard’s Concept of Moral Insanity- a Medical Theory of the Corruption of Human Nature. Medical History. 1996. Vol.40:322-324. 489 Prichard, J.C. A Treatise on Insanity and Other Disorders Affecting the Mind. London: Sherwood, Gilbert, and Piper, 1835. p.351.

242 subjugation is more complete, and the way of life differs more widely from the natural one,” and further suggested, “Perhaps nosological catalogues would afford the most convincing argument that man has departed from the way of life to which nature had destined him.”490 If his point about the pitfalls of modernity was yet unclear, his jeremiad on the same page left little question about how his contemporaries should regard their own degenerations under domestication:

The accumulation of numbers in large cities, the noxious effects of impure air, sedentary habits, and unwholesome employments; the excesses of diet, the luxurious food, the heating drinks, the monstrous mixtures, and the pernicious seasonings, which stimulate and oppress the organs;- the unnatural activity of the great cerebral circulation, excited by the double impulse of our luxurious habits, and the undue mental exertions, of the violent passions which agitate and exhaust us, the anxiety, chagrin, and vexation, from which few entirely escape, and then reacting on and disturbing the whole frame;- the delicacy and sensibility to external influences caused by our heated rooms, warm clothing, inactivity, and other indulgences, are so many fatal proofs that our most grievous ills are our own work, and might be obviated by a more simple and uniform way of life. Our associates of the animal kingdom do not escape the influence of such causes. The mountain shepherd and his dog are equally hardy, and form an instructive contrast with a nervous and hysterical fine lady and her lap dog; the extreme point of degeneracy and imbecility of which each race is susceptible.491

Yet even as Lawrence made clear the power of domestication to corrupt and disfigure, he admitted that in these deviations from nature also “lay the foundations of new breeds,” (i.e. they were also potentially a creative force).492 As he and other scientists sought to define the social meaning of the domestication analogy, their interpretations reflected a tension in the long 19th century between the sense that modern

490 Lawrence, W. Lectures on Physiology, Zoology, and the Natural History of Man London: J. Callow, 1819. p.238-239 491 Ibid. p.239. 492 Ibid. p.304.

243 civilization continued its indomitable progress and growing anxiety that the seeds of its fall had already been sown. As historian Harriet Ritvo has explained, the Victorian emphasis on cultivated civility and ordained social order encouraged the impression that domesticated animals were “perfectible” beings, “reclaimed” from brute nature. The apparently willful subordination of these animals was treated as evidence of the master’s right to rule, a justification for the rigid social order and colonial ambitions of early

Victorian Britain.493 The perfectibility of domestic creatures was thus linked with the upward climb of British industry, empire, and prestige, that is, with social advance. In the latter half of the century, however, gnawing concerns about internal decay undermined the old Victorian self-confidence. Even that greatest of evolutionary idealists, Herbert Spencer, did much hand wringing over the improvidence of the Poor

Laws.494 Despite many signals that civilization advanced apace, there existed a deepening sense that the path upon which modern society had embarked was neither as well-planned nor righteous as previously assumed. Similar doubts characterized social thought in Belle Époque France, Gilded Age America, and post-unification Germany.

This was not a rejection of the central 19th century logic of progress, but its extension. The pessimism brooding over the Progressive Era gave only the appearance of a paradox, for as historian Robert Nye observed, “the concept of degeneration was conceptually inseparable from that of progress.”495 It remained an age of ‘becoming’ rather than ‘being,’ but like the image from thermodynamics of a dimming sun, the fate

493 Ritvo, H. The Animal Estate. p.17-30. 494 Richards, R.J. Darwin and the Emergence of Evolutionary Theories of Mind and Behavior. Chicago: University of Chicago Press, 1987. p.262-263. 495 Nye, R.A. Sociology and Degeneration: The Irony of Progress. In Degeneration: The Dark Side of Progress. J.E. Chamberlin and S.L. Gilman [eds.] New York: Columbia University Press, 1985. p.49. For a similar interpretation, expressed somewhat differently, see Rosenberg, C.E. No Other Gods. p.202.

244 to which Western Europe was trending was increasingly re-imagined as a dissolution rather than an apotheosis. It was in this 19th century that Virginia Woolf employed a purebred spaniel as the title character in Flush to suggest the illusory basis of Victorian class distinctions and in which Nietzsche declared, “He who knows what goes on in menageries, doubts very much whether an animal is improved in such places. It is certainly weakened, it is made less dangerous, and by means of the depressing influence of fear, pain, wounds, and hunger it is converted into a sick animal. And the same holds good of the tamed man whom the priest has ‘improved.’”496

This new cultural mood was not solely the product of artists and iconoclasts, it was substantially mediated by science, especially after the formulation of “degeneration” as a widespread social pathology by the psychiatrist Benedict Morel and his intellectual heirs in the second half of the 19th century. These figures followed an emerging progressive tradition in which designated experts were granted the traditionally cultural or legal responsibility of defining what was normal and what was deviant or, in the new conception, pathological. Despite deliberate efforts to mitigate the influence of external values via statistical methods, scientific nosologies, and clinical detachment, psychiatrists and other experts weighing in on social questions could not escape the cultural resonances of arbitrating what is and what ought to be. For instance, Morel’s dégénérescence recalled the birth of sin and the Christian ascetic ethic of self-denial as well as contemporary anxieties over civil collapse, political radicalism, the frenetic pace of life demanded by industry, and the unceasing stimulus of the city. Thus, for the pious

496 Snaith, A. Of Fanciers, Footnotes, and Facism: Virginia Woolf’s Flush. Modern Fiction Studies. 2002. Vol.48(3):622-625. Nietzsche, F.W. Twilight of the Idols: or How to Philosophize with a Hammer; The Antichrist; Notes to Zarathustra; and Eternal Recurrence. Edited and translated by Oscar Levy. Edinburgh: T.N. Foulis, 1911. p.45.

245

Morel, what ought to be was confined to a pre-Lapsarian past with natural man situated in

God’s bucolic garden, and what is and was to be was represented by the degenerate in a depraved milieu of his own making, which in turn made him.

The normative force of the domestication analogy relied substantially upon the implicit identification of normal with natural and deviant with artificial. Nature, whether interpreted through the lens of Lamarckian equilibrium or the Darwinian struggle for existence, was wise, if not always beneficent. Artifice, or the product of human agency, seemed correspondingly unwise and tinged with more than a shade of guilt. Many who used the domestication analogy recognized that human meddling with nature had reached an unprecedented scale and extent in their time and openly worried about the outcome,

“[w]hen man in his own person, or in the organism he interferes with, so far baulks

Nature’s provisions.”497

The real concern, articulated by Darwin’s old Cambridge chum William Rathbone

Greg, was that “artificial and conventional have taken the place of natural advantages,” in the modern welfare state, and under these conditions degenerates, the unfit, and other undesirables are able to breed without encumbrance, threatening to swamp their biological and social betters.498 The heuristic utility of domestic animals was obvious, or so Darwin explained, “No one who has attended to the breeding of domestic animals will doubt that this must be highly injurious to the race of man. It is surprising how soon a want of care, or care wrongly directed, leads to the degeneration of a domestic race; but excepting in the case of man himself hardly anyone is so ignorant as to allow his worst

497 Lankester, E.R. On Comparative Longevity in Man and the Lower Animals. London: Macmillan and Co., 1870. p.86. 498 Greg, W.R. On the Failure of “Natural Selection” in the Case of Man. Fraser’s Magazine. 1868. Vol.78:358.

246 animals to breed.” However, he also offered the caveat that it would be impossible to

“check our sympathy, if so urged by hard reason, without deterioration in the noblest part of our nature.”499 This sympathy, argued Alfred Russel Wallace a few years earlier, formed the basis of human social evolution and coupled with the intellect and ingenuity unique to the human species, largely freed humanity from the vicissitudes of natural selection. The social body made up man’s natural deficits, such that even “the weaker, the dwarfish, those of less active limbs and less piercing eyesight do not suffer the extreme penalty which falls upon animals so defective.”500 For Wallace, this was no grave consequence, for in his construct, which bordered on the utopian, this sympathy led only to the greater moral and mental perfection of man. But for others, including turn of the century progressive social reformers the conflict between “hard reason” and sympathy (what Alexander Sutherland called the “moral instinct”) represented an insoluble quandary. The development of this moral instinct signaled a genuine social advance and underlay their commitment to reform; charity or reform misguided, however, only bred misery and exacerbated need.501 Thus we add to the list of fin de siecle paradoxes, between increasing prosperity and new depths of poverty, between progress and regress.

As domestication relieved animals of various of the struggles for life that they would encounter in nature, it was seen as a window into the effects of man’s evolving social condition. The use of the analogy with civilization ensured as much. Although

499 Darwin, C.R. The Decent of Man and Selection in Relation to Sex. Vol.1. New York: D. Appleton and Company, 1871. p.162. 500 Wallace, A.R. The Origin of Human Races and the Antiquity of Man as Deduced from the Theory of Natural Selection. Journal of the Anthropological Society of London. 1864. Vol.2:clxii. 501 Bender, D.E. Perils of Degeneration: Reform, the Savage Immigrant, and the Survival of the Unfit. Journal of Social History. 2008. Vol.42(1):5-29

247 this occurred in many different contexts throughout the 19th century, the uses of the analogy addressed common questions about the nature of mankind and the consequences of his departure from it, especially his entry into the modern social world. This affected not only progressive activists, but also the scientists whose work underlay the rational reforms of the period. Thus to the 1894 Congress of American Physicians and Surgeons,

University of Pennsylvania anatomist Harrison Allen suggested that the consideration of the effects of “self-imposed restraint” should precede the study of human anatomy, for

“our acquaintance with man as a domesticated animal, as a degenerate animal, and our ignorance of him as a wild and primitive animal, cause the morphologist to regard human structures with the same sort of interest that the botanist entertains for those plants which have been cultivated so long that he has lost knowledge of the typical forms of the species.”502

Domestication and Psychobiology: Identity, Approach, and Application

The experimental turn in animal psychology followed mounting criticism that the field’s past was marred by baseless introspection about the operations of the animal mind.

Such speculations paled before the tightly controlled experimental evidence of physiologists, whose experiments into the function of nerve fibers encroached upon the psychologists’ purview and undermined their claims to an independent and legitimate science. In 1894 Conwy Lloyd Morgan insisted that his fellow comparative psychologists explain the subjective experience of an animal by reference to the animal’s behavior and, then, appealing only to the simplest psychological processes sufficient to

502 Allen, H. Morphology as a Factor of Disease. p.9 see also footnote same page.

248 explain the behavior.503 Five years later the physiologists Theodore Beer, Albrecht

Bethe, and Jakob von Uexküll created a strictly objective terminology for neuro- physiological research so as to avoid the introspective and anthropomorphizing vocabulary of traditional psychology (e.g. replacing “eye” with “photoreceptor”). The new lexicon proved cumbersome and was seldom used, but it inspired reflection among comparative psychologists regarding their assumptions. One of their most prominent early leaders, Robert Yerkes, admitted in a review of the paper, “Comparative psychologists too often try to put the cart before the horse, so often in fact that most natural scientists have lost faith in their ability to progress and are inclined to ignore their work.”504 A few years later Yerkes felt it necessary to survey a number of prominent biologists for their answer to the question “Do you consider psychology a part of physiology?”505 Yerkes offered a defense of psychology as a stand-alone discipline, but his survey revealed (as far as a small and nonrandom sample group can) that the question was still an open one in the scientific community.

Experimental rigor was not an unproblematic commitment. Experimental methods demanded some measure of control, and in animal behavior study the appropriate degree of control remained a volatile subject throughout the 20th century.

Could animals be made into “organisms of convenience,” interchangeable pieces in an experiment designed to illustrate general behavioral processes, or were they specially

503 Morgan, C.L. An Introduction to Comparative Psychology. London: W. Scott Publishing Co., 1894. p.53. 504 Yerkes, R.M. Editorial: Objective Nomenclature, Comparative Psychology, and Animal Behavior. Journal of Comparative Neurology and Psychology. 1906. Vol.16:388. 505 Idem. Psychology in its Relations to Biology. The Journal of Philosophy, Psychology, and Scientific Methods. 1910. Vol.7(5):113-124.

249 adapted, demanding an appreciation of the diversity of their evolved behavior patterns?506

For some, the increasingly artificial confines of the laboratory and the arbitrariness of the tasks animals were coerced into performing were assurances of greater control, and more objective description. For others these measures of control were antithetical to the expression of “free” or “natural” behavior patterns that had evolved from the long interaction of organisms to specific environmental demands. Domestication emerged early as a means of exploring the tension between experimental control and the natural expression of behavior. Its earlier analogical and normative significances were not obliterated by the precision and objectivity of experimentalism; questions of artificial versus natural behavior, control and free will, and captive versus wild fit neatly with the existing social and scientific tropes of domestication.

Psychobiology, Behaviorism, and the Fate of the Norway Rat

The publication of Edward Thorndike’s dissertation in 1898 signaled the arrival of a truly experimental psychology in the United States. Although Thorndike rejected his faith as a young man, his stern Methodist upbringing seems to have imbued him with the seriousness and severity of interpretation that characterized his later psychological work.

He delighted in criticizing the methods of “the anecdote school,” which he identified especially with Romanes and even took Morgan to task for the latter’s credulity and failure to follow his own parsimony principle. Thorndike arrived at an even more austere interpretation of associative processes in animals than Morgan and conspicuously

506 Term borrowed from Stam, H.J. and Kalmanovitch, T. E. L. Thorndike and the Origins of Animal Psychology: On the Nature of the Animal in Psychology. American Psychologist. 1998. Vol.53(10):1135.

250 avoided the discussion of evolutionary implications present in Morgan’s work.

Thorndike designed his experiments with few expectations. Rather, they were to serve as a demonstration of method. The approach was general, substituting chicks for kittens for dogs as operators of simple puzzle boxes, which were designed to ostensibly force the animal to make associations “free from the helping hand of instinct.”507 Similarly his goal was the understanding of the process of associative learning, not of chicks, or kittens, or dogs particularly, but of animals generally. In his 1898 monograph Thorndike deliberately tried to mitigate the impact of evolutionary and ecological peculiarities in his experimental subjects so as to observe association learning uncomplicated by these considerations.

At the time of Thorndike’s publication the scope and shape that experimental research in animal psychology should take was far from clear. As the comparative psychologist Bennett Galef Jr. has explained, Linus Kline and Willard Small of Clark

University had already developed a substantially different approach than Thorndike to studying animal learning and behavior by 1898.508 Small and Kline were, as Kline himself recalled, “good fellows working together in the same laboratory mutually sharing

507 Thorndike, E.L. Animal Intelligence: An Experimental Study of the Associative Processes in Animals. Ph.D. Dissertation Columbia University, New York, 1898. p.9. Published also in Monograph Supplement No.8 Psychological Review 68-72, 248. Thorndike maintained an active interest in instincts and declared them “well worth study for their own sake.” See Thorndike, E.L. Instinct Biological Lectures from the Marine Biological Laboratory at Woods Holl. Boston: Ginn and Company, 1899. p.57-68. However, his experimental researches dealt sparingly with instinct. 508 Galef Jr., B.G. Reciprocal Heuristics: A Discussion of the Relationship of the Study of Learned Behavior in Laboratory and Field. Learning and Motivation. 1984. Vol.15(4):480-483. See also Boakes, R.A. From Darwin to Behaviourism: Psychology and the Minds of Animals. Cambridge: Cambridge University Press, 1984.. p.143-144; Stam, H.J. and Kalmanovitch, T. E.L. Thorndike and the Origins of Animal Psychology: On the Nature of the Animal in Psychology. American Psychologist. 1998. Vol.53(10):1135-1144; Logan C.A. “[A]re Norway Rats… Things?” p.290-293.

251 each others [sic.] plans and ideas.”509 Their approach stipulated that “a careful study of the instincts, dominant traits, and natural habits of an animal as expressed in its free life- in short its natural history should precede as far as possible any experimental study.

Procedure in the latter case, i.e., by the experimental method, must of necessity be largely controlled by knowledge gained through the former, i.e., by the natural method.”510 The pair explicitly highlighted the importance of evolutionary history and ecological context as a preliminary to the evaluation of learning. Their goal, as Small described it, was “the careful description of the psychic properties of special animal forms,” with each species independently analyzed. “Generalizations,” he added, “will come in due time.”511 Where

Thorndike sought to mitigate instinct through his experimental methods, Small and Kline designed their experiments with the behavioral proclivities of the animal in mind.

In the late 1890s Kline introduced the Norway rat to experimental psychology after having been convinced of the animal’s amenability to experiment by observing

Colin C. Stewart’s investigations on the activity of the rat as a measure of metabolic activity in the Biology Department at Clark. He designed his experimental methods around the assumption that the rat was both a “gnawer” and a “digger” by nature. Thus, boxes containing food were constructed that would allow the rat to gnaw or dig its way into the box to secure its reward. In a reflection on another early research interest, the

“home-finding” capacity of the rat, Kline recalled a discussion he had with his advisor

E.C. Sanford in which he mentioned a series of burrows constructed by rats under the

509 Quoted from Miles, W.R. On the History of Research with Rats and Mazes: A Collection of Notes. Journal of General Psychology. 1930. Vol.3(2):325. 510 Kline, L.W. Suggestions Toward a Laboratory Course in Comparative Psychology. American Journal of Psychology. 1899. Vol.10(3):399. 511 Small, W.S. An Experimental Study. p.133

252 porch of a cabin on his father’s Virginia farm. “These runways,” he remembered, “were from three to six inches below the surface of the ground and when exposed during excavation presented a veritable maze. Sanford at once suggested the possibility of using the pattern of the Hampton Court maze for purposes of constructing a ‘home-finding’ apparatus.”512 Kline did not make use of the suggestion. Instead, it fell to Small to introduce the Norway rat to the maze and, critically for this pairing’s later importance, in the evaluation of learning. Small, of course, emphasized “the rat’s propensity for winding passages” in his decision to use the model of the Hampton Court maze.513

Significantly, he also prefaced the publication of his experimental results with a thorough, albeit anecdotal, record of the psychological development of a litter of albino

Norway rats, evidence of G. Stanley Hall’s remaining influence at Clark.514 This respect for the natural (i.e. evolutionary) history and ecological context of the rat’s mind diverged sharply from Thorndike’s general approach, as Small was wont to demonstrate in a not-so-subtle dig at his colleague from Columbia: “Experiments must conform to the psycho-biological character of an animal, if sane results are to be obtained.”515

512 Quoted from Miles History of Research. p.331. 513 Small, W.S. An Experimental Study… II. p.208. 514 Idem. Notes on the Psychic Development of the Young White Rat. American Journal of Psychology. 1899. Vol.11(1):80-100. Small’s assertion in this article that “there is solid scientific reason that Genetic Psychology has much to hope for from minute and accurate records of the developmental periods of young animals of all species. It may never be possible to reconstruct a complete psychic organism from the evidence of a single trait- an ideal borrowed from morphology- but something surely may be accomplished towards a comparative embryology of the soul,” (Ibid., p.80) made definite allusions to Hall’s construct of “genetic psychology” and its methodological corollary, the “embryology of the soul.” Small seems have done this, however, less as an homage to Hall than as an effort to convince the elder psychologist of the importance of laboratory methods. As Hall delved more deeply into developmental psychology he “doubted whether laboratory methods could be devised that would return yields of much value” and, Small recalled, “he did not give me, at first, much encouragement in my project.” Quoted from Miles History of Research. p.333. 515 Idem. An Experimental Study…II. p.206.

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Kline and Small had already initiated their experimental program at Clark before learning of Thorndike’s work. The publication of the latter’s deliberately polemical thesis, however, sharpened the distinction between these competing visions for experimental animal psychology in the United States. Kline, in his review of the work, suggested the principal value in Thorndike’s method would be in investigating “animal psychosis” rather than the normal functioning of the animal mind.516 Small, late in his career, recalled his reaction to Thorndike’s monograph: “I studied it carefully and thoroughly- and with profit. The chief advantage I derived from it was to make me more certain that I was on the right track in seeking to conform experimental procedure to the native tendencies of my animals.”517 His early papers seem to support this recollection.

Although Small was concerned with impartiality and carefully controlled experiments, he viewed Thorndike’s commitment to objectivity as naïve and misleading. He deliberately avoided quantitative description and graphical representation (i.e. Thorndike’s learning curves) in his papers, preferring instead to describe the “conduct of the rat… as it was minutely recorded at the time of observation.” This he represented as “analogous to vital staining in histology,” but this means of description was perhaps more typical of natural history. He reasoned that “in view of the role one’s own prepossessions and limitations are bound to play in interpretations, it will be an advantage to the reader to know the observed facts upon which the writer’s interpretations are based.”518

516 Kline, L.W. Review: Animal Intelligence: An Experimental Study of the Associative Processes in Animals by E.L. Thorndike. American Journal of Psychology. 1898. Vol.10(1):150. 517 Quoted from Miles History of Research. p.333. 518 Small, W.S. An Experimental Study…II p.210.

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Small’s concern for “native tendencies,” was crucial for shaping, not only his methods, but also the questions he sought to answer. One such question arose early in his work with the rat: the impact of domestication on learning and behavior. The thought was, he claimed, “purely a by-product of the study- it was not contemplated at all in the beginning- and it takes us a little out of the straight line of exposition;” however, he assured, “the results have a suggestive value sufficient to warrant the delay.”519 He had wished to use wild (gray) rats exclusively, but the difficulties he experienced with the first series of such rats convinced him to use domesticated (albino) rats for the remaining series, a situation similar to that described by Colin Stewart from whom Kline and Small got the idea to use Norway rats.520 Domestication’s “suggestive value,” however, had more to do with Small’s “psycho-biological” view than the circumstances of the experiment. Even from the publication of his preliminary work on the development of the albino rat, Small’s respect for environmental context forced him to consider the psychic impact of “the vicissitudes of an environment different in all its factors from that of the free wild life in which [the rat’s] psychic nexus was woven in the loom of necessity.”521

519 Ibid. 520 In an exchange with W.R. Miles some thirty years after leaving Clark, Stewart explained his rationale for using albino rats: “If anyone wants to know why I changed from wild gray rats to white rats in 1895, let him work with gray rats for a year. I don’t deserve any credit for the change.” Quoted from Miles History of Research. p.334. Small, in an exchange with Miles around the same time, recalled that in 1898 he had “been carrying on with Dr. C.F. Hodge, a minor study on the transmission of acquired characters using the albino rats for experimental purposes. I had kept a fairly large number and had become rather familiar with their ‘manners and customs’ and had developed a liking for the animal… My familiarity with the albino rat was, I believe, what determined that animal as my special object of study.” This account suggests that the albino rat had been closer to his original plans than he noted in the original paper. At nearly thirty years after the experiments, though, Small admitted that his own memory might be “treacherous.” Quoted from Miles History of Research. p.332-333. 521 Small, W.S. Notes on the Psychic Development. p.100.

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His first experiments, using Kline’s gnawing and digging boxes, involved only albino rats. Even in this instance, however, Small thought to suggest that one group of rats that entered the boxes more quickly and with more consistent times was “more highly domesticated” (for having been gentled) than the others.522 He indicated that temperament, rather than intelligence may have meant the difference in their performances. His experiments with the Hampton Court maze offered him a clearer comparison of wild (gray) and domesticated (albino) rats, and thus the opportunity to evaluate the mental effects of domestication. In this set of experiments, the albino rats demonstrated a slight advantage in time taken to reach the center of the maze; the gray

(Small called them brown) rats held an edge in the number of errors committed. He credited the minor differences in performance primarily to the wild rat’s increased vigor and heightened fear. However, as Small determined, “In ability to profit by experience… the two are not far different. This conclusion does not tally well with the general opinion that animals suffer mental deterioration under domestication. However this may be with other animals, it evidently is doubtful in this instance.”523

Of the other remarkable and “unexpected” facts that emerged from the comparison, Small was particularly impressed by the survival of instincts long defunct under the conditions of domestication. He could detect, for instance, “no difference in the curiosity manifested, either in kind or degree [between his wild and domesticated

522 Idem. An Experimental Study. p.156. 523 Idem. An Experimental Study…II p.221.

256 rats]. In view of the many generations of luxurious idleness of the white rat, this profound and enduring nature of specific psychic traits is striking.”524

Perhaps the result should have been less surprising to Small, as he was, in a way, conditioned to look for just such phenomena. Instinct was fundamental to his understanding of the mind as the product of so many adaptive mental processes, and he found examples of them everywhere. In each of his three papers on the rat, he mentioned the “tenacity,” “survival,” and “persistence” of “wild traits” that had been important to the history of the species but now, especially in a domesticated state, had ceased to be adaptive. Both Small and Kline were influenced at the time by the English physician and evolutionist, Louis Robinson, whose book Wild Traits in Tame Animals (1897) highlighted the value of domesticated animals for revealing the extraordinary longevity of vestigial instincts.525

Small’s investigations into the effects of domestication raised (or, perhaps more accurately, reinforced in him) “the question whether zoological psychology may not profitably turn from its almost exclusive search for variation, to a search for the relatively invariable factors in the animal mind… out of which, in higher differentiations, human nature is made.” In highlighting the primacy of instinct, the “suggestive value” of his work on domestication became clear: “By this method, if by any, will be gathered the material for the natural history of the mind. From this source, light may be expected upon many obscure problems in individual and anthropological psychology.”526

524 Ibid. 525 Small and Kline acknowledged their debt to Robinson by punctuating their frequent use of the phrase “wild traits” and, of course, by citing his book. See Kline, L.W. Methods in Animal Psychology. American Journal of Psychology. 1899. Vol.10(2):278 and Small, W.S. An Experimental Study. p.141. 526 Small, W.S. An Experimental Study…II p.222.

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Small was forthright about finding the human in his animal subjects. For instance, he noted in his maze trials using wild gray and domesticated albino rats of both sexes, that “domestication tends to reduce the disparity between the sexes.” In trying to reconcile this with contrary evidence from contemporary anthropology (having already made the leap from domesticated rat to civilized human), he ventured in terms reminiscent of Alfred Russel Wallace: “Possibly a point in development has been reached where psychological development has attained a greater freedom than heretofore from physiological and biological determinants.”527 To realize the human in his animal subjects was inevitable, he claimed: “[I]s not a certain amount of chastened anthropomorphism a wholesome specific, a kind of saving grace against the scientific pedantry that thinks to create a new science of comparative psychology with the imperfect instruments of experiment and the law of parsimony[?]” he asked rhetorically before answering, “The real difficulty lies not in the tendency to interpret animal intelligence in the terms of human experience, for we have no other way, but in the faulty and imperfect analysis of human experience.”528 Thus, a more perfect analysis of human civilization, which Small regarded as “a kind of domestication,”529 would emphasize the evolutionarily meaningful rather than the capricious variations of human mentality, as he had done with his albino rats.

Small’s pairing of rat and maze ultimately became emblematic of the animal psychology that developed in the United States. The irony, Galef indicated, was that this occurred within the general framework established by Thorndike, in which both organism

527 Ibid. p.212. 528 Ibid. p.228. 529 Ibid. p.212.

258 and apparatus were abstracted from their ecological and evolutionary significance. Thus, by 1938 E. C. Tolman was able to affirm: “Everything important in psychology… (save such matters as involve society and words) can be investigated in essence through the continued experimental and theoretical analysis of the determiners of rat behavior at a choice point in a maze.”530

This situation was affected primarily by the rise of behaviorism in American experimental psychology. Thorndike was the movement’s most significant precedent, but even his stance was judged insufficiently objective by early behaviorists, especially his unwillingness to disavow the subjective experience of an animal as a legitimate subject for psychological study.531 Behaviorism’s founder, John B. Watson, trained at the

University of Chicago in the early 1900s, where a philosophical dispute about the proper aims of bio-behavioral research- related to that between Thorndike, Kline, and Small- was ongoing.

Jacques Loeb, a German émigré and experimental physiologist, denied the progressive evolutionism that constituted the intellectual core of the University, represented by the philosopher John Dewey, zoologist Charles Otis Whitman, psychologist James Angell, neuroanatomist Henry H. Donaldson, and anthropologist

George Herbert Mead among others. As described by Philip J. Pauly, Loeb’s scientific ideal sought prediction and control of biological phenomena via an understanding of the fundamental properties shared by all living matter. The purpose of comparative physiology (which in Loeb’s definition subsumed behavior studies) was, thus, to

530 Quoted from Galef Jr. Reciprocal Heuristics. p.482. 531 Thorndike’s own experiments were, of course, limited to the study of behavior. Boakes, R.A. From Darwin to Behaviourism. p.76.

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“discriminate between the general properties of living matter and the functions of specific organs,” i.e. to raise the general and to discount the specific differentiations of animal function.532 Watson matriculated to Chicago with the intent to work under Dewey and

Angell in the latter’s psychological laboratory, but became increasingly drawn to physiology after attending Loeb’s lectures. Watson was attracted by Loeb’s zeal for experiment and began discussions with him for possible dissertation topics in neurophysiology. Angell and Donaldson, though, dissuaded Watson from working under

Loeb and found a project for him correlating learning ability with brain growth and differentiation in albino rats, which had recently become a fixture of Donaldson’s neuroanatomical laboratory.533

Upon graduation, Watson was initially sympathetic to Angell’s evolutionarily mindful “functionalism.” Among his most significant early researches was a field study of behavior in terns on the uninhabited atoll of Bird Key in the Tortugas, which included careful analysis of their instinctive behavior patterns. He was simultaneously impressed and frustrated by the “extreme monotony, fixedness, and lack of variability in the responses” of these animals, despite his considerable efforts to disrupt these behaviors.534

The appeal to instinct, though, was unsatisfying. It seemed simply to give a name to the original mystery. Meanwhile, his laboratory studies with rats reduced the problem of

532 Pauly, P.J. Controlling Life. p.85. 533 That Watson failed to find the correlation between learning ability and neural development (he discovered that young rats could demonstrate learning prior to the myelinization of their nerve fibers), as Donaldson had predicted he would, may have heightened his skepticism toward the “functionalist,” i.e. developmental and adaptive, standpoint held by Angell, Donaldson, and the other Chicago Progressives. On the importance of evolution and development to Donaldson’s choice of the rat as an experimental animal and his attempt to cultivate it as a “biological standard against which the magnitude of diverse adaptive specializations might be measured,” see Logan, C.A. Altered Rationale p.3-24; Idem. “[A]re Norway Rats… Things?” p. 287-314. 534 Pauly, P.J. Controlling Life. p.174. See also, Boakes, R.A. From Darwin to Behaviourism. p.147-148.

260 orientation within a maze to simple kinesthetic feedback between muscles and the brain, an explanation that seemed to offer Watson not only greater objectivity, but also much more control over the behavior of his animals. The path forward was clear. In the years immediately following these investigations, Watson became more critical of the functionalist standpoint and in a 1910 letter to his friend Robert Yerkes had raised the possibility of a purely objective psychology that terminated with the description of behavior rather than the interpretation of consciousness.535 Three years later Watson made his views explicit as well as public in his monograph “Psychology as the

Behaviorist Views It,” or as it has become known to the history of psychology, the

Behaviorist Manifesto.

In a distinctly Loebian turn, Watson declared from the opening of his manifesto that psychology’s “theoretical goal is the prediction and control of behavior” rather than the mere description of mental life.536 As he had done with consciousness, Watson later proposed that the brain and instincts were beyond the pale of experimental psychology, and with them the theoretical framework that emphasized their progressive development and adaptive function. While criticism of Watson’s position was ample and few fully adopted his stridently objective stance, the suggestion that the mental life of animals, humans included, could be explained solely in terms of behaviors that were observable and predictable and that instincts and higher order consciousness could be dispensed with provided a way forward for experimental psychology. Certainly Watson’s critique that psychology’s dalliance with introspection had yielded little in terms of positive evidence

535 See Leys, R. Meyer, Watson. p.130. 536 Pauly, P.J. Controlling Life. p.174.

261 and even fewer practical applications struck a chord in a field still trying to find its experimental footing. A new generation of experimental psychologists, trained in departments and laboratories of psychology and not merely reformed philosophers or biologists, seemed to welcome an approach that relieved them of their philosophical debt to evolutionary biology and legitimized their independence as an experimental science.

Behaviorism’s hegemony over American experimental psychology in the next half century was never complete. It was, however, enough to force the psychobiological point-of-view of Kline and Small and many of the Chicago Progressives into a minority position in the mainstream of American experimental psychology.537

The stereotyped image in experimental psychology of a rat navigating a maze is more the responsibility of Watson than Small, and it held a very different meaning than

Small had imagined at the turn of the century. By the 1920s the maze was no longer an experimental facilitation of the rat’s burrowing instinct, but a generalized learning task.

The albino rat was similarly shorn of the specific differentiations that concerned

Donaldson at the time that he introduced Watson to the animal and became, as much as the maze through which it maneuvered, an instrument of the experiment. The extent to which an animal subject became a true denizen of the laboratory was, to the behaviorist, another measure of control and a further guarantee of objectivity. To the psychobiological view, the adjustment of the animal to its artificial confines raised questions about the exercise of instinct and “natural habits,” and thus about the translation of laboratory findings to the “free life” of the experimental animal.

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Freedom from the Tyranny of Instinct: Charles Otis Whitman, Progressivism, and Domestication Research at the Turn of the Century.

At the same time that Small and Kline were performing their experiments at

Clark, some of the most incisive and sweeping statements on animal behavior came, not from a psychologist, but from the doyen of American biology, Charles Otis Whitman.

His 1898 lecture “Animal Behavior” and his paper the following year, “The Myths of

Animal Psychology,” along with his own work in animal behavior suggested Whitman was little concerned with the traditional boundaries distinguishing biology and psychology.538 His view of biology was too grand to fret over the distinction. In the manner of Herbert Spencer, Whitman was committed to the idea that a generally progressive and orderly evolution guided development of mind as well as body, of the organic as well as the social and cultural. “Biology has taught us,” he declared, “what constitutes progress and those who now write history must deal with it not from the standpoint of dogmatic struggles, court intrigue, diplomacy, [and] war, but from the standpoint of evolution.”539 His orthogenesis (an internally guided evolutionism, peculiar to the late 19th and early 20th centuries) acted as a universal leaven in Whitman’s view of life, its past and future, its proper study and organization.

Pauly argued that as a young man Whitman rejected the radical Adventism of his parents and, like Thorndike and a number of other American scientists coming of age in the latter half of the 19th century, found the coherence he had lost in his new faith in the

538 Whitman, C.O. Animal Behavior in Defining Biology: Lectures from the 1890s. Cambridge, Massachusetts: Harvard University Press, 1986. p.230-272; idem. The Myths of Animal Psychology. Monist. 1899. Vol.9(4):524-537. 539 Whitman, C.O. Undated note included in the folder: Whitman, Charles Otis- Biological Farm Project in Charles B. Davenport Papers, B.D27, Series I, APS.

263 natural sciences. Although he became committed to strictly material rather than supernatural explanations, Whitman’s evolutionism remained suffused with the telos and purpose of his former faith.540

That Whitman reached such a high station in American biology is remarkable given his late introduction to academic research. His introduction to intensive biological research began only at the age of 30 at Alexander Agassiz’s school for natural history at

Penekise Island in the summers of 1873 and the year following. In the custom of

American graduates at the time, he completed his training in modern research methods in

Germany at Leipzig with the parasitologist Rudolf Leuckart. He published his first monograph on the development of a subject he encountered in Leuckart’s laboratory, the fish leech Clepsine, this at 35 years old. Whitman carried on research with Clepsine and related leeches for the next 20 years, at numerous stations, including the Imperial

University of Tokyo, the Naples Zoological Station under Anton Dhorn, the Museum of

Comparative Zoology at Harvard, the privately financed Allis Lake Laboratory near

Milwaukee, Wisconsin, Clark University (as chair of the Zoology Department), the

University of Chicago (as chair of the Division of Biology and of the Department of

Zoology), and the Marine Biological Laboratory (as its first Director). His work on leeches investigated their morphology, physiology, development, zoological relations, ecology, and behavior and in its breadth established what historian of science Richard

Burkhardt Jr. called Whitman’s

540 Pauly, P.J. From Adventism to Biology: The Development of Charles Otis Whitman. Perspectives in Biology and Medicine. 1994. Vol.37:395-408.

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“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.”541

Whitman’s status at the turn of the century owed at least as much to his administration of American biology (at Clark, Chicago, and especially the Marine

Biological Laboratory) as to his careful and comprehensive researches.542 Naturally he sought to define biology in the image of his own rigorous methods. In 1899 he delighted that “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”- i.e. the dynamic processes of life- were finally “beginning to take [their] place beside the more strictly morphological studies which have so long monopolized the attention of naturalists.”543

This was being affected through specialization, which Whitman accepted as the natural course of development (whether organic or social), and thus permitted in his administrative appointments at Clark and Chicago. In his orthogenetic ideal, he imagined that this specialization would lead inevitably to greater coordination: “An organism travels toward its most perfect state in proportion as its component cell-individuals reach the limit of specialization, and form a whole of mutually independent parts. Scientific

541 Burkhardt, Jr. R.W. Patterns of Behavior. p.22. 542 This is almost certainly an understatement. Whitman’s publications, though well regarded, numbered just over 60. E.D. Cope, by comparison published nearly 1400 articles and communications in 10 fewer years of life. Whitman, is more often remembered by historians for his administrative roles at Clark, Chicago, and the MBL than for his work on Clepsine, Necturus, and pigeons. 543 Whitman, C.O. The Myths of Animal Psychology. p.524.

265 organization obeys the same law.”544 He worried, however, that this latter stage sometimes “[lagged] lamentably behind,” leaving increasingly isolated specialists to work without thought of integration or a sense of the whole.545 So he intended to remind the Society of American Naturalists in his President’s speech for the1897 meeting in

Ithaca: “We must become more 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.”546

In similar terms, Whitman warned psychologists against viewing the animal mind too narrowly: “Comparative psychology is a science of the future; and if at present it is only a part of general biology, it follows that any attempt to soar to the ‘nature and development of animal intelligence,’ except through the aid of long schooling in the study of animal life, is doomed to be an Icarian flight.”547 It was only through sustained experience with his animal subjects, he claimed, that he was able to recognize the most relevant of their behaviors, including the “deceptive quiet” of Clepsine and the “extreme timidity” of Necturus, the genus of salamanders he had first studied while stationed at the

544 Whitman, C.O. Preface in Biological Lectures Delivered at the Marine Biological Laboratory of Woods Holl. Boston: Ginn and Company, 1891. p.iv. 545 Whitman, C.O. Specialization and Organization: Companion Principles of All Progress- The most Important Need of American Biology (Delivered 1890) in Biological Lectures Delivered at the Marine Biological Laboratory of Woods Holl. Boston: Ginn and Company, 1891. p.23. See also Maienschein, J. Whitman at Chicago: Establishing a Chicago Style of Biology? in The American Development of Biology. Ronald Rainger, Keith R. Benson, and Jane Maienschein (eds.). Philadelphia: University of Pennsylvania Press, 1988. An anecdote from Henry H. Donaldson’s memoirs suggests the extent to which the delineation of specialties at Chicago had gotten out of hand: the janitor in the as yet unfinished Science Hall, after hanging plaquards announcing the special focus of each of the laboratories, installed on his broom closet a sign reading “Broomology.” Donaldson, H.H. Memories for My Boys. 546 Whitman, C.O. Some of the Functions and Features of a Biological Station. Science. 1898. Vol.7(159):43. Whitman apparently missed the meeting, and the lecture was not delivered, except in print months later. 547 Whitman, C.O. The Myths of Animal Psychology. p.537.

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Allis Lake Laboratory in the late 1880s.548 His criticism was not limited to psychologists.

He could no more brook the mechanistic myopia of the physiologist (he almost certainly had in mind his colleague Loeb here) “proclaiming to the world that instinct reduces itself in the last analysis to heleotropism, stereotropism, and the like.” The problem, he explained, was that in this view “[t]he whole course of evolution drops out of sight altogether and things are explained as if the organic world were a chemical creation only a few hours old.”549

For Whitman, of course, the analysis of behavior could never have been limited to the immediate act, however minutely studied. Rather, behavior was the product of a deep historical and largely determinate evolutionary process. It is not surprising, then, that in his behavioral work he asserted the primacy of instinct, those behaviors that were innate and relatively ancient and, he presumed, met the most basic needs in the history of the race. Learning and conditioning might meet the immediate needs of the animal, but the capacity for these was secondary to and, indeed, derived from fundamental instinct. As the more ancient of the two, Whitman argued, “instinct precedes intelligence both in ontogeny and phylogeny, and it has furnished all the structural foundations employed by intelligence.”550

Intelligence, however, was not the inevitable outcome of instinct. Rather, instinct ensured that an animal’s behavior was biologically meaningful, but rendered the animal “quite blind” to this meaning. For the animal to gain insight into its actions, for it to become intelligent, it would need to overcome the extreme fixedness of instinct.

548 Whitman, C.O. Animal Behavior. p.288, 296. 549 Whitman, C.O. Features of a Biological Station. p.43. 550 Whitman, C.O. Animal Behavior. p.263.

267

Domestication, Whitman reasoned, offered a rare clue as to “how instinct behavior can take one step toward mind behavior.” He described an experiment in which he placed the eggs of wild passenger pigeons, “semi-domesticated” ring doves, and fully domesticated dovecote pigeons outside of their nests and observed the birds’ reactions.

Of the three “grades,” the domesticated dovecote pigeon alone retrieved two eggs, the ring dove never more than one, while the wild passenger pigeon continued to brood an absent clutch. As Whitman interpreted:

In the wild species the instincts are kept up to the higher degrees of rigid invariability, while in species under domestication they are reduced to various degrees of flexibility and there is a correspondingly greater freedom of action.551 According to Whitman the breakdown of instinctive patterns in the domesticated dovecote pigeon, “so far from indicating psychical retrogression,” was, rather, symptomatic of freedom of choice and increased plasticity: the prerequisites to intelligence. His reasoning, that these “faults of instinct” were not in fact faults, relied on his presumption that the fundamental rules underlying domestication and organic evolution were the same. “Domestication,” he explained, “merely bunches nature’s opportunities and thus concentrates results in forms accessible to observation. Natural conditions are certainly working in the same direction, only more slowly. The direction and the method of progress must, in the nature of things, remain essentially the same.”552

Thus, the luxury of superior instincts gave wild animals advantages effectively identical to the protections and provisions of animals cared for in captivity.

551 Ibid. p.267. 552 Ibid. p. 267, 270.

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This equivalence, however, only seemed to hold for mental traits in Whitman’s integrative work on pigeons. He viewed morphological and physiological aberrations less as opportunities than in the value laden terms of degeneration. In one instance, he declared the “150 ‘mongrels’ collectively known as domesticated pigeons” unsuitable for certain physiological inquiries on account of “the countless degenerations and crossings suffered by the various ‘domesticated breeds’ of pigeons since their existence as a pure, wild species.”553 With physical traits, he thought it crucial to point out the incongruities between domestication and nature. In the process he cast domestication in a very different light (capricious, impure, and, above all, unnatural) than he had done in his work on behavior. How, then, to account for the inconsistencies in Whitman’s treatment of domestication and its relations to nature?

Simply put, mental evolution was more central to Whitman’s universal progressive view than were changes to the body. Intelligence, particularly, occupied a special place in Whitman’s philosophy, as the mark of progress in nature and civilization alike. He viewed domestication as a catalyst for this process. It acted as a buffer between the animal and the exigencies of survival that made instinct so authoritative.

This “freedom of action” created choice and should be considered “the dawning grace of a new dispensation[,]… the open door through which the great educator, experience comes in and works every wonder of intelligence.” This was not only true of pigeons but of humans. Just as the influence of instinct diminished in pigeons with advancing domestication, so, Whitman claimed, “[i]n the human race instinctive actions characterize the life of the savage, while they fall more and more into the background in the more

553 Whitman, C.O. Posthumous Works Volume II, Oscar Riddle (ed.). p.173.

269 intellectual races.”554 As Pauly demonstrated, Whitman belonged to a generation and, more particularly, to a coterie of intellectuals at the University of Chicago that made little distinction between biological and cultural evolution.555 Whitman, thus, viewed the modifications of instinct under domestication not as faults that might undermine his sense of social and cultural advance, but in terms of the democratic ideals of freedom and opportunity that marked for him the height of social progress.

Whitman’s interpretation of the behavioral changes attendant to domestication in his pigeons was similar to that of Small working with his albino rats a few years later

(and without apparent knowledge of Whitman’s behavioral work). Domestication in their view had not obliterated or degraded instinctive behavior, but had released the animal to a degree from the tyranny of fear or specific behavior patterns. The similarities in their interpretations had to do in part with their approaches to studying behavior. Small and

Whitman both emphasized the necessity of long observation of the whole repertoire of behavior in a given species before any experiments could be performed. Their approach was evolutionarily and ecologically mindful, evidenced not only by their emphasis on instinct, but also by their insistence that the investigator should, in Whitman’s words,

“observe and experiment under conditions that ensure free behavior.”556 Balancing the elicitation of natural behavior with experimental control was one of the great problems facing students of animal behavior at the turn of the 20th century. Whitman was torn between these aims, but also imagined that he had found a solution.

554 Whitman, C.O. Animal Behavior. p.270, 244. 555 Pauly, P.J. Controlling Life. p.67. 556 Whitman, C.O. Animal Behavior. p.236.

270

For decades, beginning in the early 1890s, Whitman campaigned for a biological station (it would ultimately become known as the “biological farm”) that would be commensurate with his integrative vision of biology. He explained that the great problems of biology demanded “facilities for long-continued experimental study, under conditions that admit of perfect control,” yet these were problems that “cannot be wholly settled in the laboratory.” Instead, “[t]hese problems require… to be taken to the field, the pond, the sea, the island, where the forms selected for study can be kept under natural conditions and where the work can be continued from year to year without interruption.”557 Try as he might to avoid artificialities, Whitman seemed to recognize the quandary between free behavior and captivity that Helen Spurway would describe 50 years later, and this consideration influenced the naming of his idealized station:

“Biological Farm, broadly defined, is perhaps the best we can do for a name, as the work would be… upon plants and animals under cultivation.”558 Whitman was not discouraged by the realization, but included domestication among the foremost questions to be addressed at the Farm. In a letter to Conwy Lloyd Morgan explaining the facility, he noted, “I could utilize several acres in attempting to domesticate some of our wild birds like the quail, the duck, the water hen, wild turkey, the opossum, and a thousand other things. I believe the experiments in domestication of wild animals would open up a

557 Quoted from Whitman, C.O. Features of a Biological Station. p.43 (originally in the Program of Courses in Biology, University of Chicago 1892). Whitman, C.O. A Biological Farm: For the Experimental Investigation of Heredity, Variation and Evolution and for the Study of Life-histories, Habits, Instincts, and Intelligence. Biological Bulletin. 1902. Vol.3(5):214. 558 Ibid. p.218.

271 wide field for the study of habit and instinct, and for the study of variation in form and other characters.”559

Despite his relentless campaigning, Whitman never saw his vision of a biological farm realized. The station was intended to match the breadth of his approach to biology, and while both gained wide admiration, they were also sufficiently daunting to inspire few adherents.560 Without the facilities that he imagined necessary, it is difficult to determine what he thought he might learn about instinct and intelligence from the experiments in domestication. Some clue, however, may be available from Whitman’s continued researches on pigeons at his home colony, with which he became increasingly engrossed in the last decade of his life before his death from pneumonia in 1910. As his progressive evolutionism had prepared him to see, he referred again and again in his work with pigeons to the signs of incipient intelligence, the beginnings of higher, and especially human, faculties. Reflecting on the interactions of two pairs of pigeons (these were, significantly, “fully domesticated” dovecotes), Whitman remarked, “Here is recognition of partners, love; recognition of non-partners, aversion; the union of the pair owning the roost against the newcomers; and submission to the results of victory as final on the part of the conquered. Is this not human? Is such behavior conceivable without some intelligence?”561 It seems that Whitman sought from domestication not only clues about the beginnings of natural intelligence, but also the seeds of human social advance.

559 Whitman, C.O. “Part of a Letter to Lloyd Morgan” undated in Davenport Papers B.D27, Series I, Folder Whitman, Charles Otis- Biological Farm Project, APS. 560 Burkhardt Jr. Patterns of Behavior. p.27. 561 Whitman, C.O. Posthumous Works of Charles Otis Whitman, Volume III, The Behavior of Pigeons. Harvey A. Carr (ed.) Washington, D.C.: Carnegie Institution of Washington, 1919. p.154.

272

Whitman’s researches on animal behavior had little lasting impact in the United

States. Richard Burkhardt Jr. explains that Whitman had become gradually more isolated in the latter years of his life, in part because of his singular devotion to his private collection of pigeons and because American biology had moved further from the integrated ideal that he maintained, increasingly oriented toward specific problems rather than the intensive study of organisms. The shift was such that when Whitman’s extensive research notes on pigeons were finally collected and edited in 1919, their publication was “a nonevent for most of the scientific community.”562 Also in this time behavior fell increasingly under the purview of psychology in the United States, and investigators in that field possessed of a similar psychobiological view, such as Small, had either abandoned animal studies or been marginalized. Two decades into the 20th century the same basic questions about variation and evolution, for which domestication had so much appeal, were not being asked in mainstream studies of behavior. Whitman’s imperative, that “instinct and structure are to be studied from the common standpoint of phyletic descent,”563 was largely unheeded in the next half century of American animal behavior study.

Charles R. Stockard, the Rockefeller Foundation, and the Preservation of the Constitutional View of Body and Mind to Midcentury.

After 1920 there remained a small number of investigators, generally working in isolation at the periphery of biology and psychology, who still sought to integrate behavior into their pursuit of fundamental questions about evolution. For these few

562 Burkhardt Jr. R.W. Patterns of Behavior. p.33. 563 Whitman, C.O. Animal Behavior. p.262.

273 domestication remained a useful tool for engaging such questions. It also retained its suggestive appeal for clarifying the relations between organic and social evolution, this despite the emerging criticism that cultural and social development was autonomous and not predicated on biological evolution. The critique of use inheritance that began with

August Weismann in the last quarter of the 19th century left Lamarckism, so crucial to the identification of the social with the organic, on increasingly unstable footing by the first quarter of the 20th. Among the nascent social sciences, there was a growing appreciation of cultural rather than biological factors as well as of quantitative rather than qualitative description, led especially by Franz Boas, his students, and followers. Historian George

Stocking Jr. noted that the impact of “Boas’ nominalist critique was the passing of a romantic conception of race- of the ideas of racial ‘essence,’ of racial ‘genius,’ of racial

‘soul,’ of race as a supraindividual organic identity. Such notions could survive quantification only through the obscurantism of typological thinking.”564

While culturally mindful interpretations of social evolution diminished typological thinking, this should not be taken as a simple and swift displacement.

Psychology adopted cultural explanations more slowly and in a more limited fashion than anthropology.565 New evidence from emerging biological fields helped to preserve some of the essentialist and materialistic thinking that the recognition of culture sought to undercut. In the first quarter of the new century investigators began to recognize the complex relations between genetics (focused at the time on discontinuous variation in individual lineages rather than minute variation within populations), endocrinology

564 Stocking, Jr. G.W. Race, Evolution, and Culture: Essays in the History of Anthropology. New York: The Free Press, 1968. p.194. 565 Ibid. p.267.

274

(which held promise for the control of development, physiology, and behavior), and nervous function. These interactions gave credence to older, assumed connections between body and mind and revived interest in the idea of “constitutional type” along with its associated “dynamic, adaptational disease framework.”566 Around 1920, the problem of “type” seemed less like obscurantism than a legitimate subject for study in

American biomedicine.

Workers in this mold directed focus away from bacteriological models of disease to discuss the more opaque chronic diseases, which they frequently interpreted as consequences of civilization. Domestication, reputed to simultaneously modify form and behavior, occurred to some of these investigators as a useful model for investigating or explaining whole-organism interactions between physiology, behavior, and environment.

It also found use, through the analogy with civilization, as a means for discussing the long-term perils of modern life.

Historian of science Sarah W. Tracy identified Charles R. Stockard as one of the

“principal architects of the constitutional program” in the United States.567 Stockard was among the first to investigate the Gordian knot of heredity, endocrines, and behavior, culminating in his work at the experimental dog farm built for him in Shrub Oak,

Westchester County, New York. His holistic view undoubtedly influenced his ambitious research goals and convinced him that such a synthesis was possible. His understanding of domestication supported this belief and provided a crucial epistemic bridge between his work hybridizing dog breeds and his fears about the dysgenic effects of race crossing

566 Tracy, S.W. An Evolving Science of Man: The Transformation and Demise of American Constitutional Medicine, 1920-1950 in Greater Than the Parts. p.162. 567 Ibid. p.173.

275 and modern life. To understand how and why he came to this view requires a look at the contexts and contacts that shaped his early career as well as the sources of intellectual and financial support that made his comprehensive program on personality and constitution possible.

Charles Rupert Stockard was born in Stoneville, Mississippi to the local physician, Richard Rupert Stockard and his wife Ella Hyde, neé Fowlkes, in 1879.568 His remote upbringing cultivated an interest in natural history, a preparation common to many in the first generation of American experimental biologists. He graduated from

Mississippi Agricultural and Mechanical College with a B.S. in 1899 and an M.S. in 1901 before taking the role of Professor of Military Science at the school. He continued in the same role at the nearby Jefferson Military College until 1903 when he matriculated to

Columbia University to pursue a graduate degree in zoology. Stockard ultimately took his Ph.D. in 1907 under the guidance of Thomas Hunt Morgan, whose interests in embryonic development and regeneration had a profound influence on Stockard’s early career.

Stockard’s earliest published work suggests a wide scope of biological interests, including observations of nesting habits in birds, behavior and inheritance in the walking stick (Aplopus mayeri), and observations of the spoon-billed sturgeon. Near the end of his time at Columbia, his gaze increasingly narrowed upon problems of development, beginning with the artificial production of deformed embryos exposed to lithium and magnesium salts. Philip J. Pauly noted that the shift to experimental embryology was

568 Edwards, D.J. Memorial to Dr. Charles R. Stockard. Biological Bulletin. 1940. Vol.79(1):19. Edwards recalled that Stockard reflected self-consciously on the biological and sociological lessons derived from his father’s rural practice often in his later years.

276 fortuitous for Stockard’s career. A year before graduating, he was hired at the Cornell

Medical College in Manhattan as an assistant in embryology. The college was eager to assert its claims to original anatomical research, which was fast becoming the hallmark of modern medical schools, and Stockard presented a convenient solution to this need.569

Despite Stockard’s importance to the school’s image, his experiments on the brackish minnow, Fundulus heteroclitus, little impressed his senior medical colleagues at the school and by 1910 had still failed to win him proper laboratory space. Pauly argued that at this point Stockard began searching for ways to make his research more medically and socially relevant.570 He had already begun working with alcohol, ether, and other agents, after he determined that the anesthetic properties of magnesium were to blame for its teratogenic effects.571 Stockard quickly seized on the social resonance of alcohol.

Amid the din of temperance sermons, he reframed the thrust of his research, from reliably producing “monsters” with the hope to someday “obviate certain monstrous conditions,” to determining the effect of parental alcohol intake on progeny.572

569 Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? Journal of the History of Biology. 1996. Vol.29(1):6. That Stockard’s uncle, William Polk, was dean of the college at the time of Stockard’s hiring, suggests the convenience of the selection. 570 Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? p.7. 571 Stockard, C.R. The Development of the Artificially Produced Cyclopean Fish- “The Magnesium Embryo.” Journal of Experimental Zoology. 1909. Vol.6(2):332. See also Stockard, C.R. The Influence of Alcohol and Other Anesthetics on Embryonic Development. American Journal of Anatomy. 1910. Vol.10(3):369-392. 572 Stockard followed one of the senior-most faculty members at Cornell Medical, Alexander Lambert, in discussing the impact of parental alcoholism on progeny. Lambert had less than five years prior reviewed the evidence tying parental alcoholism to idiocy and epilepsy in subsequent generations. Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? p.7-10.

277

This new focus was accompanied by a new experimental animal, the guinea pig, which required a shorter logical leap to man than did the minnow.573 Through a series of mating experiments, Stockard, with the aid of his assistant, Dorothy Craig, found that if even one of the parents had been exposed to alcohol, the pairing produced higher rates of infertility, stillbirths, developmental defects, and early mortality than control animals.574

In subsequent tests, Stockard found that these degenerative effects could be perpetuated.

When the teetotal offspring of the paternal test were mated to normal animals, the resulting F2 progeny were still much more prone to developmental arrests and early mortality than were control animals, even a generation removed from the influence of alcohol. Stockard believed this line suffered from a true “germinal taint” brought on by chronic exposure to alcohol and surmised that “[w]hen the reproductive glands and germ cells became injured in this way they gave rise to offspring showing weak and degenerate conditions of a general nature, and every cell of these offspring having been derived from the injured egg or sperm cell are necessarily similarly injured and can only give rise to other injured cells and thus the next generation are equally weak and injured and so

575 on.” By 1916, with the arrival of an apparently vulnerable F3, Stockard concluded, in

573 Stockard seems to have picked up the idea to use guinea pigs from the Finnish medical scientist Taav Laitinen, whom Stockard rebuked for being “more interested in the problem of the misuse of alcohol than in the scientific study of the influence of injurious substances on the offspring.” Nevertheless, Pauly has shown that Stockard’s own research with alcohol, though carefully executed, was intimately tied to the social problem of the misuse of alcohol. Quotation from Stockard, C.R. An Experimental Study of Racial Degeneration in Mammals Treated with Alcohol. Archives of Internal Medicine. 1912. Vol.10:372; Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? p.9-12, 19-20. 574 The crucial test was mating an alcoholic male to a normal female, since the defect could only have been carried with the male gamete and not introduced in the inter-uterine environment. 575 Ibid. p.677.

278 terms evoking Morel’s grim prophecy of a half century earlier, that “the alcoholic race seems at this stage of the experiment about to fade out in the fourth generation.”576

As Pauly observed, Stockard’s work “signaled the possibility of a major convergence in biological, medical, and social thinking about the dangers of alcohol,” and the scientific, medical, and social reform communities were equally impressed with the results.577 Stockard’s results were quickly yoked to the temperance bandwagon, most notably by congressman Richmond P. Hobson in his lecture “The Great Destroyer,” while medical journals extolled Stockard’s use of animal experimentation to answer the vexed question of alcohol’s influence on progeny.578 The New York Times similarly chimed that “[t]he particular value of Dr. Stockard’s work is that he is putting all this sociological data on a definite scientific basis. Social studies are valuable, but the cry of sentimentalism and inaccuracy can be raised against persons who have a case to prove.

With the scientists it is different.”579

Pauly’s analysis revealed that even for the restrained scientist the effect of parental alcoholism on progeny was as much a social as a scientific problem. Thus, as nationwide prohibition relieved the social urgency for scientific research on the effects of alcohol, Stockard abruptly reinterpreted the results of four generations of breeding his

“alcoholized” guinea pigs. In the fall of 1920 at the Fifteenth International Congress against Alcoholism in Washington, he argued that alcohol was racially beneficial rather than detrimental. He now claimed to see a gradual improvement over the generations,

576 Stockard, C.R. A Further Analysis of the Hereditary Transmission of Degeneracy and Deformities by the Descendants of Alcoholized Animals. The American Naturalist. 1916. Vol.50(590):84. 577 Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? p.12. 578 See ibid. p.9-10. 579 “Does Alcoholism in Parents Injure Children by Heredity?” New York Times. June 9, 1912.

279 especially in the quality of the stock surviving. The improvement in the fourth generation, which he had earlier ascribed to their having only 1/16 alcoholic ancestry, he now credited to the selective effects of alcohol on gametes, culling out the weak and defective while leaving vigorous germ cells unharmed.580 To the horror of his teetotal audience, Stockard also reinterpreted the social significance of his findings in characteristically unsentimental and uncompromising terms:

There are a certain class of weak mediocre people or individuals who don’t seem to amount to anything and you want to take them on your lap- so to speak- and coddle their mediocrity, and humor them; but alcohol, if used in a eugenic way, will prevent such people from being born, and that is quite a consideration for those interested from a scientific standpoint in the progress of the world… We can’t look at this from an ethical or a humanitarian standpoint; we’ve got to consider it on a scientific basis. If you go to breed horses or dogs or cattle or pigs or any of these things, you must, and do, go at it scientifically.581 Where precisely on the road to Damascus Stockard’s conversion occurred is not perfectly clear, but its timing suggests that he had keenly honed his ability to find and exploit the social relevance of his research, all the while asserting the epistemic purity of his “scientific standpoint.” His reinterpretation of alcoholism as a symptom, rather than a cause, of degeneracy also bolstered his constitutional outlook, as he began to see degeneracy as an endogenous condition rather than one induced by some external agent.

Stockard continued to peddle his anti-prohibitionist argument in a handful of publications and lectures over the next decade, but as the ardor surrounding alcoholism waned he turned again to the wider study of constitutional abnormalities, with the goal to uncover the nature and origin of human types. To study this problem, he sought a new subject

580 Stockard had actually appropriated this argument from Raymond Pearl, this after vigorously challenging the same conclusion when Pearl initially offered it prior to national prohibition. See Pauly, P.J. How Did the Effects of Alcohol on Reproduction Become Scientifically Uninteresting? p. 15- 16. 581 Quoted from ibid. p.19-20.

280 and could find none better suited to the resolution of type and the abnormalities of constitution than the domestic dog.

Stockard laid out the scientific and philosophical justification for a study of growth types in dogs in 1923. Stockard was overcome by his luck that fanciers had prepared for him such a rich array of growth types, based, he assumed, on differences in the glands of internal secretion. “These peculiar glands,” he explained, “are the fundamental things that the breeders have unconsciously been selecting, since when they choose animals with the desired structural symptoms they also blindly choose the glandular cause. Thus, a certain gland type may be inherited and the dog breeders have produced the experimental proof of this over and over again.”582 He noted that H.

Rischbieth and Amy Barrington of the Francis Galton Laboratory had similarly suggested a likeness between dog breeds and certain human growth types, but had stopped short their analysis on the assertion that the similarities were only apparent. Stockard thought this a “vital mistake since certain animal breeds not only very closely resemble the human dwarfs but very probably arise and developed [sic.] in an exactly similar way.

The understanding of the origin and development of the one is certainly of the highest value in a biological consideration of the other.”583 Demonstrating perhaps as much his callousness as his point, Stockard pointed to “a most striking and convincing comparison” affected by “holding a fine pointed French bull dog up on its hind feet by the side of a human achondroplastic dwarf, or the identity of types is equally well shown

582 Stockard, C.R. Human Types and Growth Reactions. American Journal of Anatomy. 1923. Vol.31:273. 583 Ibid. p.264.

281 by placing the man and the dog on all fours.”584 He made similar comparisons between the exaggerated features of human acromegalic giants and St. Bernard, mastiff, and bloodhound dogs. In the case of the King Charles spaniel, he argued that it was not only

“in shape, outline, and expression almost a picture of the human midget” but also “the psychology of the two is much the same for general behavior reactions.”585 In Stockard’s view, it was axiomatic that the mind would follow the body’s constitution, and recent physiological investigations by Walter Cannon and others suggested the far-reaching influence of endocrine factors on nervous function, including behavior. The consistency of these functional states and the character of their interactions, he believed, defined distinct types, whether in dogs or man.

Stockard’s greatest influence in this line of thinking was the Scottish anthropologist Sir Arthur Keith, whom he had met during Keith’s American sojourn in

1915.586 Keith had made hormones fundamental to growth and evolution, most famously characterizing human races according to their prevailing endocrine mechanisms. As a corollary to his formulation of hormonally differentiated races, he defined racial prejudice, not as a social construct, but as an instinct, “at the very root of human mentality,” for the purpose of “safeguarding the purity of a race.”587 Civilization appeared throughout Keith’s anthropological work as a force antagonistic to “Nature’s plan of evolution.” Its modern political arrangements threatened to break down those

584 Ibid. p.268. 585 Ibid. p.271. 586 Keith recalled in his autobiography, “Of all the anatomists whom I met in New York, Dr. C.S. [sic.] Stockard (1872-1939) of Cornell University made the deepest impression on me. He, too, had realized that the hormone theory helped in the solution of many anatomical problems.” Keith, A. An Autobiography. London: Watts & Co., 1950. p.379. 587 Keith, A. The Evolution of Human Races. Journal of the Royal Anthropological Institute of Great Britain and Ireland. 1928. Vol.58:316.

282 tribal affinities that preserved race distinctions and, ultimately, the endocrine profiles fundamental to those differences. “We moderns,” he explained, “are like hill sheep turned into fenced fields with all of our wandering instincts still grafted onto our original nature. As in them, our instincts are at war with our surroundings.”588 On the question

“Is man a domesticated animal?” Keith reiterated that civilization “is a powerful selective agency… favoring those who willingly obey her behests,” but ultimately determined that man’s indomitable will prevented him from ever becoming truly domesticated.589 It was likely Keith that convinced Stockard of the particular sociological value of domestication to the investigation of type, but Stockard would take the domestication analogy much further than his colleague across the Atlantic.

For Stockard, “type” was a highly evolved adjustment of the total animal to its environment. The random assortment of chromosomes in a cross of two distinct types could, thus, only produce animals “more variable and less stable than those of long purebred ancestry.”590 It followed, he reasoned, that “[w]ild species, e.g. the wolf and jackal, are more uniform in type than their domestic derivatives, dogs.”591 Stockard borrowed from popular eugenical arguments of the time, including Alfred Mjöen’s warning about

“disharmonious” crosses between races and Keith’s concerns about man’s fitness in a civilized environment, and synthesized them into a dire prediction about the effects of human self domestication at the meeting of the Association for Research in Nervous and

588 Keith, A. Nationality and Race: From an Anthropologists Point of View. London: Humphrey Milford, Oxford University Press, 1919. p.8. 589 Keith, A. Evolution and Ethics. New York: G. Putnam’s Sons, 1946. p.42,45. 590 Stockard, C.R. Experimental Production of Degeneracy in its Bearing upon Hereditary Abnormalities of the Nervous System in Heredity in Nervous and Mental Disease: A Series of Investigations and Reports. Vol.III. New York: Paul B. Hoeber, 1925. p.207. 591 Ibid.

283

Mental Disease in December 1923. “Man all evolved in the wilds together; they all started out on the same day,” he explained, evoking the primitivism that so often accompanied sustained thinking about human domestication:

In nature, wolves in a pack are all closely alike. They are up to a standard. We do not have wolf shows; that is not necessary. Every wolf would take a ribbon. But we have a dog show because dogs are domesticated, petted, pampered and protected by man, and are not selected by nature. So we have a dog show to see whether we have any dog at all. Generally we see a monster; he borders on the monstrous type. This is what domestication has done.

Anthropologically speaking, we are domestic animals, and we have suffered just as much from it as some other domesticated animals. We have removed most of the natural agents of selection which have made every man meet a standard.592

At the meeting Stockard’s colleague at Cornell Medical, the neurologist Foster

Kennedy, rebuked him for his pessimistic view of civilization. Stockard replied with hard reason: “…[I]f we strive to breed an intellectual race and a well race, we shall never do it by breeding impaired stock… I have all sympathy with the humanitarian point of view which Dr. Kennedy so beautifully expressed, but I cannot quite believe it is philosophically sound.”593 Convinced of the soundness of his argument, Stockard prepared for its demonstration in the investigation of type and personality using pure-bred and hybrid dogs.

Stockard was aware that breeding and maintaining several dog breeds and their hybrids over multiple generations would require an unprecedented outlay of resources.

592 Ibid. p.231. 593 Ibid. p.236. Kennedy was later embroiled in a controversy for advocating that defective children, from the age of five, should be relieved of the “agony of living.” See Kennedy, F. The Problem of Social Control of the Congenital Defective: Education, Sterilization, Euthanasia. American Journal of Psychiatry. 1942. Vol.90:16.

284

Still, he began his studies in earnest, “collecting material from seemingly pure-bred dogs of various breeds destroyed at the city pound” for an investigation of their glands.594 He was quickly disheartened, however, as the uncertain age and ancestry of the animals, as well as the incidence of disease among them made them unsuitable for characterizing the glands of their respective breeds. This was the first in a long series of missteps and unforeseen complications in the execution of Stockard’s grand experiment. In May 1925, however, Stockard solved the greatest of the impediments facing his project. He had sufficiently impressed the Rockefeller-backed General Education Board to secure

$20,000 for buildings and equipment and a further $125,000 for operating costs over the next five years. 595

The timing of Stockard’s proposal coincided with the changing priorities of the

General Education Board under the direction of Wickliffe Rose. The purpose of the

General Education Board had been, since its creation in 1903, to reform and support

American higher education, including medical schools. When Rose assumed leadership of the Board in 1923, he not only expanded the scope of the agency (creating the

International Board of Education), but redirected its focus increasingly toward the support of basic science rather than education. The shift was such that Abraham Flexner,

594 Nonidez, J.F. Forward in The Genetic and Endocrinic Basis for Differences in Form and Behavior as Elucidated by Studies of Contrasted Pure-line Dog Breeds and Their Hybrids. C.R. Stockard and Collaborators. American Anatomical Memoirs 19. Philadelphia: Wistar Institute of Anatomy and Biology, 1941. p.ix. 595 See Alan Gregg to William S. Ladd June 14, 1939. RG 1.1, Series 200, Box 81, Folder 977. Rockefeller Foundation Archives, Rockefeller Archive Center, Sleepy Hollow, New York (hereafter designated RAC). Stockard was elected in the same year to the Board of Directors for the Rockefeller Institute for Medical Research.

285 secretary of the agency, complained in 1927, “[The Board] is not an education board- it is mainly a scientific research board.”596

Wider changes in the Rockefeller philanthropies in the middle 1920s also seemed at first to benefit Stockard’s ambitious plans. Erwin Embree, head of the new Division of

Studies of the Rockefeller Foundation, campaigned for a new program in “human biology” that recalled the Foundation’s earlier commitment to social application of scientific knowledge. His description of the problems to be considered as part of this program reflected many of the same social concerns that Stockard had expressed in 1923:

“… problems of race relations are pressing as never before. The crowding of population is no longer a national matter but a world issue, and questions are being raised with respect to quality as well as to numbers of population groups. The complexity of modern industrial civilization is accentuating mental and nervous disorders.597 Embree’s program never fully materialized and was aborted in the 1928 reorganization of the Rockefeller

Foundation, as much for Embree’s mismanagement as for any opposition to supporting eugenical research. As historians including Diane Paul and Kersten Beihn have shown, while officers of the reorganized Foundation sought to distance themselves from the overtly political aims of mainstream eugenics, they continued to support research into human differences with the goal of better-informed social prophylaxis.598 This support was largely directed toward research into the inheritance and control of cognitive ability

596 Flexner referred to the International Education Board in the quotation above, though it describes the shift in priorities at the domestic arm under Rose as well. Quoted from Kohler, R.E. A Policy for the Advancement of Science: The Rockefeller Foundation, 1924-1929. Minerva. 1978. Vol.16(4):488. 597 Quoted from ibid. p.495. 598 Paul, D.B. The Politics of Heredity: Essays on Eugenics, Biomedicine, and the Nature-Nurture Debate. Albany, New York: State University of New York Press, 1998. p.56-59; Beihn, K. J. Psychobiology, Sex Research and Chimpanzees: Philanthropic Foundation Support for the Behavioral Sciences at Yale University, 1923-41. History of the Human Sciences. 2008. Vol.21(2):30-34.

286 and temperament. Max Mason, upon becoming president of the Foundation in 1930, produced plans for an interdisciplinary “psychobiology” program to investigate the basis of “mental health and personality.” In the same year, the grant for Stockard’s integrated study of form, function, and behavior in dogs was renewed, but not under the General

Board of Education. It was now a Foundation project. It fit well within the long sweep of Foundation interest in the social application of knowledge concerning human mentality, beginning with their support of mental hygiene in the 1910s and ending with

Alan Gregg’s research intensive psychobiology program in the 1940s.

Despite the renewal of his grant, Stockard had little to show for the first five years of financial support. By July 1930, he had been given over $162,000 and had been promised another $250,000 over the next decade. His facility in Shrub Oak was richly supplied, but making the plant operational, that is, keeping the dogs healthy and breeding, had been a much greater challenge than Stockard had initially thought. In a short piece communicated to The Collecting Net in 1931, Stockard ostensibly set out to describe the

Experimental Morphology Farm (also called the Cornell Anatomy Farm or, most often, the dog farm), but instead provided a laundry list of the challenges he and his associates had encountered there: outbreaks of internal and external parasites and distemper, learning the distinct habits and needs of multiple dog breeds, providing suitable shelter, nutritional deficiencies in patent and commercial dog foods, improperly trained and

“superstitious” kennel men, and the provision of running water, electricity, and food storage.599 In the final report of the work performed at the dog farm, one of Stockard’s associates admitted that sorting out these problems “took several years… during which

599 Stockard, C.R. An Experimental Dog Farm for the Study of Form and Type. The Collecting Net. 1931. Vol.6(10):257, 262-264.

287

Doctor Stockard freely applied the trial and error method in view of the scarcity of data on the subject.”600

At the end of 5 years of support, Stockard had produced just 3 papers as a result to the work on dogs, and 2 of these were on subjects quite tangential to the original intent of the study.601 His first major communication on the project came in 1931, as a book entitled The Physical Basis of Personality. The book, though adapted from his Lane

Medical Lectures of a year prior, was directed toward a popular rather than a professional audience. The bulk of the volume reviewed basic principles of heredity and development as they applied to Stockard’s constitutionalist view, emphasizing especially the

“relationship between a definite physical constitution and its characteristic functions and behavior.”602 He also reiterated many of the suggestive hypotheses from his 1923 papers on type and nervous and mental disease. Of sixteen total chapters, only one included any new evidence from experiments conducted at the dog farm. Even here, he was careful to remind his readers that his study “at this time is but in its beginning. So that our present discussion is more profitably confined to the question of inheritance and development of the modified structures and shapes which have appealed so directly to the fanciers of dogs, and which are so exactly comparable to important conditions in man.”603

Stockard’s description of crosses between the dachshund and the Boston terrier is typical of his analysis. These breeds were selected for the localized dwarfism they

600 Nonidez, J.F. Forward in The Genetic and Endocrinic Basis. p.ix. 601 Stockard, C.R. Inheritance of Localized Dwarfism and Achondroplasia in Dogs. Anatomical Record. 1928. Vol.38:29; idem. Rickets in Dogs as Probably Related to Sex. American Journal of the Diseases of Children. 1928. Vol.36:310-314; idem. The Presence of a Factorial Basis for Characters Lost in Evolution: The Atavistic Reappearance of Digits in Mammals. American Journal of Anatomy. 1930. Vol.45:345-377. 602 Stockard, C.R. The Physical Basis of Personality. New York: W.W. Norton & Company. 1931. p.v. 603 Ibid. p.227.

288 displayed: limited to the legs in the dachshund and to the head and tail of the terrier. The

F1 hybrid progeny were relatively uniform, bearing shortened legs like those of the dachshund parent while the head fell somewhere between the characteristic shapes of the parent breeds. The F2 progeny were much less uniform than their parents’ generation had been. Short legs appeared again in ratios indicative of a single factor dominant, while the head and tail of the Boston terrier were much more variable and were ascribed to multiple factors. Stockard could report from these crosses and a similar breeding scheme between dachshunds and French bulldogs that “localized achondroplasia in two separate skeletal regions may be inherited in completely independent fashion.”604 Without physiological or histological analyses, however, he was left to assume that the glands of internal secretion were the modifying factors, that they displayed varying levels of “disturbance,” and that these qualities were heritable. Lacking behavioral data, Stockard’s mentions of behavior were largely impressionistic, as with his description of the F1 hybrids: “… in behavior, they show rather the nervous, jumping, noisy disposition of the Boston terrier than the quiet and shy demeanor of the dachshund.”605 Yet he remained convinced that

“it will be found that the characteristic behavior of each breed fits in a very definite way the form and appearance of the animal and follows what we know about the modified behaviors of peculiar types of human beings… [In the way that] the quiet, determined,

604 Ibid. p.243. 605 Ibid. p.239. Stockard did achieve some behavioral data that extended beyond impression. In a cross of a basset-hound sire and a German shepherd dam, in which the hybrid progeny were kept exclusively with their mother, the young exhibited recognizable instinctive traits peculiar to the basset-hound breed (tracking with noses down and barking as they run). These traits, Stockard asserted, were “entirely unlike the reactions of their shepherd mother.” Ibid. p.227.

289 and rather severe attitude of the stocky achondroplasic human dwarf is very closely paralleled by the demeanor and voice of the bulldog.”606

Despite meager progress, Stockard still had the confidence of Rockefeller

Foundation officers in the early 1930s. Priorities at the Foundation were shifting again at this time. Warren Weaver, Mason’s protégé, fellow physicist, and pick to lead the division of natural sciences, sought to bring the rigor and methods of the physical sciences to bear on the biological problems of interest to the Foundation. Although

Stockard’s analyses did not touch directly on the biochemical basis for the genetic, developmental and behavioral phenomena he studied, their promise to uncover the relationship between these phenomena was sufficiently close to Weaver’s emphasis on

“vital processes” to win praise from his division. Frank B. Hanson, Weaver’s associate director, summarized the impression Stockard had made on Weaver and him in a 1934 interview: “S.’s program and setup make an impressive showing. The genetic-endocrine combination is an important aspect, and the fact that the work is done on such a large scale and variable mammal makes it unique.”607 Stockard may have been helped by

Weaver’s initial reliance on Alan Gregg’s greater expertise in biology, which shaped a unified agenda for the divisions of medical sciences and natural sciences under the rubric of Gregg’s “psychobiology.”608 Again, despite significant reorganization, Stockard’s research program fit neatly with both the scientific and social priorities of the Foundation.

606 Ibid. p.226. The frontispiece to the book seeks to make this point by presenting pictures of humans with exaggerated features juxtaposed with images of dogs ostensibly showing the same or like distinctive traits. 607 Hanson, F.B. Interview C.R. Stockard with W. Weaver June 2, 1934. R.G. 1.1, Series 200A, Box 81, Folder 976. RAC. 608 See Kohler, R.E. The Management of Science: The Experience of Warren Weaver and the Rockefeller Foundation Programme in Molecular Biology. Minerva. 1976. Vol.14(3):289-290.

290

Weaver’s far-reaching report to the Foundation in 1934, for instance, nearly enumerated the aspirations and assumptions of Stockard’s investigation of genes, endocrines, and behavior in domestic dogs:

Can man gain an intelligent control of his own power? Can we develop so sound and extensive a genetics that we can hope to breed, in the future, superior men? Can we obtain enough knowledge of physiology and psychobiology of sex so that man can bring this pervasive, highly important, and dangerous aspect of life under rational control? Can we unravel the tangled problem of the endocrine glands, and develop, before it is too late, a treatment for the whole hideous range of mental and physical disorders which result from glandular disturbances…? Can we release psychology from its present confusion and ineffectiveness and shape it into a tool which every man can use every day? Can man acquire enough knowledge of his own vital processes so that we can hope to rationalize human behavior? Can we, in short, create a new science of man?609

Stockard was more productive under his second Rockefeller grant. He had advanced significantly in his breeding experiments, included histological investigations to determine if endocrine glands varied in quality as well as gross morphology, and initiated more rigorous psychological tests to gauge the influence of type on behavior.

Rockefeller officers seemed satisfied that Stockard was beginning to deliver on some of the ambitious aims of his research program. In 1933, Alan Gregg commented in his diary that “[Stockard] has pretty good evidence that the principle of harmonious development as claimed by [William E.] Castle and others is not verified in several instances. The F2 generation quite frequently inherited genetic factors that seriously handicap the total organism,” the racial and social implications of which Stockard had explored a decade earlier.610 In the intervening years Stockard had refrained from such bald social

609 Quoted from ibid. p.291. 610 Gregg, A. Diary- March 11, 1933. RG 1.1, Series 200A, Box 81, Folder 976. RAC.

291 pronouncements in his published work, but his comments at the 1937 American Eugenics

Society conference on medicine and eugenics make clear that concerns about the quality of human stock under the modifying influence of civilization still motivated much of his thinking. “…human beings are not isolated from the rest of the animal kingdom,”

Stockard asserted at the conference, “If we are going to continue to live in this world and not face ultimate extermination, we must give thought to the effect of the artificial conditions of civilization, as they affect human breeding… We have never been willing to face these questions in a large enough way.”611 Undoubtedly, this is precisely what

Stockard had intended to do in his investigation of “type” in domestic dog breeds and their hybrids.

As the decade of support drew nearer to its end Stockard was under increased pressure to produce his summary and synthetic statement on the relative influence and interactions of genetics and endocrines in the formation of type. Support from the

Rockefeller Foundation was still largely steadfast, but Alan Gregg’s assessment of the project in May 1938 expressed some doubt about Stockard’s management of the project, noting especially his apparent inability to delegate and unwillingness to cooperate with or train others.612 In October of 1937 Stockard set about summarizing a dozen years’ work, but almost as soon as he began, his health started to fail him. He had contracted lung cancer and died in April 1939.

Workers remaining at the Dog Farm were eager to learn if the project would be continued in the conspicuous absence of its founder. A week prior to Stockard’s death,

611 Summary of the Round Table Conference Held at the New York Academy of Medicine, Wednesday, April 21, 1937. American Eugenics Society Papers 575.06.Am3, Folder AES-Conference on Medicine and Eugenics. APS. 612 Gregg, A. Diary- May 28, 1938. RG 1.1, Series 200A, Box 81, Folder 977. RAC.

292 however, Alan Gregg and William Ladd, Dean of Cornell Medical, had already begun planning for the dissolution of the farm.613 Stockard’s death, Gregg explained with more candor than he had shown previously, “resemble[d] in its effect the death of a lone farmer at harvest time… Stockard was a dominating and forceful character more impressive than tolerant, more likely to direct than to develop his juniors and disposed to dispute rather than collaborate with his colleagues. It is necessary to know this in order to understand the major failure of this project- that Stockard trained no one and no group competent to carry on the work he started. His response to the opportunity was to take everything in his own hands, his reaction to overloading was a stubborn and energetic insistence in just the same direction.”614 Gregg determined that the funds remaining on the grant should be used to bring Stockard’s unfinished manuscript to publication. The manuscript as

Stockard left it was, one reviewer commented, surprisingly “well put together, though it lacks [a] final chapter as summary.” Stockard’s associates in psychology, W.T. James and O.D. Anderson contributed sections on “Morphologic Form and its Relation to

Behavior” and “The Role of the Glands of Internal Secretion in the Production of

Behavioral Types in the Dog,” respectively, and the Wistar Institute published the collected work in 1941.

Stockard’s posthumous opus, The Genetic and Endocrinic Basis of Form and

Behavior, revealed the grand scale of his operations and perhaps more clearly the enormity of the problem he set before himself. From the beginning, he wished that it would “be clearly understood that our aim is to give an experimental analysis of

613 Gregg, A. Diary- March 28, 1939. RG 1.1, Series 200, Folder 977, RAC. 614 Gregg, A. Cornell University- Stockard Dog Farm June, 1939. RG 1.1, Series 200, Box 81, Folder 978. RAC.

293 constitution in a comprehensive manner and not simply to report on the genetics of isolated characters among dogs… the various problems concerned are so closely interrelated that they cannot be fairly considered or properly analyzed in any other way.”615 No problem could be dealt with in isolation and the range of possible explanations for a given phenomenon was dizzying.

For the analysis of these problems, Stockard sought an array of extreme features in his dog breeds, reasoning that “[i]f a correlation exists between the inherited qualities of endocrine secretions and peculiarities of structural features, no doubt such associations can be more readily recognized and analyzed in fully manifested cases.”616 He included in his crosses short-legged breeds such as the dachshund and bassethound, giant breeds like the Great Dane and St. Bernard, the svelte saluki, and short-faced breeds of various size such as the English and French bulldogs, the Pekingese, the Boston terrier, and the

Brussels griffon. These anomalous types, Stockard recognized, could only be considered anomalies by reference to a standard or archetypal dog. For this role, he selected the

German shepherd.

The reasoning Stockard gave for this choice is both puzzling and telling: “This dog, in its general type, structure, functions and behavior, deviates very little from a standard or control type.” The shepherd was, of course, meant to be a standard and yet had to be measured itself against a higher standard, the wolf. Stockard had chosen the shepherd because “no other breed of dog differs so little from the present living species

615 Stockard, C.R. Genetic and Endocrinic Basis. p.8. 616 Ibid. p.18.

294 of wild Canidae.”617 Still, he admonished that it must be “…realized that these animals are reared and kept under the most abnormal conditions… These dogs are not usually permitted to breed normally as wild animals would, have too little exercise and freedom, and eat an artificial diet, all of which might tend to modify their endocrine glands.” This, however, only helped the comparison to the other dog breeds, “since all the animals are kept under the same uniformly regulated regime.”618 At one and the same time the

German shepherd was conspicuous in its wildness and familiar in its domesticity; it was both natural and unnatural. Even as Stockard began recording rigorous anatomical and histological measures of difference, he maintained a separate, more arbitrary scale of deviance upon which dog breeds were judged more or less “wild.” He was convinced that these differences and the concept of type they helped to define had a basis in the biology of the animal, not in culture. By identifying “normal” as primordial in the dog, he could also casually endorse a standard model of man very different than city pent modern man: “No other dog [than the shepherd] fits so closely into the pattern which one would imagine for the early hunting and fighting associate of prehistoric man.”619

For Stockard, variation was not neutral. He referred to mutations, sports, and modifications of development as “violations of their original type and size.” Thus, his assertion that “man and the dog probably show the widest and most highly modified

617 Ibid. p.40. Stockard was not alone in making this likeness. The breed’s founder, Max von Stephanitz asserted that the shepherd’s ancestry was derived solely from the wolf, while other breeds possessed some jackal or mongrel form of ancestry. Through his management of the Verein für deutsche Schäferhund (Society for the German Shepherd Dog), the shepherd became an emblem of racial purity and the wisdom of natural (as opposed to social) order. See Sax, B. Animals in the Third Reich: Pets, Scapegoats, and the Holocaust. New York: Continuum International Publishing Group, Inc., 2000. Esp. p.83-84; Skabelund, A. Breeding Racism: The Imperial Battlefields of the “German” Shepherd Dog. Society and Animals. 2008. Vol.16:354-371; Wipperman, W. and Berentzen, D. Die Deutschen und ihre Hunde: Ein Sonderweg der Mentalitätgeschichte? Munich: Seidler, 1999. 618 Ibid. p.42. 619 Ibid. p.40.

295 deviations from stem type” cannot help but reflect a normative judgment on the human agency that brought both species to their current state.620 As he had done nearly two decades earlier, Stockard maintained that hybridization was the primary culprit and produced a specious and sweeping history to support his assertion that what was true of dogs was also true of man:

Since prehistoric time, hybrid breedings of many kinds have occurred at random among the different races of human beings. Such race crossings may have tended to stimulate mutations and genetic instability, thus bringing about freak reactions and functional disharmonies just as are found to occur among dogs. The chief difference has been that in dogs a master hand has selected the freak individuals according to fancy and purified them into the various dog breeds. No such force regulates the mongrel mixing of human beings… Mongrelization among widely different human stocks has probably caused the degradation and even the elimination of certain human groups, the extinction of several ancient stocks has apparently followed the extensive absorption of human slaves… [A] very definite correlation between the amalgamation of the whites and the negroid slaves and the loss of intellectual and social power in the population will be found. The so-called dark ages followed a brilliant antiquity just after the completion of such mongrel amalgamation.621

Stockard believed that he had the necessary evidence for a demonstration of his concept of type and the perils of hybridity. While the gross morphology of the endocrine glands failed to correlate with any particular type, histological analyses of the glands pointed the way to characteristic anomalies, such as the irregular appearance of thyroid follicles in short-muzzled breeds.622 In a cross between short- and long-muzzled breeds,

Stockard found striking disharmonies between the upper and lower jaws, producing both

620 Ibid. p.22,19. 621 Ibid. p.37-38. 622 Ibid. p.423-440. This evidence was not unequivocal, however, as the thyroid was found to be highly variable within and between breeds. Most of the short-muzzled breeds had “hyperactive” thyroids to match their nervous and excitable temperaments. However, in the example of the lethargic English bulldog, Stockard was forced to claim that this hyperactivity of the thyroid epithelial cells was only apparent and not real. p.427.

296 overshot and undershot mandibles. Crosses between breeds with loose skin, such as the

English bulldog and basset-hound produced in the second hybrid generation individuals with skin “sufficient to cover twice [their] body size… This disharmony,… although differing in many ways from the misfit between upper and lower jaws…, is in some respects a comparable structural mixup resulting from hybrid confusion of genetic qualities.” To these he added the “rabbit-like” stance of the dachshund-Brussels griffon hybrid, the crowded teeth and bulging eyes of short-faced breeds, and others. “The one character common to all these breeds and hybrids in which structural misfits occur,”

Stockard explained, “is an abnormality of the pituitary gland.” He was able to minimize the differences between these phenomena by arguing that they were affected not by the random appearance of distinct mutations, but by modifications to a more basic and relatively ancient endocrine system, which underlay all growth and expression. That these disharmonies were definitely heritable, Stockard argued, “…may mean that the related peculiarity of the pituitary is the primarily inherited character, while the structural derangements are secondary results due to the failure in harmonious growth regulation by the genetically modified pituitary gland.”623

The crucial behavioral side of Stockard’s study was the least advanced of his research objectives at the time of his death. Still, Stockard’s appointed assistants, W.T.

James and O.D. Anderson, students of the Pavlovian psycho-physiologist Howard

Liddell, were able to report from their conditioned salivary and avoidance reaction experiments: “We know what is meant by a pure constitutional type, and have demonstrated how they have arrived. We have analyzed the behaviors of hybrids

623 Ibid. p.330, 334-335.

297 obtained by crossing widely differenct [sic] types and shown how physiological form, glands, and behavior vary among these dogs.”624 They were able to achieve this in part by focusing on the wide view of the integrated organism, referring to specific behaviors

“only as an indicator of the total behavioral complex… In the present experiments the interest is rather in how the reflex phenomena perform as a unified pattern, and how they differ among dogs, and in the total behavioral picture of each dog.”625 Thus in James’ attempt to link behavior and morphology, he did not use any of the structural peculiarities

Stockard had studied, but designated two body types according to a general body shape index derived from the relative height and width of the animal’s chest. He then correlated the wide- and narrow-bodied types he identified with lethargic and active temperaments, respectively- just as Stockard had done with his lateral (brachycephalic, or hypothyroid) and linear (dolichocephalic, or hyperthyroid) human types in 1923.626

James and Anderson largely reinforced Stockard’s notion of type as a true breeding complex of interrelated form and behavior in the execution and interpretation of their experiments on the behavior responses of purebred and hybrid dogs.

In 1942 an aging William E. Castle reviewed Stockard’s posthumous volume for the Journal of Heredity. As might be expected, Castle seized on the genetic lessons from

624 James, W.T. Memorandum on the Cornell Anatomy Farm. R.G. 1.1, Series 200, Box 81, Folder 977. RAC. Stockard seemed willing to delegate to his associates in the behavioral work to a degree he was not prepared to do with the morphological or physiological work. Nevertheless, Gregg reported that during Anderson’s tenure at the Dog Farm, Stockard had “treated him like a doormat.” See Gregg to C.C. Little July 17, 1945. R.G.1.2, Series 200A, Box 134, Folder 1190, RAC. It is worth noting that despite Stockard’s shared interest with Whitman and Small in the importance of adaptation to environment, he allowed behavioral experiments to be performed that made no pretense of reproducing behaviors natural to the animal. His interest, though, was not in the behaviors themselves, but in the ability to characterize behavioral types and, if possible, to disassemble these complexes by crossing. 625 James, W.T. Section VI Morphologic Form and Its Relations to Behavior in Stockard, C.R. and Associates. The Genetic and Endocrinic Basis for Form and Behavior. p.530. 626 Ibid. p.598-601.

298

Stockard’s work including the hereditary basis of structural deformities like achondroplasia, the shortened face of certain breeds, “screwtail,” ear form, hair length, independent assortment of the upper and lower jaws, among others. He treated these modifications largely independently, noting only in passing Stockard’s conclusion that

“gene mutations in the chromosome are responsible for changes in body type but they may and doubtless do act through the agency of hormones produced in endocrine glands.” Castle indicated that Stockard had not demonstrated the endocrine basis of achondroplasia as he had not corrected the purported endocrine imbalance “by injection of hormones or ingrafting [sic.] normal endocrine glands, a procedure which restores normal form and function in a race of hereditarily dwarf mice studied by Snell,

MacDowell, and Smith” In his own interpretation, Castle asserted the primacy of the mutation itself and reserved judgment on the action of the gene.627 He had, though, larger objections.

“Stockard’s experimental studies are a model of painstaking thoroughness,” wrote

Castle, “but when he allows himself to expand beyond their scope he is less restrained.

Thus he offers the suggestion, without any evidence which justifies it, that racial degeneration among humans is a necessary consequence of race mixing.”628 More than a decade earlier he had discredited Alfred Mjöen’s analogy between domestic rabbit

627 On the point of achondroplasia, Stockard had also reserved judgment in an early section of the book (p.146), but in later sections tries to distinguish between the relative influence of genetics and endocrine constitution in a series of crosses between breeds with localized chondystrophies. Stockard’s general conclusion on these influences was ambiguous, but is in line with the conclusion Castle derived from the work. See Castle, W.E. Dogs and Human Crosses: A Review of The Genetic and Endocrinic Basis for Differences in Form and Behavior by Charles R. Stockard and Associates. Philadelphia, Wistar Institute of Anatomy and Biology, 1941. Journal of Heredity. 1942. Vol.33(7):249-252. 628 Ibid. p.251-252.

299 crosses and human miscegenation, and now he alleged a “mental bias” in Stockard’s work that not only influenced his overzealous interpretations, but led to basic

“misstatement[s] of fact.”

The example Castle shared was Stockard’s claim that “During recent years wild stocks of rabbit have been crossed in numerous ways under domestic conditions. Some of these stocks have lately been reported by Greene, Hu, and Brown (’34) of the

Rockefeller Institute to give rise to structural distortions and deficiencies comparable in many ways with those known for man or dog.” That the rabbits were wild gave a sense of primitive purity to the animals, which was subsequently corrupted by racial crossing.

However, as Castle pointed out, the rabbits had not been recently wild, but were derived from common European rabbit stock, domesticated some 1000 years. Castle was personally familiar with these stocks from having worked with “two of the most striking

‘structural deficiencies’ mentioned by Stockard as originated in the Rockefeller Institute experiments, namely dwarf and brachydactyl.” He explained that “[b]oth are true gene mutations, recessive in inheritance, such as put in an appearance when a race of animals is inbred so as to uncover them. There is no necessary antecedent period of race crossing in rabbits or in other domestic animals, such as rats, mice, guinea pigs, cattle and horses, before gene mutation occurs. It would be more correct to state that inbreeding rather than outcrossing, occasioned the appearance of these degenerative changes. But their causation is not to be ascribed to either process, but to spontaneous gene mutation.”629

Stockard believed that gene mutations might be the proximate cause of the monstrosities he encountered, but the ultimate cause could not be spontaneous. Even as

629 Ibid. p.252.

300 he sought the precise biological mechanisms for these degenerations, he had in mind that they were rooted in “violations” of the pure and proper order of nature. They were consequences, not random events. Domestication spoke to Stockard’s concerns about constitution- the correlation of parts, especially of mind and body, into an adaptive whole- principally by demonstrating what would happen in the absence of the factors responsible for maintaining this balance. From the earliest conception of Stockard’s dog project, domestication served as a model of deviance from an idealized type. Thus, by use of a value-laden analogy accompanied by a virulent racism, hybrids between formerly “pure-line” dog breeds became an expression of Stockard’s sociological concerns about the breakdown, under civilization, of biological barriers to miscegenation.

At the beginning of Stockard’s investigation of type among domestic dog breeds, the constitutional viewpoint was at its height in American biomedicine. By his death in

1939 the outlook was in decline, as social scientists increasingly explained group differences by reference to culture and many biologists became more interested in the differences within groups than between them. The constitutional viewpoint was more and more burdened by its frequently racist and eugencist trappings, especially with the polemical work of the late leader of constitutionalism, William Sheldon.630 It became increasingly isolated in extra medical fields like psychiatry where it was still crucial to explaining ties between the body and mind. The publication of Stockard’s posthumous volume scarcely registered in the scientific world of 1941.

No one was more discouraged by the outcome of Stockard’s study than Alan

Gregg. He deemed the project “an acute disappointment,” not for any perceived excesses

630 Tracy, S.W. An Evolving Science of Man. p.178.

301 of interpretation or anachronistic views, but rather for the “scattered and inconclusive state of the work” at the time of Stockard’s death.631 Stockard had fallen short of his chief aim to demonstrate the mutual determination of mental and physical type through his investigation of genetic and endocrine factors in dogs. Gregg had for years shown a keen interest in the biological basis of mental traits and shared a sociological concern with Stockard over “the economic, moral, social, and spiritual losses occasioned by the feebleminded, the delinquents, the criminally insane, the emotionally unstable, [and] the psychopathic personalities” whose numbers seemed only to grow.632 His promotion to director of the recently renamed Division of Medical Sciences (significantly, the former

Division of Medical Education) in 1930 precipitated a steep decline in funding for medical education and a subsequent increase in support for basic research in

“psychobiology.”633 Since the middle 1920s, Gregg had also closely followed the development of “the so-called constitutional school of medicine” in part because he saw

“possibilities in this field which might be of considerable interest in the study, not only of disease and its incidence, but of the behavior of individuals according to certain type in which there may be demonstrable correlation between morphological characteristics and function and conduct.”634 It is unsurprising, then, that Gregg had taken a conspicuous interest in Stockard’s investigation into these very questions.

As early as 1934, Gregg had suggested to Stockard the possibility of breeding a uniform dog for experimental purposes, along the lines of what the Wistar Institute had

631 Gregg, A. Cornell University- Stockard Dog Farm 632 Quoted from Paul, D.B. The Politics of Heredity. p.60. 633 Ibid. p.55. 634 Gregg to Walter B. Cannon March 5, 1936 RF, RG 1.1, Series 200A, Box 77. RAC.

302 achieved with the Norway rat.635 In a letter sent in the fall of 1941, Gregg’s old Harvard colleague and Director of the Roscoe B. Jackson Laboratory in Bar Harbor, Maine, C.C.

Little proposed a similar idea, breeding a standard line of dogs for cancer research.

Gregg was supportive in his reply, but by this time he had begun to consider what else might be accomplished by selective breeding of the dog, especially in connection with his primary interest: a demonstration that intelligence was limited by heredity rather than by its cultivation through education. Less than a month after Stockard’s posthumous work appeared in print, Gregg had proposed breeding dogs of exceptional intelligence and pleasant temperament. Such dogs, he reasoned, would not only convince the scientific community of the heritability of mental traits, but could be sold as pets to apprise the public of this fact and perhaps to soften opinion toward animal experimentation.636 In a similar letter to Robert Yerkes more than two years later, Gregg reiterated the value of using dogs: “…nobody cares much whether rats are intelligent or stupid, and except for their depredations we give little attention to the behavior of rats. If it were possible to develop a line of small dogs of conspicuous intelligence… we should have a demonstration of the rule of inheritance of intelligence which would carry interest and possibly conviction to a very large number of persons.”637 Both recipients agreed with

635 Gregg, A. Cornell University- Stockard Dog Farm 636 Gregg to Little November 12, 1941. RF, RG 1.1, Series 200D, Box 143, Folder 1775, RAC. See also Gregg to Little January 3, 1944. RF, RG 1.1, Series 200A, Box 133, Folder 1189, RAC. 637 Gregg to Yerkes February 24, 1944. Robert Mearns Yerkes Papers. Ms. 569, Series I, Box 22, Folder 400, YUL.

303

Gregg about the human sociological value of this study and did not fail to see the connection to Stockard’s work.638

Historian of science, Diane Paul explains that from the beginning of the study,

Gregg had been deliberate about its social agenda. He had initially named Oscar

Anderson, Stockard’s former associate and a psychologist interested in genetics, as a potential lead investigator, but that post ultimately fell to John Paul Scott, a geneticist with an interest in behavior. At the time of his appointment, Scott prepared a brief memorandum to explain his own concept for the project. Gregg felt that Scott’s approach was largely congruent with his own, but lest there be any confusion, Gregg felt the need to “risk a short paragraph in the hope of defining as clearly as I can, not the exact procedures nor the immediate specific research objectives [these he would leave to

Scott], but rather certain needs that the fields of both medicine and education experience.” Gregg summarized his point at the end of the paragraph: “…if in a decade or two we could show the role of selective mating in creating a dog of high intelligence and marked social stability the influence of such findings upon educational theory would be considerable and in exactly the right direction since they would supplement the present absurd over-emphasis on education all by itself without reference to the capacities of the creatures to be educated.”639 In the following year the Foundation funded a conference at the Jackson Laboratory on “Genetics and Social Behavior” with a body of invitees distinguished in the field of behavior studies. Gregg had also invited a handful

638 Little to Gregg January 6, 1944 RF, RG 1.1, Series 200A, Box 133, Folder 1189. RAC. See also Yerkes to Gregg March 31, 1944 Robert Mearns Yerkes Papers, Ms. 569, Series I, Box 22, Folder 400. YUL. 639 Gregg to Little March 16, 1945. RF RG 1.1, Series 200A, Box 134, Folder 1190, RAC.

304 of journalists, whose role as the “Committee on Social Interpretation” was to reveal to the public not only the results of the study but their meaning.640

There was also a deliberate effort not to repeat the mistakes of Stockard’s project.

Scott, for instance, was encouraged to collaborate with and train numerous investigators, one of whom, John L. Fuller, ultimately co-authored the major reports of the study. Scott and Fuller explained that Stockard had been too convinced that the glands were at the root of various breed differences and because his experiments were “not designed to test a Mendelian hypothesis, his results were inconclusive… [W]e decided to set up our experiment in the broadest possible way, realizing we could not predict the important results in advance… Both from this viewpoint and the simple statistics of life insurance tables, we knew it was imperative that this project be a cooperative venture, not a one- man show.”641 Gregg, no doubt, made sure of this.

Scott and Fuller confronted other ghosts of Stockard’s study without recalling his work. Like Stockard they paused on the question of why domestic dog breeds were so variable, and what this meant for their human counterparts. They began with a similar hypothesis, that due to the greater number of generations that dogs have experienced since becoming domesticated “the genetic consequences of civilized living should be intensified in the dog [versus the human]… In short, the dog may be a genetic pilot experiment for the human race.”642 They further supposed that because potentially harmful mutations were deliberately preserved in dogs, “[w]e might then conclude that

640 Paul, D.B. The Politics of Heredity. p.68-72. 641 Scott, J.P. and Fuller, J.L. Genetics and the Social Behavior of Dogs. Chicago: University of Chicago Press, 1965. p.6-7. 642 Ibid. p.397.

305 the dog is genetically weak compared to the ancestral wolf, and that we ought to attempt to restore the uniform appearance and greater vigor of these ancestral animals.”643 Unlike

Stockard, though, their emphasis was not on breed specific or individual differences, but on social behavior. In fact, they found that breed differences in intelligence and temperament were not consistent and seemed to be situational rather than fixed by heredity. Even dogs that performed well in their battery of psychological tests would often do so in different ways, suggesting that “nothing like the general-intelligence factor sometimes postulated for humans” could be found, at least in dogs.644 While Scott and

Fuller did assume that uniformity follows natural selection, they did not correlate this uniformity with purity or perfection. They concluded that “the development of all-round capacities to meet a variety of environmental conditions leads not to a superb development of these capacities but to a balance between them… This suggests that the idea that natural selection will produce a super-man or super-animal of any sort is an unobtainable myth… The super-man is not to be found as an individual but as a well- developed human society.”645 They frequently cited Theodosius Dobzhansky and others who asserted the hidden extent and importance, both biological and sociological, of diversity within populations. From this viewpoint, very different from that held by

Stockard more than 25 years earlier, Scott and Fuller concluded “that a relaxation of natural selection in the dog has not produced a continuing process of degeneration but

643 Ibid. p.401. 644 Quoted from Paul, D.B. The Politics of Heredity. p.73. 645 Scott, J.P. and Fuller, J.L Genetics and the Social Behavior of Dogs. p.403.

306 rather has set up a new condition of genetic balance which permits a wider degree of variation.”646

As Diane Paul explains, Gregg was again disappointed in the results of a decade- long study of inherited behavior in dogs, not necessarily for Scott and Fuller’s denial of the degenerative consequences of domestication, but more for their inability to support the social agenda he had laid out at the beginning of the investigation. Nevertheless, the

Rockefeller Foundation and other patrons continued to support projects straddling the border between biology and psychology and “were usually able to identify scientists whose views were congruent with their own,” i.e. strongly hereditarian and focused on mental traits.647 Gregg’s psychobiology program (later renamed psychiatry) remained a robust source of support for biologically oriented behavior research until his retirement in

1954. He had funded Clyde Keeler’s study of pigment gene pleiotropy, which had begun with the observation that coat color genes seemed also to determine temperament and the hypothesis that domestication had been the result of a relative few single gene mutations rather than a centuries long process of sedulous selection.648 Gregg and other officers at the Rockefeller Foundation had also supported work in physiological psychology performed at the Phipps Psychiatric Clinic at the Johns Hopkins University.649 One of

646 Ibid. p.408. 647 Paul, D.B. The Politics of Heredity. p.74-75. As evidence of Gregg’s disappointment, Paul notes that “Had the results been more to Gregg’s liking, there existed a committee to ensure that its message was brought to the public. As it turned out, the study was effectively buried.” 648 Keeler, in private correspondence, had also advanced the domestication-civilization analogy. In a letter to Robert Yerkes, Keeler suggested that as with his rats and domestication, “civilization rides forward on recessive mutant genes.” He argued, though, that for an animal to be “level headed and sluggish” was requisite of civilization rather than an unfortunate consequence of it. See Keeler to Yerkes October 8, 1941. Robert Mearns Yerkes Papers. Series I, Box 28, Folder 529. YUL. 649 Gregg’s Medical Sciences Division ultimately funded the Phipps Clinic for 15 years, between 1933 and 1948, in the amount of $529,000, with an additional $100,000 terminal grant to aid Johns Hopkins while it assumed responsibility for the Clinic. Richter received a portion of this funding,

307 the chief beneficiaries of this largesse, the psychobiologist Curt P. Richter, later became perhaps the greatest midcentury explicator of the domestication analogy in American psychology.

Domestication and Decline: The Degeneration Thesis of Curt P. Richter

Curt Richter’s childhood and early education would seem to have prepared him more for life as an engineer than for a career in psychology. Indeed, his German immigrant father had just such plans for Curt, ostensibly so that he could take over the family business, manufacturing iron and steel parts for use in construction. Some of his earliest memories were of the factory in Denver, playing with tools, shaping metal, and constructing widgets, but this era of free pay was cut tragically short. Curt was only eight when his father was fatally shot in a hunting accident, and he remembered little of the man, save his stern discipline and his desire for Curt to attend a German Technische

Hochschule. Curt’s mother kept the factory afloat in her husband’s absence, and Curt proved similarly resourceful in fending for himself. Except for his athletic prowess, the young Richter did not fare well in school, part of a pattern of aimlessness that

along with his fellow psychobiologist at the Phipps and leader of the Pavlovian Laboratory, W. Horsley Gantt. Despite the lack of existing correspondence between the two, Richter seemed to have enjoyed a close relationship with Gregg. Richter wrote to Gregg’s wife upon his passing: “Your husband was my guardian angel for so many years. I just wanted to tell you my gratitude for all that he did for me.” Richter to Mrs. Alan Gregg November 24, 1958. Alan Gregg Papers, MsC 190, Series II, Box 22, Folder 4. Alan Mason Chesney Medical Archives of the Johns Hopkins Medical Institutions, Baltimore, Maryland. Gantt similarly expressed psychobiology’s debt to Gregg in his essay, Retrospect and Prospect: “The laboratory which was initiated and christened by Adolf Meyer owes a great deal to the support of the Rockefeller Foundation under the aegis of Alan Gregg, who has been responsible for the incubation of many eggs which would have otherwise perished from the cold.” See in Gantt, W.H. (ed.) Physiological Bases of Psychiatry. Springfield, Illinois: Charles C. Thomas, 1958. p.13. Richter was subsequently funded through the Rockefeller International Health Board for his project on Rodent Ecology and Control, which was subsequently transitioned to the School of Hygiene and Public Health also at Hopkins.

308 characterized his early life. Without reflection, Richter followed his departed father’s wishes and left home after his graduation from high school in 1912 to attend the

Technische Hochschule in Dresden. He remained there until April 1915, but growing resentment toward foreigners after the outbreak of war paired with a rapidly declining interest in engineering convinced him to return to the U.S.

Richter enrolled immediately at Harvard, not for some sudden clarity of what to do with himself, but rather because he had only the vague notion that he did not wish to take over the factory in Denver or to enter business. In quick succession, he either became disillusioned with or was dissuaded from continuing with his initial interests in international diplomacy, economics, and European history. Richter’s first brush with success came in a class on the philosophy of nature with E. B. Holt, who had been a student of William James. Holt managed to stimulate Richter’s first genuine interest in an academic subject, this in psychology, especially the ideas of Freud and the conditioned reflex. Holt, an explicator of radical empiricism, was also likely Richter’s first real introduction to the philosophy of American pragmatism. Richter followed Holt’s course with readings from Freud, Adler, Brill, and other prominent psychoanalysts, and classes in psychopathology and experimental behavior study. The latter course, taught by Robert

Yerkes, refined Richter’s interests and further convinced him to follow experimental psychology. Yerkes also exposed Richter to John Watson’s Animal Behavior and encouraged him to pursue graduate work with Watson at Hopkins. At this point in his career, Richter had almost no experience with biology, though he remembered in an autobiographical essay several lunches with the physiologist J. L. Henderson and the

309 biologist William Morton Wheeler (who imbibed the wide definition of biology used by his mentor, Charles Otis Whitman), which had a “great and lasting effect” on him.650

After graduation from Harvard and a brief and uneventful stint in the military,

Richter took Yerkes’ advice to study under Watson in his new laboratory in the Phipps

Psychiatric Clinic at the Hopkins Hospital in 1918. Richter later admitted that at this point he knew almost nothing about Watson or his approach to animal behavior study.

What he had read of Animal Behavior amounted only to “snatches here and there… [A]t that time, and for a long time later, I had no idea about what Watson meant by

‘behaviorism.’ For me ‘behaviorism’ simply meant ‘behavioural [sic.].’”651 His initial meeting with Watson hardly gave him a better sense. As Richter recalled many years later, Watson assigned Richter to a space in his laboratory and instructed him where to find lodging with medical students. He then left Richter with only the brief directive: “I want you to know that I am only interested in getting a good piece of research. You do not have to take any courses or attend any lectures. You are strictly on your own.”652

Late in his life, Richter described this as an Archimedean point in his life, one that

“changed it from a long period of inactivity to a career of intense scientific activity.”653

He couched his explanation in anachronistically ethological terms, suggesting that

“Watson’s ‘freedom of research’ announcement had released my innate gene.” He

650 Richter, C.P. It’s a Long, Long Way to Tipperary, the Land of my Genes in Studying Animal Behavior: Autobiographies of the Founders. D.A. Dewsbury (ed.). Chicago, University of Chicago Press, 1989. p.369. Wheeler, like Whitman, maintained a profound interest in animal behavior and was one of the first to introduce the term “ethology” to the United States. 651 Ibid. It should be stated, however, that Watson’s own concept of behaviorism was shifting at this point, from one that was more ecologically and evolutionarily mindful to something like the engineering ideal described by the late Philip J. Pauly, which denied the preexistence of adaptive behavioral patterns, or instincts. 652 Ibid. p.371. 653 Ibid. p.357.

310 further noted, “There is probably real irony in the fact that Professor Watson, the great advocate of behaviorism and the negator of all hereditary beliefs, should be the one who released my gene, by entirely hereditary mechanisms.” The irony was only apparent as, at this point, Watson had not repudiated instinct. Nevertheless, it is true that from this point Watson moved steadily away from the concept of instinct, while Richter moved deeper and deeper into the nature of innate behavior. As Richter recalled, “From the beginning of my studies on animal behavior in Watson’s lab [which began with Watson placing a cage containing 12 albino rats in Richter’s lab space without instructions], my interest focused entirely on what animals do on their own, that is, their innate behavior, not what they can be taught to do.”654

This divergence was not only the result of Watson’s hands-off approach to mentorship; it was further affected by Watson’s removal from Hopkins following an extramarital affair less than two years after Richter’s arrival. Richter had imbibed

Watson’s insistence on experimental rigor, but his primary influence at Hopkins was not the man who inspired him to enroll there, but Adolf Meyer, director of the Phipps Clinic and one of those instrumental in Watson’s ouster.655 Meyer was one of the chief advocates for psychobiology in the United States. Psychobiology, in Meyer’s estimation, finally made psychology objective by eliminating the “medically useless contrast of mental and physical” and grounding behavior in the terms of adaptation and response of

654 Ibid. p.377. 655 Meyer, who had initially collaborated with Watson, became increasingly suspect of him, especially as the orthodoxy of behaviorism began to harden and as Watson applied its principles to human psychiatry. For more on the interaction of these figures, see Leys, R. Meyer, Watson, and the Dangers of Behaviorism and Scull, A. and Schulkin, J. Psychobiology, Psychiatry, and Psychoanalysis: The Intersecting Careers of Adolf Meyer, Phyllis Greenacre, and Curt Richter. Medical History. 2009. Vol.53:5-36.

311 the total organism to its surroundings. “What is important to us,” he explained, “is the activity and behavior of the total organism or individual as opposed to the activity of a single detached organ. It is more than cerebration; we must take our domain broadly as behavior and passive and constructive adaptation of the entire individual.”656

Meyer set about making a proper home for psychobiology at the Phipps Clinic. In short order after Watson’s departure, Meyer appointed Richter as research director of the newly renamed Psychobiology Laboratory. Meyer turned down offers from more accomplished psychologists, like Robert M. Yerkes, because, as Jay Schulkin suggests, the young, unpublished, and unproven Richter was more amenable, more malleable to

Meyer’s philosophical vision. As Meyer himself wrote, “I trust Richter will make safer contact.”657

Richter had already shown an interest in innate behavior and a view toward its adaptational significance in his dissertation. Although Meyer worried about Richter’s

“difficulty in getting focused,” he saw promise in the younger man’s apparent ingenuity in the laboratory, which might provide the empirical backing to Meyer’s largely philosophical construction of psychobiology. Meyer drew heavily from the pragmatists

James and Dewey, whose psychology was grounded in contemporary biology, especially the notion of the functional utility of behavior, and whose philosophy insisted that even something as abstruse as the mind could be laid bare by experiment. As Richter recalled,

“[Meyer] had a very important influence on my whole development, he was far and away the most widely-read man, the greatest scholar I had ever come into contact with. He had

656 Meyer, A. Objective Psychology or Psychobiology with Subordination of the Medically Useless Contrast of Mental and Physical. Journal of the American Medical Association. 1915. Vol.65(10):860. 657 Quoted from Schulkin, J. Curt Richter: Psychobiology and the Concept of Instinct. History of Psychology. 2007. Vol.10(4):327.

312 a great fund of information and high ideals and tolerance for all kinds of work.” This influence, especially his faith in the power of experiment, seems to have been transmitted more through ideal than example, as Richter explained, “[Meyer] had little idea of what could be worked out experimentally.”658 Nevertheless, Meyer remained a firm source of support and guidance for Richter in his first 20 years at the Phipps Clinic.

Richter rewarded Meyer’s investment in him by carrying out a prodigious number of experiments that were largely consistent with Meyer’s vision of psychobiological research. From the introduction of his first publication, Richter explained that his approach would begin with “the study of the responses of the total organism- intact- and in the situation in which it ordinarily finds itself. Here the biological aspects of the problems stand decidedly to the fore… It is the behavior of the organism that is of most interest, what the organism does, and how it works.”659 As concerned as Richter was with studying behaviors that were spontaneous and natural for the animal, he was also enchanted by the control offered by the laboratory. For him these two goals, of maintaining control and simultaneously allowing the animal freedom of behavior, were not problematic. Like Kline and Small before him, Richter designed a cage that ostensibly permitted the rat to perform natural behaviors, such as nest-building, burrowing, huddling with other rats, running, climbing, and jumping. “In our artificially constructed environment practically every variety of behavior observable in a natural environment is obtained,” Richter explained, “…and if we can judge how normal the environment is by the type of rat that it produces, the success of the method was

658 Quoted from Anne Roe interview of Curt Richter March 1952. Anne Roe Papers B.R621, Folder Richter, Curt 1, p.6. APS. 659 Richter, C.P. A Behavioristic Study of the Activity of the Rat. Comparative Psychology Monographs. 1922. Vol.1(2):2.

313 unquestionable.”660 At this point in his career, Richter intuitively identified “normal” with natural and simultaneously tried to downplay and defend the artificiality of his test cages by reference to the ostensibly natural behaviors they elicited. The need to reconcile the importance of free, unencumbered, and “natural” behavior and the rising esteem of experimental control had produced similarly peculiar mental constructs in recent years.

Nenozo Utsurikawa, for instance, while working in the lab of Robert and Ada Yerkes, defined “naturalistic data” in the most oxymoronic of terms, as those observations made

“under the natural cage conditions.”661

For the bulk of Richter’s career in the laboratory he did not dwell long on theory.

He characterized his approach as loose and tentative, an organic process by which he allowed new questions to arise out of his earlier work. From the time that Watson left him alone with a dozen rats, Richter claimed to have been driven from question to question by his observations of their spontaneous behavior.662 Kymograph recordings of every movement taken by his rats revealed a cyclical regularity to their periods of activity and inactivity, which became the basis of his seminal contributions to the concept of biological clocks. He then determined the impact of altering temperature, light, and the availability of food and water on these cycles of activity and tried to relate “this diffuse, undirected activity to one of the animal’s most important specific responses, its food-

660 Richter, C.P. Animal Behavior and Internal Drives. The Quarterly Review of Biology. 1927. Vol.2(3):341-342. 661 Credit for uncovering the Utsurikawa example belongs to Richard W. Burkhardt, Jr. Quoted from Burkhardt Jr, R.W. Patterns of Behavior. p.10. 662 In his interview with Anne Roe, he described the basic development of his research: “I was interested in what makes an organism move around and so on, and I began to set up cages to see if just through watching I could see id I could find out anything and that has led from one thing to another.” Anne Roe interview of Curt Richter March 1952. Anne Roe Papers B.R621, Folder Richter, Curt 1, p.6. APS. See also, Richter, C.P. …Land of My Genes. p.370-375.

314 seeking activity.”663 From these observations, Richter questioned what effects alcohol would have on spontaneous activity. He discovered that rats stopped consuming food in direct proportion to the quantity of alcohol they consumed, suggesting a self-regulation of caloric intake664. This result, in turn, precipitated dozens more experiments on self- regulatory, or homeostatic behaviors. There is a strong logical continuity through much of Richter’s work, and he remained mute on many theoretical issues, such as the definition of instinct. As Jay Schulkin explains, “We do not know Richter’s views on

[the precise definition of instinct] because this is not the level of discourse in science in which he was engaged. He assumed a concept of instinct and then set about demonstrating its validity.”665 There is reason, though, to doubt that Richter’s research agenda was quite as organic as he portrayed it throughout his career. His emphasis on drives was the influence of his former professor, Holt and perhaps his early readings from

Freud. His concept of homeostatic behavior was derived, of course, from Cannon and

Bernard before him. These influences undoubtedly shaped the kind of questions Richter asked of his rats. Still, it is remarkable the degree to which Richter was able to live within the image of the independent and sober experimentalist, to remain aloof from the controversies complicating experimental animal psychology at the time.

This becomes more remarkable as we see how close he came to the heart of some of these controversies. Most notably, Richter was asked to attend the 1954 conference on instinct in Paris, which became briefly the center of the debates over the merits of instinct

663 Richter, C.P. Activity of the Rat. p.3. 664 Richter, C.P. A Study of the Effect of Moderate Doses of Alcohol on the Growth and Behavior of the Rat. The Journal of Experimental Zoology. 1926. Vol.44:418. 665 Schulkin, J. Psychobiology and the Concept of Instinct. p.335.

315 as an organizing concept, as it was one of the first meetings between prominent leaders in

American comparative psychology and European ethology.

Less than a year before the meeting, Daniel S. Lehrman, at the time a relative unknown among American comparative psychologists, issued a withering critique of some of ethology’s most widely held assumptions, exposing some of the fundamental differences between American and European students of behavior. Lehrman argued that the distinction raised in ethology between innate and learned behaviors did not acknowledge the development of those behaviors, including those that ethologists claimed were innate. He further claimed that ethologists too often assumed homology in the physiological mechanisms underlying the behavior of very distantly related animals.

This thinking, he finally leveled, led to unsubstantiated leaps from the behaviors of animals to the actions of humans. As Burkhardt’s analysis has shown, nonepistemic factors played a role as critical to this emerging controversy as the epistemic differences between the ethologists and comparative psychologists. Lehrman, who was Jewish, was concerned that ethologists’ assumptions had been pressed into the service of Nordic racial mythology. He pointed to Konrad Lorenz’s oft-repeated analogy between “the effects of civilization in humans and the effects of domestication in animals,” which ostensibly revealed “the involution or degeneration of species-specific behavior patterns and releaser mechanisms because of degenerate mutations,” acquired upon the relaxation of natural selection. Lehrman clarified the social dimensions of this controversy: “[Lorenz] presents this as a scientific reason for societies to erect social prohibitions to take the place of the degenerated releaser mechanisms which originally kept races from interbreeding. This is presented… in the context of a discussion of the scientific

316 justification for the then existing (1940) German legal restrictions against marriage between Germans and non-Germans.”666 At the Paris Conference Lehrman kept up his critique of Lorenzian ethology, joined by his mentor, T. C. Schneirla, Helen Spurway

(whose demeanor was described as “frightful” by one attendee), and others.667

Richter, for his part, was mute on the controversy. Amid the heated exchanges at the conference, Richter’s remarks were limited to a comparatively dry question about experimental methodology. For his own paper, he had been asked to discuss his research on innate salt-seeking behaviors in the Norway rat. The topic (and Richter’s interpretation) aligned him somewhat more closely with the host ethologists than with his psychologist compatriots, but aroused no undue attention.668 Jay Schulkin suggested that

Richter preferred to focus on behavioral analysis rather than to weigh in on the nature- nurture debate and its implications, adding that “it is not clear that [Richter] ever thought deeply about these concepts.”669 However distant Richter wished to remain from the polemics of the Paris conference, he was already engaged in a protracted meditation of one of its central controversies.

Beginning in the middle 1940s, Richter had begun making comparative studies of wild caught and domesticated Norway rats and by the time of the conference was

666 Lehrman, D.S. A Critique of Konrad Lorenz’s Theory of Instinctive Behavior. Quarterly Review of Biology. 1953. Vol.28:354. Lehrman had included more on the political subtext of the debate in his initial draft of the article, but was persuaded to hew more closely to his scientific arguments by several prominent psychologists and the editors of the Quarterly Review of Biology. See Rosenblatt, J.S. “Daniel Sanford Lehrman, June 1, 1919-August 27, 1972.” Biographical Memoirs of the National Academy of Sciences. 1995. Vol.66:227-246. See also, Burkhardt Jr., R.W. Patterns of Behavior. p.384- 390. 667 Burkhardt Jr., R.W. Patterns of Behavior. p.392-394. 668 Indeed, in later life he described the expression of behaviors as the “release” of “genes” and identified himself more closely with zoologists (which was the background of most ethologists) than with psychologists. However, his comfort in the laboratory and use of the Norway rat were more characteristic of American psychology. 669 Schulkin, J. Psychobiology and the Concept of Instinct. p.337-338.

317 peddling the culturally freighted analogy between domestication and civilization with the aim to illustrate disquieting trends in human social evolution. With the help of colleagues, he published dozens of papers stemming from these comparative studies, including a handful of synoptic reviews. The title of the last of these, Rats, Man, and the

Welfare State (1959), suggests that the sober scientist had found a cause beyond the laboratory.

The wild gray Norway rat had early entered Curt Richter’s imagination. In work stemming from his doctoral thesis, Richter suggested that his initial preference was for observing behaviors in the wild rat under natural conditions but abandoned the plan as he became charmed by the control offered by the laboratory. War would finally introduce

Richter to the wild rat some thirty years later.

As Axis powers established a stronghold in north Africa, rodent populations swelled in North American cities deprived of red squill, a coastal plant native to the

Mediterranean and the chief source of rodenticide at the time. Concern was also raised that rats and other rodents might be employed as agents of bacteriological warfare. The effort to combat the rodent problem was centered in Baltimore, as much for its resident rat expert as for its role as a port city. The National Research Council (NRC) of the

National Academy of Sciences selected Richter to lead a new Rodent Control Program for his expertise in dietary self-selection in rats. Richter’s research revealed the Norway rat’s impressive ability to discriminate between beneficial and harmful substances, an apparently adaptive trait given the rat’s inability to vomit. He had recently determined that phenyl thiourea, a compound made famous by geneticists for distinguishing hereditary “tasters” from “non-tasters” in the human population, was highly toxic to the

318 rats in his laboratory, which consumed it readily. The NRC provided Richter with a small grant to investigate the utility of phenyl thiourea on wild rat populations, hoping the compound would prove as tasteless to these rats as it had been to the albinos in his laboratory. Richter found, however, that the wild caught rats brought into the laboratory were much more cautious of the toxin than the albinos had been. This result precipitated a search for a universally tasteless thiourea compound, but more importantly it also caused Richter to consider what other differences might exist between the wild gray and domesticated albino forms of the Norway rat and what the nature of these differences might be.670

Richter soon made it a matter of course for all investigations in his laboratory to be performed on both the domesticated and the wild form of the Norway rat. Distinctions between the two proliferated. Domesticated rats experienced much higher rates of audiogenic seizures (and died more frequently from them) than wild rats as a result of various treatments including blasts of pressurized air and being fed a diet deficient in magnesium. Body weights varied, especially in the wild caught rats, though the wild form was capable of much heavier weights than the domesticated albinos in Richter’s colony. The domesticates had smaller livers, hearts, brains, preputial glands, and adrenals relative to body weight than their wild counterparts, but larger pituitaries and thymuses. Thyroids remained the same size under domestication, but were apparently

670 The Rodent Control Program was ultimately renamed the Rodent Ecology Project after control of the program shifted from Richter’s psychobiological laboratory to the School of Hygiene and Public Health, also at Hopkins, on the belief that social architecture would be a more effective long term means of managing rodent populations than targeted poisoning programs. See Richter, C.P. Experiences of a Reluctant Rat Catcher: The Common Norway Rat- Friend or Enemy? Proceedings of the American Philosophical Society. 1968. Vol.112:403-415; Keiner, C. Wartime Rat Control, Rodent Ecology, and the Rise and Fall of Chemical Rodenticides. Endeavour. 2005. Vol.29(3):119-125.

319 less efficient as indicated by histological studies of the glands. The gonads were initially larger in the domesticated rats, but after 8 months, became larger in the wild rats. Richter attached the greatest significance to the changes in the adrenal and reproductive glands, especially for their value in explaining far ranging differences in physiology and behavior.

Richter and his associates measured adrenal function indirectly by determining the concentration of stored lipids, cholesterol, and ascorbic acid in the adrenal cortex, all of which correspond to the secretion of stress hormones from the cortex. They discovered that various sources of stress- cold temperatures, fighting other rats, swimming, irradiation, starvation, and injection with desoxycorticosterone acetate

(DOCA) or adrenocorticotropic hormone (ACTH; both hormones were administered to stimulate the synthesis or release of steroids from the adrenal cortex)- depleted lipid, cholesterol, and ascorbate levels in domesticated rats while having little effect on these adrenal markers in wild rats. Adrenalectomy, in turn, had a lesser impact on the domesticated form, requiring only a small supplement of salt to offset the sodium regulatory functions of the missing glands. Adrenalectomized wild rats, on the other hand, invariably died regardless of the amount of salt they consumed. Even when administered products of the cortex such as DOCA, cortisone, or both, these rats did not survive with any regularity. “The results of these various experiments on the adrenal,”

Richter explained, “indicate that the domesticated rat is much more able to compensate for the loss of the adrenal secretions than is the wild rat. In other words, during the process of domestication the adrenal secretions have come to play a comparatively less

320 important part.”671 Richter even speculated that the increased size of the pituitary in the domesticated rat was a compensatory adaptation, which allowed for more ACTH to stimulate the diminished adrenals.

As the relative value of the adrenals declined under domestication, Richter argued, the role of the sex glands- the ovaries and testes- grew in the life of the animal.

He found that domesticated rats reached sexual maturity earlier, reproduced sooner and at all times of year rather than only in the spring and fall, and appeared to be more fertile.

After gonadectomy, domesticated rats showed a steep decline in spontaneous running in a revolving drum, while similarly altered wild rats experienced little to no decline in spontaneous activity. Richter concluded, “[D]uring the process of domestication adrenal hormones have become less important and gonadal secretions more important in the life of the Norway rat.”672

Richter had amassed an impressive record of the changes attendant to the domestication of the Norway rat. Given his commitment to homeostasis and the wisdom of the body, these changes- anatomical, physiological, and behavioral- were, perforce, adaptive. In the wild state, Richter was fond of explaining, the rat was always in peril and ever in need: hunted by predators, fighting fellow rats, prodded by hunger and thirst.

Only the wiliest, the most aggressive, the fiercest could survive such pressures. By contrast, “[i]n the protected environment of the laboratory, the energy needed for survival, the supplying of food, water, nesting material, etc. comes from laboratory

671 Richter, C.P. Domestication of the Norway Rat and its Implication for the Study of Genetics in Man. American Journal of Genetics. 1952. Vol.4:279; originally read at the meeting of the American Society of Human Genetics, Ithaca, New York, September 9, 1952. 672 Ibid. p.280.

321 helpers. The rats do not have to do any work… In this protected environment it is the tamer, more gentle, rather than the fiercer, more aggressive, rats that survive- that is, the rats ‘fittest’ for this type of environment. Thus it is seen that ‘natural selection’ does not always operate to eliminate weaker individuals.”673 Richter recognized the contingency of fitness in the artificial environment of the laboratory, though his punctuation belies how foreign the concept of fitness for the created environment was to conventional thinking. The point seems more begrudging as he consistently characterizes the domesticated form of the Norway as less efficient, perspicacious, industrious, physiologically competent, and stable than the wild form and asserts that of the varieties arising among the domesticates, “very few if any… would have even the least chance of surviving in the wild unprotected environment.”674 Throughout the 1950s, Richter’s lectures and writings on the subject of domesticative changes turned repeatedly, and perhaps inevitably, toward man and suggested his own discomfort with a world he viewed as valuing security above freedom.

The leap to man was unproblematic for Richter. He insisted the Norway rat paralleled man “in various ways, particularly in dietary needs, geographic distribution, world population, and colony formation.”675 The disparity in the sophistication of the social experiences of rats and humans held little interest for him; it was enough that their physiologies were alike. If exposed to similar adaptive demands, he reasoned, the two species should mirror one another in response:

673 Richter, C.P. Rats, Man, and the Welfare State. The American Psychologist. 1959. Vol.14:22. 674 Ibid. Richter was so assured of this truth, he did not bother to follow through with his plans for long-term studies of domesticated varieties released into the wild. Such an effort, evidently aborted, is discussed in Richter, C.P. The Effects of Domestication and Selection on the Behavior of the Norway Rat. Journal of the National Cancer Institute. 1954. Vol.15(3):735. 675 Richter, C.P. Domestication… and… Genetics in Man. p.274.

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It seems likely that man, like the wild rat, originally lived in an environment where his fitness, hence his survival, was measured by his physical activity, aggressiveness and ability to withstand violent change. But with the growth of communities and the consequent increase in daily peace and stability for the individual- a transformation like that undergone by the rat in his introduction to the colony life in the laboratory- the qualities of physical strength, fierceness and aggressiveness would no longer be at such a premium. Hence would result the increase and perhaps even the predominance of the progeny of the weaker, milder, “better adjusted” individual.676

Richter acknowledged that it was not possible to compare the domesticative changes in primitive and civilized man as he had done with wild and domesticated rats, but offered the suggestive contrast of Australian aborigines, whose constant striving and hardship had left them in strikingly good health, and the inhabitants of the modern welfare state, “protected in practically every situation from the cradle to the grave” and yet plagued by any number of “untoward happenings.”677 Among these ill consequences were increased rates of chronic disease: rheumatoid arthritis, skin diseases, asthma and other hypersensitivity disorders, diabetes, mental diseases, hypertension, arteriosclerosis, and cancer. Replacement hormones could reverse or mitigate many of these conditions, but only temporarily, thus suggesting the underlying deficiency of the glands.678 He

676 Richter, C.P. Domestication of the Norway Rat and its Implications for the Problem of Stress. Research Publications: Association for Research in Nervous and Mental Disease. 1949. Vol.29:44. 677 Richter, C.P. Rats, Man, and the Welfare State. p.18. 678 Richter had been less physiologically deterministic in the past, describing the differences in individual albino rats according to the richness of their experience: “An individual brought up in the multiple cage [with apparatus designed to allow for a range of different behaviors] is far more intelligent than one raised in the ordinary running cage. Within the multiple cage, the animal shows all kinds of constructive and imaginative activities, rarely, if ever, seen in an ordinary laboratory rat, and on occasions when it escapes, it avoids recapture with extraordinary success.” This lesson in the value of individual experience seems to have been forgotten in Richter’s later assessment of the abilities of the domesticated albino rat. Richter, C.P. Animal Behavior and Internal Drives. p.341.

323 concluded, “[W]e ultimately may find that a high price has been paid in corticoid hormones for the security provided by a controlled environment.”679

Throughout the 1950s, Richter became less circumspect about the analogy between domestication and civilization; his discussion became less technical, more political, his tone less descriptive and more prescriptive. Embracing the roles of amateur sociologist and historian, he blended epidemiological findings and public health statistics with allusions to the hedonism and decadence of 2nd century Rome and Pharaonic Egypt, tropes familiar to an older generation of degenerationist literature. His central physiological mechanism likewise reiterated longstanding associations between civilization, luxury, and sexual perversion. Richter’s biographer Jay Schulkin (an accomplished physiologist in his own right) described the putative shift from the primacy of the adrenal glands to the gonads in the domestication of the Norway rat as “nice evolutionary physiology hand waving.”680 It betrayed, however, much more. It signaled

Richter’s discomfort with what he claimed were the symptoms of these changes in humans, especially “the increased interest in sex and sex activity… [and] the high birth rates at all levels of the population.”681 Domestication was, for Richter, ultimately a seduction, one that offered security in exchange for dominion. He captured this sentiment in an extraordinary obiter dictum during his presentation at the 1952 meeting of the American Society of Human Genetics: “[George] Orwell’s idea of man’s life in

1984 closely resembles that of our domesticated Norway rat- ‘happily’ living out its

679 Richter, C.P. The Effect of Domestication on the Steroids of Animals and Man in CIBA Foundation Symposium- Steroid Hormone Administration. Vol.3, Book II of Colloquia on Endocrinology. G.E.W. Wolstenholme (ed.). London: Churchill, 1952. p.106. 680 Schulkin, J. Curt Richter: A Life in the Laboratory. Baltimore: Johns Hopkins University Press, 2005. p.94. 681 Richter, C.P. Rats, Man, and the Welfare State. p.23.

324 caged existence.”682 Ironically, Richter’s solution to the incongruity between man’s natural frame and the created environment was yet more control, specifically a new agency to manage the “conservation and preservation of human resources just as now other commissions deal with the conservation and preservation of our soil, forests, and fisheries.”683

Reaction to Richter’s reviews of his domestication research had been mixed from the beginning, though few had failed to note the political and sociological implications of his work. In the discussion following Richter’s 1949 presentation before the Association for Research in Nervous and Mental Disease, Hans Selye announced that he was in almost complete accord with Richter’s conclusions. Neuropathologist and scientific administrator Frank Fremont-Smith, however, asked Richter if the recent actions of

American soldiers in war did not suggest the “probability that under appropriate environmental encouragement, the human race can once again respond adequately” to stress and if their heroics would not compel him to revise his “gloomy and fatalistic outlook.” Richter could not provide a brighter outlook; he merely conceded, “I have expressed a rather gloomy view. I hope many things I have said about the rat are not true for man.”684 Robert Morrison of the Rockefeller Foundation was ambivalent, hopeful for

“the sociological and physiological speculations to be derived,” but concerned that

Richter had become “over enthusiastic in analogizing man with the domesticated rat.”685

682 Richter, C.P. Domestication… and… Genetics in Man. p.283. 683 Richter, C.P. Rats, Man, and the Welfare State. p.26. 684 See discussion in Richter, C.P. Domestication… and… Stress. p.45-46. 685 Morrison, R.S. Memorandum of Interview with Curt P. Richter, January 13, 1948. RG 1.1, Series 200A, Box 95, Folder 1147. RAC.

325

As the severity of the consequences for human self-domestication grew in

Richter’s estimation, so did the breadth and intensity of the commentary directed toward his work. Leonard Carmichael declared Rats, Man, and the Welfare State “a most significant paper” in 1959.686 Conservative columnist Holmes Alexander immediately saw the policy implications of Richter’s findings. He warned that a “food stamp

(giveaway) bill” that had recently passed the House of Representatives might result in the

“untoward happenings” Richter described in his recent paper. To avert such disaster,

Alexander thought that Richter’s findings should command the attention of no less than

President Eisenhower and the aspirants seeking his job. “Dr. Richter’s findings,”

Alexander assured, “would give Candidates Nixon, Rockefeller, Kennedy, Johnson,

Symington, Humphrey and others something new which they could ‘view with alarm.’”687 The bulk of the commentary following Rats, Man and the Welfare State was, however, negative. The raft of criticism that followed Richter’s paper in The American

Psychologist indicated that the political basis of Richter’s interpretations had not escaped notice. “Shades of Malthus,” declared Solomon Kaplan, who also rephrased Richter’s conclusions thus: “Scientists to Supply Get Tough Policy for Crocks Coddled by

Creeping Socialism.” James Crumbaugh argued that rather than revealing any great truth

686 Carmichael to Richter March 27, 1959. Leonard Carmichael Papers B.C212, Series I, Folder “Richter, Curt P,” APS. 687 Alexander, H. On the Political Front. The Reading Eagle. August 27, 1959. Alexander described himself in the article as Richter’s friend; the two had been neighbors in the Bolton Hill district of Baltimore in earlier years. He left the city for the greener pastures of the nearby Capital, but remained friends with Richter. He learned of Richter’s domestication experiments at least by 1958, when he visited Richter’s laboratory and reported in unmistakably political terms the “lessons” to be derived from the rat: “Rat study shows that a protected environment makes its creatures become milder, weaker, easier to handle and less concerned with personal liberty. A certain amount of savagery is needed for a people to keep its freedom. The mutation that a protected environment- a welfare state- performs upon our bodies is a tendency toward rot.” Alexander, as surely as John Dos Passos (see following pages), helped shape the political implications of Richter’s domestication research. See Alexander, H. Lessons from the Rat. Los Angeles Times. June 16, 1958. Part III, p.5.

326 about civilized man, Richter’s domestication analogy “misses the whole point of civilization.” Kaplan agreed: “To draw a parallel between domestication and civilization is to devalue the mastery which man has wrestled from nature and to dim the hope of attaining the greater mastery which more and more will separate civilized man from the domesticated rat, the coddled crock and passive dependency upon scientific commissions whose advice is more honored by emoluments than by observable consequences.”688

These and other forms of public rebuke did little it seems to shake Richter’s confidence in the domestication analogy, his ideological commitments, or his self-image as an objective reporter. They did, however, seem to chasten his pen and polemics.689 He did not offer another review of the domestication work after 1959, save for brief mentions in reminiscences of his career.

In more than 100 papers and presentations on other subjects, Richter had never approached the overtly political tenor of his reviews of the domestication work. In his later recollection of this period, he scarcely acknowledged the candor or the extraordinary reach of his interpretations. The work on domestication, he maintained, had proceeded as his other work had done, each problem arising organically from a previous observation.

His research was unfettered by agendas, spontaneous, like the behaviors he hoped to

688 The commentaries, including Crumbaugh’s and Kaplan’s, appear in The American Psychologist. Vol.14(9):592-595. 689 Leonard Carmichael seems to have come closest to convincing Richter to write on the subject again. In March 1969, he requested another copy of Rats, Man, and the Welfare State (now a decade old) from Richter. In April Carmichael, Richter, and Wilbur Garrett, an editor at National Geographic (where Carmicheal was vice president for research and exploration met for lunch to discuss Richter submitting a new article on rats for the magazine. Garrett reported to Carmichael in December that since the luncheon, he had heard nothing from Richter. Carmichael contacted Richter and learned that he was “still thinking about the rat article.” The article was never published, and may never have been submitted. See Carmichael to Richter March 5, 1969, Garrett to Carmichael December 1, 1969, and Carmichael to Richter December 4, 1969. Leonard Carmichael Papers B.C212, Series I, Folder “Richter, Curt P,” APS.

327 elicit in his laboratory animals. Richter had naturalized his scientific development, beginning with the “release of [his] gene,” the moment John Watson had left him alone with a dozen rats and a fertile imagination. In the midst of his work on domestication,

Richter warned the National Research Council’s Committee on Medicine of the pitfalls of

“design research” and reminded them that:

in the past great discoveries have with few exceptions been made by individuals, often working in great isolation; that some of the most important discoveries have been made without any plan of research… that discoveries have resulted from a general state of what Alan Gregg has called puzzlement… a state of mind which would not lend itself to any accurate verbal description; that there are researchers who do not work on a verbal plane, who cannot put into words what they are doing- whose thinking functions in terms of experiences, subconscious observations- who don’t know what they are after until they actually arrive at their discoveries.690

Richter had tried to cultivate this image of himself: a lone hunter after knowledge armed with an unspoiled naiveté. A quotation from one of his scientific heroes, François

Magendie, adorned the wall of his laboratory: “I compare myself to a scavenger; with my book in my hand and my pack on my back, I go about the domain of science, picking up what I can find.”691 Ironically, this romantic self-image had also driven him to some of his most baldly political assertions. As historian of science Edmund Ramsden has shown, Richter’s characterization of the wild rat reflected his own sense of rugged individualism, forged in the resourcefulness of his youth on the “frontier.”692 We must also consider, however, factors external to Richter, in his social milieu, and in contexts closer to this outburst of social criticism.

690 Richter, C.P. Free Research versus Design Research. Science. 1953. Vol.118:91-92. 691 Quoted from Schulkin, J. Curt Richter. p.116. 692 Ramsden, E. Rats, Stress, and the Built Environment. History of the Human Sciences. 2012. Vol.25(5):130.

328

Henry H. Donaldson and Adolf Meyer, perhaps the two men most responsible for establishing the domesticated albino rat as a standard laboratory animal, were both key influences on Richter’s early career.693 Richter had been introduced to Donaldson, perhaps by Watson, during his doctoral research on the spontaneous activity of the

Norway rat. At the time Donaldson was the foremost expert in the biology of the animal and was engaged in a long-term study of the effects of its domestication in the laboratory, an investigation that became an important precedent for Richter’s much later work.

Richter reported that he met with Donaldson frequently over the years and that the two were close. There is little to suggest, however, that they discussed the effects of domestication; Richter expressed interest in the phenomenon only years after

Donaldson’s death. Donaldson had not been coy about his intentions to apply the lessons learned from wild and domesticated rats to man, but was never so incautious as Richter in outlining the social implications of his studies. It seems unlikely that Richter derived the association between domestication and social degeneration from Donaldson. The specter of social degeneration was especially live in Donaldson’s generation, and he turned a skeptical eye to the notion. He decried the tactics of the “alarmist school” of Nordau and

Lombroso, which, he claimed, found “an historical parallel in the method of making witches once used in Salem.”694

Meyer, whose influence and, indeed, control over Richter’s early career was unparalleled, was equally critical of any generalized concept of degeneration. “Grown up in the Darwinian movement,” he declared as early as 1896, “we can hardly realize the

693 On Donaldson and Meyer’s roles in making the albino Norway rat into a standard research animal, see Logan, C.A. Altered Rationale. p.3-24. 694 Donaldson, H.H. Review of Genius and Degeneration by W. Hirsch. American Journal of Sociology. 1897. Vol.2(6):874.

329 primitive meaning of the general expression degeneration, as used by writers who first introduced it… The originally perfect man has been swept away by the doctrine of evolution.”695 Having matured in the clinical atmosphere of the asylum, Meyer even criticized more evolutionarily palatable iterations of degeneration theory for their vagaries of diagnosis and arbitrary assumptions of normality and deviance. In his psychiatry, he eschewed generalities and stressed the need to view the individual in the totality of his or her own life. It is remarkable that Richter, who drew so much from the philosophy of Meyer and gave the latter’s vision of psychobiology life in the laboratory, should have advanced a sweeping theory of constitutional and social decline more than a half century after his mentor emerged as one of the most vocal critics of such theories.

For Richter the appeal of domestication lay in its broad explanatory powers. It simultaneously altered anatomy, physiology, and behavior and promised to reveal the nexus of their interaction within the evolutionary adaptive framework of Richter’s psychobiology. It allowed him to point to a single root cause for the physiological shortcomings of his albino rats and, by analogy with civilization, for many of the perils facing modern humans. Tellingly, Richter cited his domesticated rats’ “inadequate equipment for meeting stress” as an example of Hans Selye’s “diseases of adaptation.”696

Disease, in Selye’s concept, was a nonspecific condition, a failure of the animal to adapt to its circumstances. Selye explained that he arrived at this idea initially as a medical student in Prague where he mused on the similarities of many disease symptoms and the condition of “just being sick.” Years later, while puzzling over experimental

695 Meyer, A. A Review of the Signs of Degeneration and of Methods of Registration. American Journal of Insanity. 1896. Vol.52(3):344 696 Richter, C.P. Domestication… and… Stress. p.43.

330 results in which rats responded in a stereotyped fashion to a range of different tissue extracts, he thought again of the nonspecific nature of disease and put forth his new interpretation in his 1936 Nature paper, “A Syndrome Produced by Diverse Nocuous

Agents.” As historian of science Russell Viner has argued, Selye’s unified concept of disease was less likely the result of “youthful intuitions and inspirational flashes” than it was the “influence of antireductionist and holistic movements in European and American medicine” during Selye’s intellectual maturation.697 These movements, of course, included the constitutional approach of Charles Stockard, Lewellys Barker, and George

Draper whose definition of disease as “the expression of a conflict between a given external or environmental agent (such as a bacterium, a chemical toxin, physical violence, or psychic stress) and a given human being” came into print a month before

Selye’s “Syndrome” paper.698

Curt Richter had forged his outlook in a similar intellectual atmosphere (he and

Selye even shared time briefly at the Johns Hopkins Hospital campus). The two shared an abiding commitment to an adaptationist philosophy and an admiration for one another’s work. Selye had opened the discussion panel for Richter’s first synthesis of the work on domestication in 1949; he gushed, “I have followed Dr. Richter’s work for many years now and I so heartily agree with all his conclusions that I think I am the wrong man to discuss the paper. I feel that the interpretation of his data which he has given is very well supported.”699 It seems clear that Richter modeled his use of domestication as a

697 Viner, R. Putting Stress in Life: Hans Selye and the Making of Stress Theory. Social Studies of Science. 1999. Vol.29(3):394. 698 Quoted from ibid. 699 Richter, C.P. Domestication… and… Stress. p.45.

331 unifying causal mechanism, at least in part, after Selye’s use of stress and the general adaptation syndrome to explain the broad range of human health and misery. After all,

Selye had successfully, albeit controversially, popularized these concepts in the scientific world and beyond, spawning dozens of “institutes of stress” and even an incipient philosophy and ethics around the stereotyped response to stress he had described. Selye had made “stress” into a social as well as a physiological phenomenon. His ideas were a crucial precedent for Richter’s thinking about domestication and foray into sociology.

The tenor of Richter’s musings on domestication and degeneration, however, depended in part on a more unlikely source, the American novelist John Dos Passos.

Richter had met Dos Passos in June 1934 when both served as groomsmen in the wedding of W. Horsley Gantt.700 Dos Passos was then at the height of his literary fame, having written the first two installments of his opus U.S.A. trilogy with the third in preparation. His was a “fiction of despair,”701 and the degenerations he depicted, both individual and socio-historical, were the results of false experience, of artifact, of being untrue to one’s nature.

Dos Passos had always been troubled by pretenses. He was born a bastard and lived under his mother’s surname until his father, John Randolph Dos Passos, acknowledged paternity when the younger Dos Passos was 16. Despite his reticence,

John Randolph financed a superlative education for his son, steeped in the classics and including the requisite tour of Europe and the Middle East. The itinerant Dos Passos finally landed at Harvard where, perhaps for his own rootlessness, perhaps for the

700 Virginia Spencer Carr Interview with Curt Richter March 21, 1977. Virginia Spencer Carr Collection, John Dos Passos Records, Series 1, Box 2, Folder 115. 701 Colley, I. Dos Passos and the Fiction of Despair. Totowa, New Jersey: Rowman & Littlefield, 1978.

332 affectations and effete posturing of the “Harvard aesthetes” he encountered there, he began his search for “the core,” the fundamental and most essential moral qualities of an individual.

Critic John Diggins explains that even early in Dos Passos’s career “[His] moral orientation [was] clearly retrospective,’ a feature that troubled his friends on the radical, revolutionary left.702 He, thus, located the “core of value” in nature and the distant, romantic past and always looked there first to ground it. Fresh from college, Dos Passos lamented that the problem with American writing was that it lacked an “unconscious intimacy with nature… No ghosts hover about our fields; there are no nymphs in our fountains; there is no tradition of countless generations tilling and tending to give us reverence,” whereas “[i]n other countries literature is the result of long evolution… fervid with primitive savageries, redolent with the old cults of earth and harvest.”703 Modern man was similarly handicapped. Driven by pretenses, the pursuit of money, or his ideals and not the natural expression of emotion and instinct, his life was “from one point of view a constant succession of thwarted desires.”704

This basic tension runs throughout Dos Passos’s early fiction. In Streets of Night the character Wenny seeks to express his Dionysian nature against the backdrop of effete intellectualizing and repressive Puritanism in Cambridge, Massachusetts. He is always in

“search of words” to express the ripe “joy and agony” he feels, and dreams at one point that he found them on a “yellow, half-obliterated parchment” that would “resolve the

702 Diggins, J.P. Visions of Chaos and Visions of Order: John Dos Passos as Historian. American Literature. 1974. Vol.46(3):331. 703 In John Dos Passos: Travel Books and Other Writings 1916-1941. New York: Literary Classics of the United States, 2003. p.588. Originally appeared in the October 14, 1916 issue of The New Republic. 704 Quotation from Clark, M. Dos Passos’s Early Fiction, 1912-1938. Cranbury, New Jersey: Associated University Presses, 1987. p.25. Originally from Dos Passos’s college essay “Art and Baseball.”

333 festering chaos of the world into radiant Elysian order.”705 In Manhattan Transfer and throughout the U.S.A. trilogy the “old words,” the “clean words our fathers spoke,” are meant to confer a similar clarity, though most of the characters in the titular city and country understand only a perverted meaning of this pure and primitive wisdom.706

Dos Passos’s emphasis on the core seems odd at first, as one of the most consistent criticisms of his fiction was that he treated his characters with insufficient depth, that they were merely tumbled and blown about by ambiguous and irrepressible forces beyond their control. Fellow Harvard alumnus Malcolm Cowley tacitly agreed by including Dos Passos in the tradition of naturalism in American fiction, which he described in two words as “pessimistic determinism.” Cowley suggested that the authors of American Naturalism (London, Norris, Dreiser, Steinbeck, among others), took as their archetype Émile Zola, who explained that his first duty as a writer was to “study men as simple elements and note the reactions… [to] subject men and women to things.”707 Indeed, Dos Passos would later describe his style in a letter to Edmund

Wilson as “the behavioristic method,” that is, the “method of generating the insides of the characters by external description.”708 However, as literary critic Michael Clark points out, the inspiration for Dos Passos’s method seems to have been drawn more immediately from the pragmatism of James and Dewey. His description of the “behavioristic method” and his preoccupation with “the core” might well have been lifted, Clark explains, from

705 Quotations and analysis from Clark, M. Introduction to Streets of Night by John Dos Passos. Cranbury, New Jersey: Associated University Presses, Inc., 1990. p.17. 706 See Vanderwerken, D.L. Dos Passos and the “Old Words”. Ph.D. Dissertation, Rice University, 1973. 707 Cowley, M. “Not Men”: A Natural History of American Naturalism. The Kenyon Review. 1947. Vol.9(3):414-415. Emphasis original. 708 Dos Passos to Wilson June 27, 1939 reprinted in The Fourteenth Chronicle: Letters and Diaries of John Dos Passos. Townsend Ludington (ed.). Boston: Gambit, Inc., 1973. p.522.

334 an excerpt in James’s Psychology: “The very core and nucleus of our self, as we know it, the very sanctuary of our life, is the sense of activity which certain inner states possess.

This sense of activity is often held to be a direct revelation of the living substance of the soul.”709 Dos Passos was not content to theorize about the core of his characters, rather he sought to test their moral mettle against the dominant forces of the day and to observe their behavior objectively.

Dos Passos’s interest in the objective description of behavior brought him perhaps closer into the fold of psychology, especially experimental psychology, than any writer of his generation. His envoy to the field was W. Horsley Gantt, whom Dos Passos had met during his Soviet sojourn in 1928 while Gantt was working in the laboratory of Ivan

Pavlov. Adolf Meyer summoned Gantt back to his native Mid-Atlantic the following year to establish a Pavlovian Laboratory at the Phipps Clinic at Hopkins (alongside

Richter’s Psychobiology Laboratory). Dos Passos paid visits to Gantt both at his home in

Baltimore and at the laboratory, especially after Dos Passos settled on his father’s old farm in northeast Virginia with his second wife, Elizabeth, in 1949. While psychiatrists had rebuked Zola and other naturalist authors of the late 19th century for pretending toward science,710 Gantt welcomed Dos Passos into the laboratory. Dos Passos made original descriptions of subject animals there, and Gantt asked him at one point to contribute to a conference on physiological psychology in honor of the 30th anniversary of the Pavlovian Laboratory.

709 Quotation from Clark, M. Dos Passos’s Early Fiction. p.26. 710 See Pick, D. Faces of Degeneration: A European Disorder, c.1848-1918. Cambridge: Cambridge University Press, 1989. p.74-76.

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The result, a short paper entitled “Objective Description,” idealized its subject matter and revealed the extent to which his stark pragmatism was colored by a persistent romanticism. Objective description began, he claimed, with a “virginity of the perceptions,” a subordination of self-consciousness “analogous to the hunter’s” mental state. “[T]o drop into the innocent and naïve state of mind,” offered a sublime connection to the absolute, “a flitting hint of that first fine careless rapture,” a “primeval happiness… You look out at the world with a fresh eye as if it were the morning of the first day of creation.”711 Dos Passos recycled many of these phrases in a strikingly different context, the individualist political essay “A Question of Elbow Room,” drafted a few years later. The detachment that Dos Passos praised in both papers- “a happy self- forgetfulness where there is no gap between observation and description”712- recalled the state of “puzzlement” that Curt Richter had recently extolled, that pure and unprejudiced product of “experiences [and] subconscious observations.”713 By tilting at hardening bureaucratic tendencies, Dos Passos came even closer to the political message of

Richter’s “Free Research versus Design Research”: “Such a view is unlikely to result from the labors of research teams or sponsored surveys. The prime discoveries are more likely to be made by solitary individuals, who have managed by hook or crook to find the elbow room they need to look about them, and the selfsufficiency [sic.] they need to observe the world objectively.” The political dimension of these works suggests that the

711 Dos Passos, J. Objective Description in Physiological Bases of Psychiatry. W. Horsley Gantt (ed.). Springfield, Illinois: Charles C. Thomas Publisher, 1958. p.39-40, 43. 712 Dos Passos, J. A Question of Elbow Room. The National Review. January 25, 1958. 713 Richter, C.P. Free Research versus Design Research. p.92.

336 sympathies between Dos Passos’s and Richter’s views relied on more than a shared, naïve pragmatism.

It may be said that Richter met Dos Passos twice- as an acquaintance at Gantt’s wedding in 1934 and as a close friend after Dos Passos moved to Virginia in 1949- and that each time he met a different political animal. Critic Iain Colley explains, “[A]s a man [Dos Passos] is so elusive as to be a source of embarrassment… but the most generally known fact about [him] is that he switched political sides.” When Dos Passos met Richter in 1934 he had just begun his descent from the “perihelion” of his enthusiasm for the radical left.714 Personal tragedy affected the final break in 1937; Dos

Passos’s longtime friend, Jose Robles, had disappeared while serving Loyalist forces in the Spanish Civil War and was reportedly executed by communists who accused him of spying for Franco’s fascists.715 This and other contributing factors notwithstanding, the span of Dos Passos’s political swing in the coming years- from the New Masses to the

National Review, from William Zebulon Foster to Barry Goldwater- was remarkable and

714 In January of that year, Dos Passos was listed as a primary contributor for the New Masses. February, however, brought tension. Communists had intruded on a socialist rally in Madison Square Garden, ending in a violent melee rather than radical unity. Dos Passos signed a letter with 24 others denouncing the counterproductive actions of the Party and sent it to the editors of the New Masses. The editors wrote their reply to Dos Passos, disregarding the other signers as Trotskyites and fair- weather friends, pleading with him to reaffirm his fealty. Instead their letter reaffirmed Dos Passos’s longstanding distrust of too-strong orthodoxy; he broke ties with the New Masses. See Hicks, G. The Politics of John Dos Passos. The Antioch Review. 1950. Vol.10(1):93. 715 Even before Dos Passos learned of the execution, he was beginning to learn the capriciousness of Party favor as his star declined in the eyes of Soviet officials. His friend Ernest Hemingway was in favor then, and his defense of the executioners was Dos Passos’s final disillusionment with the radical left. For more on the fallout Dos Passos had with Hemingway, see: Pizer, D. The Hemingway- Dos Passos Relationship. Journal of Modern Literature. 1986. Vol.13(1):111-128 and The Breaking Point: Hemingway, Dos Passos, and the Murder of Jose Robles by Stephen Koch. Berkeley, California: Counterpoint Press, 2005.

337 easy enough to caricature.716 In Richter, however, Dos Passos seemed to have found a sympathetic friend, one who had undergone a significant rightward shift himself.

Curt Richter’s time at Harvard overlapped briefly with Dos Passos’s tenure, though he was distant from the affluent and artistic circles in which Dos Passos traveled.

He was, however, privy to the broader intellectual climate in Cambridge and the youthful enthusiasm for revolutionary politics. Richter recalled in an autobiographical essay that during his tenure at Harvard he had “become an active reader of The Nation, The New

Republic, and The Masses”- all leftist journals for which Dos Passos was a contributor during his most radical period. In the next line Richter claimed that these interests fell off in the coming decade, as “’biology’ would take over my life, practically to the exclusion of everything else.”717 Given the political flavor of his domestication work in the 1950s, we may be excused some incredulity. This body of social commentary does, however, stand out among Richter’s other works, a distinct phase beginning in 1949 and ending in 1959.

Richter may well have arrived at the analogy between domestication and civilization without Dos Passos’s help, but the hardening political tone of his reviews and the historical and cultural allusions he used, suggest that Dos Passos played an increasingly large role in shaping the dire implications he derived from it. Timing is significant here. Dos Passos became a more conspicuous presence in and around

716 Hemingway obliged: “He has the irreplaceable early training of the bastard, and a latent and long denied love of money. He ends up rich himself, having moved one dollar’s width to the right with every dollar that he made.” See Hemingway, E. A Moveable Feast. New York: Charles Scribner’s Sons, 1964. p.208. 717 Richter, C.P. …Land of My Genes. p.369.

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Baltimore in 1949, the year leading up to Richter’s first use of the analogy.718 In the coming months, with Gantt as his guide, the normally shy and retiring Dos Passos was adopted into Baltimore’s social life, including the 14th West Hamilton Street Club where

Richter was a member along with Hamilton Owens, Louis Azrael, and H.L. Menken of the Baltimore Sun, Ogden Nash, Milton Eisenhower, and others of Baltimore’s social and intellectual elite. Richter and Dos Passos became fast friends over weekly lunch meetings and monthly dinners at the Club and at the steeplechases, the premier social event of the spring in Baltimore. By 1952 Dos Passos and his family were living in

Baltimore during the week to be closer to friends and the cultural benefits of the city, returning to the farm in Virginia on weekends and during the summer months. Richter’s review of his domestication research in that year followed the format of earlier iterations, but struck a tone distinctly more political than those before it. The changing selection pressures and adaptive constraints he mentioned previously now took specific identities:

718 Dos Passos had begun trying to establish a working farm on his father’s property in northeastern Virginia at least by 1948, but legal disputes with his half brother Louis, travel arrangements, and preparing for his wedding to Elizabeth prevented the Dos Passoses from firmly establishing themselves in the Northern Neck of Virginia until late in 1949. See Carr, V.S. Dos Passos: A Life. Evanston, Illinois: Northwestern university Press, 2004. p.385-386. Dos Passos’s effort at farming, though ill fated, was philosophically revealing. His pragmatism had always made him sympathetic to the tangible and productive merits of unskilled labor, which at an early age he naively surmised, “must make one lay hold of the elementary facts of existence in a wonderful way” (Quotation from Clark, M. Dos Passos’s Early Fiction. p.27). His sympathies shifted along with his politics in the decade of the 1940s, from the industrial worker (most likely organized) to Jefferson’s yeoman farmer (a truer emblem of independence he judged). In that decade Dos Passos, most likely through the influence of fellow writer-turned-farmer Louis Bromfield, became involved with the Friends of the Land, a multipartisan social and political organization that promoted ecologically mindful modern farming techniques as well as a Romantic view of the spiritual and community benefits of rural life. Dos Passos had long been influenced by the Jeffersonian ideal of self-governance and individual freedom, though it would be fair to say that this influence was at its greatest in the late 1940s and early 1950s, as he began experimenting with farming and prepared for his biography The Head and Heart of Thomas Jefferson. Dos Passos contributed to the Friends of the Land’s quarterly magazine The Land and to the organization’s anthology Forever the Land: A Chronicle and Anthology. Russell and Kate Lord (eds.). New York: Harper and Brothers, 1950. It is worth noting that Curt Richter presented a paper on diet selection in wild and domesticated rats at the group’s 1946 Conference on Conservation, Nutrition, Health, and Happiness in Athens, Ohio.

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“family welfare, social security, unemployment insurance, old age pensions, police protection”- all devices by which modern man was “protected almost from the cradle to the grave.” More telling, he equated social security with the “control of man” and finally with the “caged existence” of the domesticated Norway rat.719 The following year he decried the bureaucratization of research as science became increasingly reliant on government subsidies.720 Richter’s social and political concerns at midcentury were significantly Dos Passos’s too- the corruption of pastoral innocence by modern life and the myopic exchange of freedom for security- which found their fullest expression in

1959’s Rats, Man, and the Welfare State. In the same year Dos Passos offered a defense of the fundamental continuity of his art and politics, which might well have served as the

719 Richter, C.P. Domestication… and… Genetics in Man. p.283. Richter’s 1952 paper also included allusions to the decline of some of the great civilization of Antiquity, and he admitted his recent interest in ancient civilizations to Anne Roe in his interview with her that year. It is unclear if this was in any part Dos Passos’s influence, though Dos Passos had retained a longstanding interest in the lessons of ancient cultures since his father introduced him to Gibbon’s Decline and Fall sometime before his 10th birthday. In the interview with Roe, Richter also mentioned reading The Mother Country, a defense of capitalism by Sidney Smith, former secretary of Britain’s National Anti-Corn Law League, written in the immediate wake of the Communist Manifesto. Smith’s proposal, to depauperize Britain by leasing small plots of “waste” land to the country’s poor for them to cultivate, resonated with Jefferson’s vision of the yeoman farmer and with Dos Passos and Richter’s shared ideals of self-government and individual freedom. Indeed, Smith’s words, though written in 1849, might have served as an epigraph at the heading of one of Richter’s reviews or the beginning of one of Dos Passos’ chapters one hundred years later: “The wise impulses of our instinctive nature have been overlaid and smothered by over-government. We have substituted artificial state-craft for the wise economy of providence. We have over-civilized ourselves into a state of property run mad. Indeed, the extent of the interference we advocate with the existing arrangements of our social polity, may be reduced simply to a demand for a recurrence to the natural order of things, and the suggestions of common sense.” Smith, S. The Mother Country: or the Spade, The Wastes, and the Eldest Son, an Examination of the Condition of England. London: William S. Orr & Company, 1849. See also Roe Interview of Richter March 1952. p.7. 720 Richter, C.P. Free Research versus Design Research. p.91-93. He expressed a similar opinion, privately, in an interview with Anne Roe nearly a decade later when he complained that government subsidies for research most often meant “pouring great sums of money out to supporting people who really don’t want to work, who do this because it’s thrown in their laps and so that it clutters up many of the labs… [S]ome people think that’s wonderful- having a lot of workers around- so-called workers- and I think that’s just a snare and a delusion. It really slows things up terribly.” from Anne Roe interview of Curt Richter October 1962. Anne Roe Papers B.R621, Folder Richter, Curt 1, p.5-6. APS.

340 summary statement of Richter’s evolving concern over human self-domestication: “the basic tragedy my work tries to express remains monotonously the same, man’s struggle for life against the strangling institutions he himself creates.”721

Domestication and the Psychobiological Critique of Behaviorism at Midcentury

Curt Richter remained a champion of the Norway rat, wild and domesticated, claiming in 1968 that if granted “ the power to create an animal most useful for all types of studies on problems concerned directly or indirectly with human welfare, I could not possibly improve on the Norway rat.” Given Richter’s sociological concerns, he came to value the domesticated variety not despite its degeneracy but precisely because of it.

However, as Edmund Ramsden has argued, the image Richter created of the albino rat- defective, adulterated, improvidentially synthetic- raised (or rather reinforced) doubts about the use of a creature so contrived for clarifying natural behavioral mechanisms.

This had been the problem of domestication for psychobiologists since Willard Small asked what happened to the rat’s “psychic nexus” since its removal from the “loom of necessity.” But Small’s viewpoint had lost, and domestication hardly figured into the triumphant behaviorism except to make the rat more amenable to the tightly controlled conditions of the laboratory. Richter’s construction of the domesticate-as-degenerate may have failed to rouse a slumbering public to the dangers of self-domestication, but it resonated with a widening critique of the conventions of American comparative psychology by that minority of researchers still concerned with the adaptive value of

721 Dos Passos, J. “Looking Back on U.S.A.” New York Times. October 25, 1959. p.5.

341 behavior, not as a conditioned response in the individual, but as reactions fixed in the phyletic history of the species. This critique, which coincided with a growing appreciation of European ethology, seized upon the image of the albino Norway rat as an artificial denizen of the laboratory- degenerate, incomplete, and thus unrepresentative of animal behavior generally.

There had always been willing critics of behaviorism’s dominance of comparative psychology, but the flourishing of such criticism after midcentury had its seed in Frank

Beach’s landmark 1950 review, “The Snark was a Boojum.” In the Lewis Carroll poem from whence Beach derived his unorthodox title, a party hunting “Snarks” is given a haunting admonition: “If your Snark be a Snark, that is right: Fetch it home by all means- you may serve it with greens… But oh… beware of the day, if your Snark be a Boojum!

For then you will softly and suddenly vanish away, and never be met with again!” Beach explained the allusion: “Thirty years ago in this country [the Comparative Psychologist] went Snark hunting… Instead of animals in the general sense he found one animal, the albino rat, and thereupon the Comparative Psychologist softly and suddenly vanished away.”722 He analyzed the articles published in the pioneering Journal of Animal

Behavior from 1911 to 1946 and noticed that as the number of articles grew, there was a corresponding decrease in the diversity of species represented. After 1920, mammals dominated, and from 1930 fully more than half of the papers published dealt exclusively with the albino Norway rat. Such a psychology, whatever its merits, could hardly be called comparative. “You will recall,” begged Beach, “that the Pied Piper rid Hamelin

Town of a plague of rats by luring them into the river with the music of his magic flute.

722 Beach, F.A. The Snark was a Boojum. The American Psychologist. 1950. Vol.5(4):115.

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Now the tables have turned. The rat plays the flute and a large group of human beings follow.”723 He also found that the paucity of species used was correlated with a lack of diversity in the kinds of behavior studied in the major journals of comparative psychology. More than half of all papers dealt with conditioning and learning. Perhaps, he jabbed, a new journal title would better suit the habits of the field: “The Journal of Rat

Learning.”724

Beach’s review did not dwell on the issue of domestication nor did it disparage the albino rat as a degenerate, but it established the format for the major critique of behaviorism around midcentury and spurred the rise of an alternative and previously marginalized approach to comparative psychology in the United States. The association of the rat and maze was familiar to everyone in American comparative psychology, but after Beach’s paper this emblematic pairing seemed to illustrate the poverty rather than the strength of the methods of the field. He challenged the fundamental behaviorist assumption sanctioning the ubiquity of the albino rat, namely, that this single species was somehow representative of animal behavior in general and human behavior in particular.

For the limitations of their methods and subjects, Beach argued that the behaviorists failed even in their stated goal to give a comprehensive picture of animal learning.

Tellingly, he lamented that the dominance of such a narrow view had caused psychology

“to neglect many complex patterns of response that stand in urgent need of systematic analysis. The best example of this tendency is seen in the current attitude toward so-

723 Ibid. p.117. 724 Ibid. p.119.

343 called ‘instinctive’ behavior.”725 He provided an example of the dangers of this tradeoff, citing rat studies on hoarding, which gave conflicting results on the rat’s general ability to learn to hoard food or other items. Beach explained, “Now this is not surprising since

Norway rats rarely hoard food under natural conditions. Would it not seem reasonable to begin the work with an animal that is a natural hoarder?”726 Beach was advocating

Small’s argument from 50 years prior: that the experiment should be made to fit the animal and its natural proclivities, not the other way around. The price paid for comparative psychology’s narrowness and the path forward were equally clear:

European students in this field have justly condemned Americans for the failure to study behavior in a sufficiently large number of representative species. And non-psychologists in this country are so well aware of our failures to develop the field that they think of animal behavior as a province of general zoology rather than psychology… [T]o maintain our status as indispensible contributors to the science of behavior, we will have to broaden our attack upon the basic problems of the discipline. This will sometimes mean sacrificing some of the niceties of laboratory research to deal with human beings under less artificial conditions. It may also mean expanding the number of non-human species studied and the variety of behavior patterns investigated.727

Although Beach’s paper was widely read, its recommendations were not so readily heeded. A decade after its publication, William F. Dukes and Richard E. Whalen independently updated Beach’s analysis of subject choice in the Journal of Comparative and Physiological Psychology and determined that save for a greater interest in primates, the albino rat remained disproportionately represented in its publications. Whalen also determined that the predominance of learning and conditioning studies had hardly

725 Ibid. p.121. 726 Ibid. p.123. 727 Ibid. p.123-124.

344 lessened over ten years; he concluded, “[T]he unfortunate possibility exists that animal psychology will remain the science of rat learning.”728 Around the same time, Morton E.

Bitterman explained that the behaviorists’ assumption of generality in the learning process had not only sanctioned the concentration on the albino rat, but had bankrupted the field in other ways. This fundamental assumption, he argued, encouraged the sense that the purpose of experimentation was merely to reinforce convictions already held.

Only by adopting a truly comparative approach, might the field “come again to a kind of research that is at once more satisfying and more productive. Its function… inquiry, not proof.”729

Despite these pessimistic analyses, others perceived a growing sense that the field was becoming enlightened according to Beach’s view. Keller and Marion Breland for instance, cited “a continuing realization by psychologists that perhaps the white rat cannot reveal everything there is to know about behavior” at the opening of their coyly titled review, The Misbehavior of Organisms.730 Significantly, the subject of their review was not the white rat at all, but the theoretical basis of behaviorism, for which the white rat served as a convenient totem. The Brelands held an interesting vantage point upon the debate over methods in American comparative psychology at midcentury. The pair had begun their graduate work under B.F. Skinner’s direction at the University of Minnesota and followed him into the war effort in 1943, conditioning pigeons to guide bombs by pecking at crosshairs on a glass screen. They remained keepers of the behavioristic faith

728 Whalen, R.E. Comparative Psychology. The American Psychologist. 1961. Vol.16(2):84. See also Dukes, W.F. The Snark Revisited. The American Psychologist. 1960. Vol.15(2):157. 729 Bitterman, M.E. Toward a Comparative Psychology of Learning. The American Psychologist. 1960. Vol.15(11):711. 730 The title referenced B.F. Skinner’s The Behavior of Organisms (1938). Breland, K. and Breland, M. The Misbehavior of Organisms. The American Psychologist. 1961. Vol.16(11):681.

345 for a number of years, transforming their wartime experience with pigeons into a new field of applied animal psychology, even initiating a commercial business furnishing zoos, museums, movie studios, department stores, fairs, conventions, and various other exhibits with animals trained according to Skinner’s operant conditioning techniques.

However, it was in moving “further and further from the security of the Skinner box,” that by 1961 the Brelands noticed a “persistent pattern of discomforting failures… of conditioned operant behavior.”731

Many of their animals began exhibiting behaviors different from those for which they had been successfully conditioned. The pattern of this drift, the Brelands concluded, was consistent and unmistakably in the direction of “instinctive behaviors having to do with the natural food getting behaviors of the particular species.”732 The Brelands expressed frustration, not so much with the prepotency of instinctive behaviors, but with their initial inability to explain the phenomenon given their background in behaviorism.733 This they regarded as a “clear and utter failure of conditioning theory,” and although they identified their new perspective with European ethology, it stood also as the central tenet of the tradition of psychobiology begun in their native country more than 60 years prior: “[T]he behavior of any species cannot be adequately understood,

731 Ibid. 732 Ibid. p.683. 733 Ibid. Marian Breland (Bailey after her second marriage) later suggested that William Verplank’s 1955 review of ethology, a key introduction for many American psychologists to the ideas of Lorenz and Tinbergen, had given her and her husband a suitable alternative framework to ease their apostasy. See Bailey, R.E. and Bailey, M.B. A View from Outside the Skinner Box. The American Psychologist. 1980. Vol.35(10):942. For Verplank’s article, see Verplank, W.S. Since Learned Behavior is Innate and Vice Versa, What Now? Psychological Review. 1955. Vol.62:139-144.

346 predicted, or controlled without knowledge of its instinctive patterns, evolutionary history, and ecological niche.”734

The Brelands’ statement concerning psychology’s misplaced faith in the white rat seemed prescient in the years following its publication. A growing number of comparative psychologists in these years heaped blame for their field’s shortcomings on the albino rat’s diminutive and, crucially, domesticated frame. In 1964 J. Lee Kavanau reflected on the impact of those measures of control- confinement and compulsory experimental regimes- that had become indispensible to comparative psychology’s claims to objectivity. By “depriving animals of natural outlets for activity,” Kavanau explained, these artificial conditions dramatically altered their behavior. In his own experiments he found that wild deer mice would repeatedly exercise whatever means of control were available to them toward the opposite extreme of the restrictive conditions imposed.

Apparently unconcerned by the traditional bugaboos of animal behavior, anthropomorphizing and subjectivity, Kavanau interpreted his results thus, “These animals find it rewarding to attain and to exercise a high degree of control over their environment, perhaps in partial substitution for the freedom of action enjoyed in the wild, but denied by confinement.” His criticism was not limited to the artificiality of experimental conditions, but extended to the animals that had long been exposed to the

“insults” of the laboratory. Domesticated rats and mice, he asserted, “are hundreds of generations removed from the wild. Their bland behavior tells us mainly how animals react to experimental regimes after many of the characteristic adaptive responses of the species have been largely or completely lost. Domestic animals remain convenient

734 Ibid. p.683, 684.

347 vegetalized strains for physiological studies, but only wild animals provide the full range and vigor of responses upon which solutions to the central problems of behavior must be based.”735 The will Kavanau had located in wild animals was not to be found in their domesticated counterparts. An animal so deprived, it would seem, could not help becoming depraved.

The image of the degenerate albino rat gained wider currency as the decade wore on. Roy Robinson’s standard The Genetics of the Norway Rat, peddled the view that the provisions of the laboratory would, and frequently did allow for “even an ‘idiot’ rat [to] flourish in such an environment”736 Robert B. Lockard explained that the albino rat, albinus as he preferred it, had been aberrant even before it had entered the laboratory.

Citing recent evidence that wild rats trapped by customary methods were usually neither random nor representative samples of the wild population, Lockard speculated that laboratory stocks of albino rats must have been derived from such guileless captives.737

735 Kavanau, J.L. Behavior: Confinement, Adaptation, and Compulsory Regimes in Laboratory Studies. Science. 1964. Vol.143:490. Kavnau claimed, in a follow-up letter to the editor of Science that the response to his report had been enormous and universally positive. He quoted from these in his later memoirs, including this note from an unnamed zoologist: “Some of the points made by you seem fairly obvious to anyone who knows anything about animals… I am glad to see white rats and mice reduced to just what they are- spiritless, vacuous vestiges of their original wild forebears- with all their real vigor leached out by countless generations of domestication and inbreeding. Experiments with these damn things are so far removed from the real world that a new discipline is virtually involved- the laboratory biology and responses of white rats and mice.” Quoted from Kavanau, J.L. J. Lee Kavanau: A Man for All Sciences, Some Arts, and Technology. 2008. Web Manuscript: https://www.eeb.ucla.edu/kavanau/preface.php. Accessed April 6, 2013. 736 Robinson, R. The Genetics of the Norway Rat. Oxford: Pergamon Press, 1965. p.514. Robinson was an amateur geneticist, rather than a psychologist, but his book was a standard reference for psychologists at least as much as it was for geneticists. 737 Lockard, R.B. The Albino Rat: A Defensible Choice or a Bad Habit? The American Psychologist. 1968. Vol.23(10):735. Robert Boice, whose study Lockard cited, drew a very different lesson from his own research. In later years, Boice suggested that the unrepresentative sampling of wild gray rats had skewed the picture of the wild rat as it was understood by psychologists claiming dramatic differences in behavior and physiology between wild and domesticated forms of the species. See Boice, R. Some Behavioral Tests of Domestication in Norway Rats. Behaviour. 1972. Vol.42(3-4):198- 231, especially 200-204.

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Once in the protective environment of the laboratory, “with selection pressure no longer keeping the functions [of albinus] intact, its parts [disintegrated] in whatever sequence the supporting biology [failed].” From a long list of charges against albinus, some supported by previous research, others which he deduced, Lockard concluded, “It is at least a waste of time, if not outright folly, to experiment on the degenerate remains of what is available intact in other animals.”738

The divide between natural and artificial loomed large in Lockard’s indictment of the albino rat, as it had in Kavanau’s. The albino rat’s domestication, or perhaps more accurately, “laboratorization” was “absolutely unique in natural history,” and made the albino rat into “one of the more domestic animals, least related to its ancestors.”739 So mercurial was Lockard’s albinus, so forcefully adapted to sterile cages, fluorescent lights, and the whims of the researcher, that it hardly merited the title ‘animal.’ Rather, it was a

“commercial product,” and he glimpsed in its future development a repellant scene of futile detachment:

One can imagine a sort of ‘schmoo’ in a germ free environment, surrounded by life-support systems and fitted with prosthetic devices, attended by technicians and still studied by scientists; its riddles of ‘the learning process’ still unsolved for continued survival in the new habitat favors a baffling learning process, one which is composed of pieces no longer harmoniously interlocking and which changes in time rapidly enough to make past findings discrepant, all the while producing enough variability to keep the entire enterprise confused.740

Domestication was not a neutral term for Lockard; it hardly could be. He did not have to seek its normative dimensions or the attendant dichotomy between nature and

738 Ibid. p.740, 739. 739 Ibid. p.737. 740 Ibid. p.740.

349 artifice in 19th century accounts by naturalists and anthropologists. They were on display in several of the sources (including recent works by Richter, Lorenz, and Spurway) he cited in his case against the albino rat, and not coincidentally. Lockard shared with

Richter and Lorenz a belief in the adaptive value and evolutionary basis of behavior, if not their respective sociological and political concerns. Rather, Lockard adapted the construct of the domesticate-as-degenerate to the critique of American comparative psychology begun by Beach in 1950.

His ambition (and Kavanau’s as well) had been wider than denigrating the albino rat as an experimental subject; it was in challenging the fundamental behaviorist assumptions that permitted the rat’s ubiquity in the field. Kavanau signaled as much in a follow-up to his original report. Emboldened by the positive responses to his report, he felt confident that “[a] new era in the rationale and design of psychologically oriented animal experimentation may indeed be in its inception.”741 Three years after his condemnation of albinus, Lockard also turned to a broader critique of the field.

Comparative psychology was dead, he declared, and its replacement had emerged in departments of zoology. Comparative psychology (i.e. behaviorism) had taken a fatal draught of narrow scope and dogmatic environmentalism, and even as it tried to incorporate physiological explanations, this was but “a fragment of biology… lacking the dynamic processes of biology: adaptation, natural selection, ecological significance, and evolution.”742 Meanwhile the alternative, or “modern,” approach, which Lockard

741 Hawkins, J.D. and Kavanau, J.L. Wild and Domestic Animals as Subjects in Behavior Experiments. Science. Vol145(3639):1462. 742 Lockard, R.B. Reflections on the Fall of Comparative Psychology: Is there a Message for Us All? The American Psychologist. 1971. Vol.26:169.

350 identified with Darwin, Whitman, and the European ethologists, emphasized precisely these dynamic processes.

Behaviorism’s air of objectivity and rigor was significantly bound up with the image of the albino rat as a scientifically standardized animal. The criticism from psychobiology, however, recast the albino rat as a domesticated rather than a standardized animal and, thus, undermined many of the premises at the heart of the behaviorist program. By pushing for greater control and generality, psychobiologists claimed, the behaviorists had inadvertently created an animal that was less stable, more mutable, and unrepresentative of anything in nature. According to Kavanau and

Lockard’s ecological perspective, if the conditions of life in the laboratory were artificial, the animals raised therein and their reactions could not be otherwise. The albino rat, as a domesticated animal, thus embodied the growing critique that behaviorism produced artifacts, not facts. This is not to suggest that the critique of the albino rat was disingenuous or entirely allegorical. Kavanau, Lockard, and others raised legitimate questions about the appropriateness of the animal for investigating various problems in psychology. However, certain of their claims743 and, more pointedly, their rhetoric betrayed a prejudice in their thinking about domestication and domesticated creatures, one that was predicated on the culturally constructed and scientifically sanctioned image of the domesticate-as-degenerate.

Lockard was roundly criticized for prematurely calling the death of comparative psychology. A fledgling professor at the University of Missouri, Columbia, however,

743 For instance, when Kavanau was asked to justify his claim that the learning capacity of wild deer mice “far exceeded” that of domestic rodents, he could only point to “the known performances of domestic animals,” not to his own or any other specific experimental results. See Hawkins, J.D. and Kavanau, J.L. Wild and Domestic Animals. p.1462.

351 painted a picture of the field even grimmer than Lockard’s: Robert Boice suggested that if comparative psychology had not already fallen, “it may be that [it] never had the elevation from which to fall.” Boice declared that for all the “cathartic” self-criticism of the previous two decades, any comparative psychologist would still have to “admit to the overwhelming influence of Watsonian environmentalism around us.” He lamented that a truly comparative psychology was likely to remain “peripheral in the American scene for a long time,” as zoologists viewed comparative psychologists with suspicion and

“psychology in general [was] more interested in the mentalistic than natural adaptations of the little zoo we keep in their departments.”744 Even as Boice tacitly endorsed

Lockard’s call for a “biology of behavior,” he began to question one of its central justifications, the condemnation of the domesticated Norway rat as an experimental subject.

By the early 1970s Boice had been studying the effects of domestication in the

Norway rat for several years. It had been the central organizing concept behind his 1966 dissertation at Michigan State University and the subject of several research papers since his graduation. In the course of his investigations, he uncovered what he believed to be the two assumptions that stymied psychological research on the question of domestication. First, most researchers assumed the behavioral differences between wild and domesticated animals were already well established. He determined that this impression was founded on culturally familiar notions of degeneracy rather than experimental demonstration. In agreement with Kavanau, he identified the second assumption as a belief that wild rodents were completely intractable and dangerous to

744 Boice, R. On the Fall of Comparative Psychology. The American Psychologist. 1971. Vol.26:858- 859.

352 work with. In 1971 he outlined methods for “laboratizing” the wild gray rat, not, as

Kavanau had suggested, to replace the albino rat, but rather to prepare for comparative studies in order to experimentally evaluate the differences between the wild and domesticated varieties.745

Boice’s effort to uncover objective facts about the effects of domestication was not unprecedented, but the reflexivity he applied to the understanding of domestication in

American comparative psychology was novel.746 In an autobiographical account he claimed his initial interest in the subject was inspired by questioning the domestication- degeneration thesis, rather than an intrinsic interest in domestication. Concerned that he might never formulate an interesting dissertation topic, Boice overheard a fellow student asserting the association of domestication and degeneracy with such zeal that he felt compelled to investigate the relationship himself. His preliminary research revealed little to substantiate the dire consequences of domestication. Colleagues and senior researchers discouraged him from pursuing the question, convinced that maintaining a colony of wild gray rats would be impossible.747 His own experience trapping and keeping gray rats convinced him otherwise, and he soon began questioning the source of the assumptions surrounding the use of wild and domesticated rats. The normative meanings of domestication were never far from Boice’s mind, as evidenced by the title of

745 Boice, R. Laboratizing the Wild Rat (Rattus norvegicus). Behavior Research Methods and Instrumentation. 1971. Vol.3(4):177. 746 Boice undoubtedly benefitted, though, from the reflexive look that Helen Spurway had applied to the discussion of domestication in European ethology in the 1950s. 747 Boice, R. Domestication and Degeneracy in Comparative Psychology: An Evolutionary Analysis of Animal Behavior. M. Ray Denny (ed.). New York: John Wiley & Sons, 1980. p.87-88.

353 his paper at the 1971 meeting of the Midwestern Psychological Association,

“Domestication: Degeneracy, Fitness, or Unnaturalness.”748

Boice’s preliminary work “laboratizing” the wild gray rat produced a chance discovery that called into question some of the previous notions regarding the temperament of the wild rat and its physiological and behavioral distinction from the domesticated albino. In his ignorance about trapping rats, he set out with a single trap for the local landfill- “a fortuitous choice,” he later reported. Not only did the landfill provide rats in numbers, despite Boice’s inexperience and meager equipment, but the rats he acquired were much more tractable and disease free than he had anticipated. As he tried trapping elsewhere, near apartment buildings, packing plants, and feed lots, he found the rats there more difficult to capture and “as impractical to cage or breed as the literature suggested.”749 To account for this discrepancy, Boice hypothesized that the differential stability of the habitats from which the rats were drawn had produced very different samples of the respective rat populations in these areas.

Boice drew on John B. Calhoun and D.D. Thiessen’s work on the sociology of rats and mice, respectively, to suggest that rats sampled from stable habitats (alleys, apartment complexes, and industrial buildings) were more likely to be on the social periphery of the rat colony.750 Frequently bullied by more dominant rats and denied access to surer sources of food, these rats were more likely to seek bait left in traps. They

748 See Hill, W.F. Proceedings of the Forty-third Annual Meeting of the Midwestern Psychological Society. The American Psychologist. 1971. Vol.26(9):782. The paper was presented at a symposium Boice co-chaired on the vexed question of the implications of domestication for man. Other participants in the symposium included J. P. Scott, who had previously denied that domestication in the dog led to degeneracy, and Edward O. Price, who stumbled upon the problem of domestication at about the same time as Boice and likewise criticized the prejudices that attended its study. 749 Boice, R. Domestication and Degeneracy. p.88. 750 Boice, R. Some Behavioral Tests of Domestication in Norway Rats. p.200.

354 also bore the marks of their social status, visible as scars on their tails and backs. Boice determined that these scar-marked rats were frequently diseased, emotional, and uncooperative in the laboratory. Rats sampled from unstable habitats (regularly bulldozed landfills) were much less frequently scar-marked. When scarred and non- scarred rats were compared in the laboratory, the scarred rats were generally more emotionally reactive: they were submissive to unscarred rats in encounters, bred poorly with small litters, and offered poor maternal care. Unscarred wild rats bred comparably to domesticated albinos and were no more difficult to handle.751 Boice repeated Richter’s

1954 experiment, in which wild rats drank excessive amounts of water, ostensibly because of their large adrenal glands. He found, however, that only the scarred wild rats maintained excessive water intake. In a short time, unscarred rats took in water at the same rate as domesticated albinos.752 Boice surmised that the wild rats used in Richter’s study and in other studies claiming a wide disparity in the emotionality of wild and domesticated rats had been scarred rats taken from the periphery of colonies in stable habitats. He concluded further that the general impression given in the literature of the wild rat as highly emotional, vicious, and intractable- and thus wholly distinct from the domestic albino- was due to unrepresentative sampling.

Boice’s program of research was less about evaluating the effects of domestication per se than about evaluating the perception of domesticated animals in contemporary comparative psychology. He focused upon the putative association of domestication and degeneration, which he recognized as a cultural construct. Whether

751 Ibid. p.203-204. 752 Boice, R. Excessive Water Intake in Captive Norway Rats with Scar-Markings. Physiology & Behavior. 1971. Vol.7:724-725.

355 viewed positively, as freedom from “innate dispositions,” or more often negatively as the breakdown of harmoniously adaptive behavior patterns, Boice argued that “[n]either view has much basis in fact.”753 His own experiments on the social behavior of wild, domesticated, and hybrid rats upheld that the varieties displayed important differences in sociality, but denied that aggressiveness and dominance hierarchies had been obliterated in the domesticated rat. Wild rats were less playful in agonistic behaviors, kept a greater distance from newly introduced rats, and showed a wider range of elicited social signals than domesticated rats. However, domesticated rats displayed a greater richness of social behavior when placed into cages with wild rats than they had when housed together, suggesting that the social behaviors were not lost in the domesticated variety, but had a higher threshold of elicitation.

Boice’s main concern, however, was the purported difference in learning ability between the wild and domestic rat. “History,” he wrote, “reveals a generous attitude regarding the learning abilities of the wild rat.”754 He referred to the maze-running studies of Small at the turn of the century and of Calvin Stone in the early 1930s. While

Small denied support for the already familiar association of domestication and degeneration, Boice seized on his explanation that the poorer performance by the wild rats in reaching the center of the maze may be explained away by their greater fear and timidity rather than differences in learning ability. Similarly, he criticized Stone for suggesting that wild rats “may actually have a greater native learning ability” than

753 Boice, R. Behavioral Comparability of Wild and Domestic Rats. Behavior Genetics. 1981. Vol.11(5):546. 754 Boice, R. Some Behavioral Tests of Domestication in Norway Rats. p.213.

356 domestic rats when his experiments did not show any such advantage.755 More recent comparisons between wild and domestic rats were even less useful Boice claimed, pointing to a study by Robert Powell and Grant Morris that compared domesticated

Norway rats (Rattus norvegicus) with wild cotton rats (Sigmodon hispidus), belonging to an entirely different genus. Boice’s own comparisons, between wild and domestic

Norway rats, showed superior conditioning in the domesticated rather than the wild variety in simple tests measuring licking in anticipation of a water drop and in avoidance tests using electric shock. His critics suggested that wild rats had been too uncomfortable under the experimental conditions to demonstrate their superior learning and recommended comparing both varieties in a maze, which more closely approximated the natural capacities of the rat. After selecting unscarred rats and modifying the design of

Stone’s maze to eliminate distracting noises, Boice found that even though the wild rats completed the maze with apparent ease, the domestic rats performed better still. He suggested this was because the domesticated rat was busy exploring during lulls in maze- running, while wild rats usually rested or practiced grooming behaviors. Even when wild rats were gentled to the point of docility or pre-trained in other learning tasks, they failed to match the performances of their domestic counterparts.

Boice then decided to investigate the role of behavioral flexibility in learning by forcing wild and domesticated rats to adjust their drinking habits to a diurnal rather than

755 Stone, though, was much more tentative in this suggestion than Boice suggests. He had mentioned it as a hypothetical factor, among several others, and concluded, “Which… of these suppositions is responsible for the differences obtained [between different coat color strains of rats] cannot be determined from the information at hand.” Boice’s overreach in his analysis of Small and Stone’s interpretations suggests that the depiction of domesticated animals as intellectually degenerate had reached later authors such as Kavanau, Robinson, and Lockard by means other than these particular precedents, perhaps by longstanding cultural notions concerning the nobility of wildness and the poverty of domestication. Quotation from Stone, C. P. Wildness and Savageness in Rats. p.51.

357 the usual nocturnal schedule. In this test, the domesticated albinos proved much more capable of adjusting their drinking habits than the wild gray rats. Boice indicated that if this greater flexibility held for behaviors generally, “[t]his could mean that learning… is more natural for the domestic rat. Then the study of laboratory learning might best be modeled on the domestic rat, an organism as artificially flexible as man.”756 Boice’s pointed phrasing evoked many of the fundamental tensions surrounding research on domestication in comparative psychology in order to suggest how domestication had produced adaptive, not degenerative, changes in the albino rat and made the animal a better, not a poorer candidate for answering some problems in the field.

In a final bid to silence his critics, Boice placed the burden of change upon the domesticated rat by attempting to “feralize” them in an 84 m2 outdoor pen. His goal was to determine if burrowing behavior, ubiquitous in the wild and useless in the laboratory, had vanished from the domesticated rat’s behavioral repertoire after countless generations in captivity. The experiment was reminiscent of Henry H. Donaldson’s failed attempts to feralize the albino rat early in the century, but was concerned primarily with the persistence of ancestral behavior patterns rather than changes in the anatomy of the animal. Boice found that the albino rats began constructing burrows and running paths immediately upon release into the outdoor pen. Their burrows were qualitatively indistinguishable from those constructed by wild gray rats and carried them through two

Missouri winters with little mortality.757 Not only this, but the albinos “[displayed] all

756 Boice, R. Some Behavioral Tests of Domestication in Norway Rats. p.226. Italics added. 757 Boice, R. Burrows of Wild and Albino Rats: Effects of Domestication, Outdoor Raising, Age, Experience, and Maternal State. Journal of Comparative and Physiological Psychology. 1977. Vol.91(3):652. Boice was not the first to observe apparently feralized laboratory rats in a wild or semi-natural setting. Donaldson’s associates achieved at least one generation of feral albino rats in

358 the dynamic social postures (e.g. flank presentation) often thought to be absent in laboratory rats” and by the second generation bred outside, they were “as neophobic and intractable as are many wild rats.”758

Although Boice’s feralization studies indicated that many of the changes associated with captivity were reversible, he emphasized that they “[did] not demonstrate that genetic differences between wild and domestic rats are insubstantial…

Domestication, however complex the interplay of early experience and genetics, is not undone in a single generation.”759 It was a legitimate biological and psychological phenomenon, a kind of adaptive fitness, and a problem worthy of study rather than speculation. Boice’s research had not been directed toward resurrecting the albino rat as an experimental subject, but was rather concerned with separating the animal from the myth surrounding its captivity and domestication. For psychology, he explained, this myth was instantiated by the differing methodological aims of behaviorists and psychobiologists: “Rarely is captivity so distorting as claimed by its critics

[psychobiologists] or facilitating as claimed by its proponents [behaviorists].” He advocated a “compromise stance” by combining typically behaviorist concerns such as

the Dry Tortugas, but the conditions there were deemed too inhospitable to maintain a colony in perpetuity. Nevertheless, at the end of his feralization experiments, Donaldson remained confident that feralization could be accomplished. In 1936 Arthur Svihla of Washington State College described a population of feral white and spotted rats surviving on the Hawaiian island of Lanai, where, he argued, they thrived only by the benefit of “negative natural selection” pressures. Jeff Minkler of the University of Minnesota and F.D. Pease of the Missoula, Montana Health Department described a colony of some 2000 feral albino rats living at a dumping ground near Missoula, presumably begun with laboratory stock cast off from the nearby University of Montana. Minkler and Pease also concluded that the albino colony had only been able to survive for lack of predators or competition from wild gray Norway rats. For Donaldson’s feralization studies see Section 1 of this volume; see also Svihla, A. The Occurrence of Albino and Spotted Rats Under Feral Conditions. The American Naturalist. 1936. Vol.70(729):403-404 and Minkler, J. and Pease, F.D. A Colony of Albino Rats Living Under Feral Conditions. Science. 1938. Vol.87(2264):460-461. 758 Boice, R. Behavioral Comparability. p.551. 759 Ibid.

359 early experience with traditionally psychobiological concerns like ecological relevance in

“comparisons across wild and captive contexts.”760 He (and others of his generation including Edward O. Price and Bennett Galef) also provided a more sophisticated view of domestication’s psychological effects by incorporating factors such as sociality and maternal influence into his studies. However, despite Boice’s novel approach to studying domestication, his assertions of its intrinsic interest to psychologists, and his reflexivity concerning its association with degeneration, he maintained that its primary value was allegorical. He argued in his first review of domestication in contemporary psychology that notwithstanding the generalities about the albino rat that had been rightly debunked,

“[t]here is one way the domestic rat could be a good model that has been strangely overlooked by psychologists: The domestic rat may be a good model for domestic man!”761

Boice’s turn to the human subject should not be surprising. In comparative psychologists’ work with animals, they seemed always to be working toward the ancient metaphysical maxim: know thyself. Boice seemed to grudgingly accept that comparative psychology would remain committed to this narrow goal in his response to Lockard’s obituary for the field: “[a]nimals will perhaps always have limited value as models for human behavior but only as models, not as animals.”762 While behaviorists were able to approach this self-knowledge through a presumed generality of behavioral processes among vertebrates, psychobiologists, concerned with the particularities of phylogenetic

760 Boice, R. Captivity and Feralization. Psychological Bulletin. 1981. Vol.89(3):418, 419. 761 Boice, R. Domestication Psychological Bulletin. 1973. Vol.80(3):227. Boice’s statement seems almost incredible given his familiarity with the literature on domestication, especially works by Willard Small, Curt Richter, Helen Spurway, and Irenäus Eibl-Eibesfeldt (a disciple of Konrad Lorenz). Surely, he had encountered the domestication analogy before. 762 Boice, R. On the Fall of Comparative Psychology. p.858.

360 history and ecological milieu, sought evidence of genuine homologies between animal and human behavior. Failing concrete evidence of continuity, they often relied on analogy to suggest the relevance of their research on animals for man. Thus anthropomorphism became a more common conceit among psychobiologists than the ostensibly more objective behaviorists. For instance, at the turn of the century Small suggested it was folly, and worse yet, affectation to avoid interpreting animal behavior in human terms, “for we have no other way.”763 No dogmatic canon or extent of experimental control would unburden the psychologist of the painstaking task of observing individual animals and analyzing their behaviors or remove the lens of human interpretation.

Weighing the effects of domestication on animal behavior only further ensured that man would bear a role in the interpretation. It was unavoidable, or so Helen

Spurway argued at midcentury: “All… cultivations can be considered as commensalisms of which the common feature is the dominant partner, Homo sapiens. Therefore they cannot be discussed without continual attention to our own behavior… The adjective

‘domestic’ describes human behavior to the commensal,” not the other way round.764

Culpability inhered in this arrangement. What the animal became was a reflection of the master’s conduct, hence “[t]he application of pejorative (and I suggest guilt-ridden) adjectives to these animals,” as Spurway explained.765

It was the particular difficulty of analyzing domestication, especially for those

20th century scientists trying to determine its effects on behavior, to distinguish the literal

763 Small, W.S. An Experimental Study…II p.228. 764 Spurway, H. The Causes of Domestication. p.327, 355. 765 Ibid. p.356.

361 from the figurative. Domestication described, however crudely, the concrete changes in the physiology and behavior of an animal line. It simultaneously described a relationship between human agency and natural order, such that even as methods emerged to measure and evaluate the extent of those more concrete changes, domestication evoked familiar, value-laden, and sometimes contradictory impressions of the human condition.

Questions of subjugation, transcendence, and free will followed discussion of stereotyped behaviors and changes in glandular function. Domestication, as a human activity whose consequences were borne literally in the flesh, resonated with the long-standing tension within American animal behavior study over the appropriateness of experimental control and the expression of free or “natural” behavior. Moreover, it remained, even in the confines of the laboratory, a potent model for understanding the effects of human agency, whether triumphant or improvident.

362

Conclusion

[B]iologists and psychologists before the pet terrier or hunted fox often become like Samson shorn… [Anecdotal] observations may tell us, if the observer is perfectly reliable, that a certain thing takes place, but they cannot assure us that it will take place universally among the animals… Nothing but carefully designed crucial experiments can.

-E.L. Thorndike, 1911.766

Among psychologists as among biologists there are those who are willing to shut themselves up in the narrow lane of experiment… That a pair of eyes is not all that is requisite as an observer, Dr. Thorndike’s work but too pointedly exemplifies. I venture to think that in all cases it is a question of whose eyes, or, in other words, the training those eyes have had, and still more of the intellect that passes judgment on what is seen.

-T.W. Mills, 1899.767

Although recent scholarship has largely dispensed with the view of domestication as predicated on control, concepts of domestication leading well into the 20th century were grounded in this view, such that the tractability and putative universality of some laboratory animals became synonymous with the experimental ethos of control.

Domestication, to the extent it was addressed at all, was frequently regarded as an innocuous assumption, preliminary to objective experimental work. As we have seen in the preceding pages, however, assumptions about domestication were never inconsequential.

Robert Boice described the situation for his own field of comparative psychology as late as 1980: “There is a quandary here. The same researchers who insist on empirical evidence for their own specialized study of behavior have been content to speculate about

766 Thorndike, E.L. Animal Intelligence: Experimental Studies. New York: Macmillan and Co., 1911. p.25. 767 Mills, T.W. The Nature of Animal Intelligence and the Methods of Investigating It. Psychological Review. 1899. Vol.6(3):262-263.

363 the effects of domestication on behavior. In fact, domestication is a researchable phenomenon.”768 The question Boice failed to ask, one that has driven much of the present volume, is whether subjecting domestication to experiment would quell speculation about its effects. Of course, by 1980, numerous researchers tried to show that domestication was amenable to experiment. What they demonstrated in the process, perhaps more forcefully than any single conclusion from their tests and trials, was that domestication did not instantiate experimental ideals of control and objectivity; it forced their reevaluation.

In a positivistic view that emerged around the turn of the 20th century, it might be expected that experiment would have purged folk wisdom, breeders’ superstitions, specious tropes and analogies, and other “sordid interests” from the scientific understanding of domestication. Metaphors, though, are persistent. The cultural resonances of domestication proved superbly adaptable in the hands of researchers charged with describing the phenomenon objectively. These investigators predicated discussions of the social disparity of the sexes, miscegenation and race theory, and the promise and anxieties of modernity upon their studies of domesticated animals, surely more than the evidence merited.

To point out the speculations of domestication researchers is not to decry their interpretations or to deny their science was properly experimental. It is, rather, to recognize the frequently unacknowledged assumptions that informed this narrow field of experimental study. It is to question from whence these ideas came, rather than assume the knowledge from these studies emerged entirely within the parameters of the

768 Boice, R. Domestication and Degeneracy. p.84.

364 experiments themselves. Finally, it is to acknowledge that scientific understanding of domestication was not extricated by fiat from its wider cultural understanding.

To recall a question posed at the beginning of the present investigation, how did these cultural notions about domestication coincide and conflict with the ideals of experimental biology and psychology that emerged around the turn of the 20th century?

Their interaction, it is clear, did not follow a single path, but was significantly determined by the local contexts of domestication researchers, including the particular social and political motivations of the investigator. At the same time, the experimental investigation of domestication was not an atomized experience. The cultural identity of domestication was by no means universal, but some views extended beyond intensely local contexts, and discussion of domestication in social and scientific contexts made certain interpretations more common. Thus, while different investigators rarely agreed on the specific effects of animal domestication, the notion that their experimental results provided a window into human nature, agency, and posterity was a theme assumed by many, bemoaned by some, and familiar to all.

In the first half of the 20th century, domestication researchers could claim support for their generalizations by appealing to the purported universality of domesticative effects. They emphasized common patterns across a range of domesticated species: coloration patterns, the retention of juvenile characters into adulthood, breaking with seasonal reproductive patterns, neuropathic ticks, and changes in temperament and intelligence, many of which co-occurred. These commonalities reinforced the idea that domestication was a shared experience, that if relieved of the struggle for existence

(simultaneously an assumption and a generalization), animals would respond predictably.

365

Critics at least from the middle of the 20th century have argued that the focus on common features of domestication overlooked important distinctions, conveniently neglected species that do not share in these features, and failed to consider more problematic examples such as semi-domesticated species. 769 Such common patterns and the interpretations they inspire, however, remain popular even in the recent literature on domestication, as evidenced by German zoologist Helmut Hemmer in 1990:

There is a network of relationships that links seemingly independent factors such as stress and psychosocial tolerance, behavioural flexibility, activity and intensity of action, aptitude for life in social groups and differentiation of social relationships, sexual and aggressive reactions, and even pigmentation and bodily development, into a closely interwoven system. That this network could also serve to provide a more profound knowledge of human nature independently of the ‘domestic animal’ phenomenon may be mentioned in passing. The parallels- and any ideas implicit in them- will not escape the attentive reader.770

Cultural ideas about domestication encouraged similar interpretations. It is, of course, too easy to point to a nebulous set of ideas floating in the cultural ether and simultaneously too difficult to defend or demonstrate the precise effects of such factors.

The notions of a late 19th century East London dog fancier were not the same as those of a contemporary goatherd in Eastern Europe. For the historical actors featured in this work, Americans working primarily at the border of biology and psychology in the first half of the 20th century, local circumstances and particular social and political motivations were no less important.771 Their backgrounds must be understood

769 For an early critique, see Spurway, H. The Causes of Domestication. p.354-355. 770 Hemmer, H. Domestication: The Decline of Environmental Appreciation. Cambridge: Cambridge University Press, 1990. p.viii. 771 An interesting extension of the current study would be to compare the interpretations of American domestication researchers with those of similarly interested scientists in other countries. These might include studies of 19th century acclimatization experiments in France, concurrent studies of domesticated animal behavior and physiology in Germany, and the longitudinal study of

366 individually, but the ideas they ascribed to domestication were not independently conceived. These investigators encountered notions of domestication so common and diffuse as to engender renunciations of pervasive superstitions and old wives’ tales or, alternatively, unreflective acceptance of such commonplace ideas. The power of metaphors and analogies, which these investigators employed routinely, relied in part on their cultural familiarity.

Commonalities in domestication researchers’ interpretations also rested upon similarities in their philosophical approaches to biology. None of these investigators were adherents of what the late Philip J. Pauly termed the engineering ideal in American biology (or its intellectual heir in psychology, Behaviorism). Their view of domestication precluded it. In Section III of this work, I identified a “psychobiological” alternative to the engineering ideal, Behaviorism, and extreme reductive and positivist approaches to biology and psychology in general, which by the middle 1920s occupied a minority stake in biobehavioral studies in the United States. Researchers holding this view were no less committed to experiment or intellectual rigor than those possessed of the engineering ideal, but they rejected the latter’s assumption that rigor and objectivity were synonymous with control. To the psychobiologist, the laboratory was not a

“placeless place” that derived authority from its putative neutrality but a new set of contingencies to which animals were forced to adapt themselves. The animals were, in turn, the products of countless adaptive changes prior to their captivity. The psychobiologists’ thinking about animals and their conditions of life was historical,

behaviorally selected foxes, mink, and rats conducted by Dmitri Belyaev and his associates in the Soviet Union from the middle of the 20th century. These may shed more light on the importance of national context in explaining concepts of domestication and its application to the human world.

367 contingent, and relational. This thinking drove their concern for holism, for the integrated, adapted nature of life, for the relations between body and mind and between the individual and society. It was tied also to their fondness for analogizing.

That the psychobiologist considered not only living nature, but knowledge to be contingent and relational, Willard Small’s example is perhaps the best illustration. At the turn of the 20th century, Small was at the forefront of American experimental psychology and at the same time was an outspoken critic of the unwarranted authority of experimental results in the field. In his mind, knowledge did not emerge fully formed from the narrow parameters of the experiment, but was circumscribed by the

“prepossessions and limitations” of the investigator. His defense of the experiential basis of interpretation bears repeating, as a window into psychobiological thinking in general:

Is not a certain amount of chastened anthropomorphism a wholesome specific, a kind of saving grace against the scientific pedantry that thinks to create a new science of comparative psychology with the imperfect instruments of experiment and the law of parsimony[?]… The real difficulty lies not in the tendency to interpret animal intelligence in terms of human experience, for we have no other way, but in the faulty and imperfect analysis of human experience.772

Small’s defense was, of course, a reaction to specific, reductive approaches to comparative psychology, namely Edward Thorndike’s positivistic experimentalism and

(C. L.) Morgan’s Canon. Institutions such as these were intended to proscribe the influence of individual experience from experimental interpretation, to render experimental results more general, more repeatable, and ostensibly more objective. As

Robert J. Richards explained, “…Morgan devised his principle as a methodological restraint on experimental observation and interpretation. The canon did not arise out of

772 Small, W.S. An Experimental Study… II. p.228.

368 experience, but scientific experience arose out of it.”773 Small’s objection to this approach was not only methodological but epistemological. He thought the particular experiences, norms, and values of the investigator would inevitably shape his or her interpretations and the knowledge they produced. Placing Procrustean limitations on scientific inquiry merely reflected one particular set of values at the expense of others.

Small recognized that investigators are actors, not benign recorders. They do not uncover knowledge, but construct it with reference to their prior experience, their values, and motivations. This is essentially the point raised by T. Wesley Mills (whom Small named as an inspiration for his own work) in the epigraph to this chapter.

Small failed to establish any school based on his “psycho-biological” view.

Indeed, following his work on domesticated and wild Norway rats he abandoned the field for a more promising career in education. In the following decades American experimental psychology counted Morgan and Thorndike, not Small, among its intellectual forebears. Still, there persisted what may be called a psychobiological alternative to overtly positivistic views at the margins of both biology and psychology. It may not be surprising to find, especially in the early decades of the 20th century, examples of interdisciplinary investigators questioning the validity of traditional distinctions between fields whose borders they traversed. This coincided with the efflorescence of pragmatism, which defined itself largely by exposing conventional distinctions as received ideas that constrained thought, not concrete reflections of immutable differences. Many psychobiologists in these years challenged earlier dichotomies between body and mind, organism and environment, individual and society.

773 Richards, R.J. Darwin and the emergence of Evolutionary Theories of Mind and Behavior. p.395.

369

Domestication, as it simultaneously modified form, function, and behavior, seemed to reify new ways of thinking about interrelations rather than distinctions. It appealed to psychobiologists precisely because it transgressed disciplinary boundaries and questioned conventional dichotomies.

Few domestication researchers acknowledged the dynamic, contingent, and relational nature of scientific knowledge as explicitly as Small, but their work and interpretations signaled that experiment did not entail a commitment to a radically reductive view of life and, more broadly, did not unequivocally reorder the flow of knowledge, with science the source and culture the sink. We may then wonder at the example of Henry H. Donaldson, who oversaw the largest and longest-lived experimental program on domestication in the United States and concurrently contributed to the development of the albino rat as a standard experimental tool, a product of and for the laboratory. As Cheryl Logan has demonstrated, Donaldson’s initial interest in the rat was not in making a generalized mammalian model (as the animal later became), but in locating a model specifically for human neural development. The animals he brought into the laboratory were not interchangeable cogwheels in an experimental machine, but historically distinct, uniquely adapted species and strains. Even as Donaldson had helped to establish the albino rat as the mammalian model par excellence, he was sure to include the caveat: “one hastens… to disclaim the suggestion that [the Wistar Rats] furnish standards of the same type as are to be had in the physical sciences. In the very nature of the case such accuracy and constancy is unattainable, for all animals at all times are in a state of flux…. Really all that can be said… is that ‘once upon a time’ there was a group of albino rats living under moderately favorable conditions which had the various

370 characters and values here recorded.” Donaldson’s hiring at the Wistar Institute in 1906 was a harbinger of the Institute’s entry into a new experimental era, however, his appointment was as much dependent upon his sharing a pre-experimental progressive view of life with the Institute’s philosophical founders, an outlook he maintained throughout his life.

Clyde E. Keeler is, at first blush, another troubling example. Keeler proposed the most radical reorientation of the concept of animal domestication, ascribing the bulk of the complex phenomenon to one or perhaps a handful of single gene mutations and largely downplaying the roles of phylogenetic history and changing conditions of life.

However, his causal mechanism, pigment-gene-pleiotropy, situated these few genetic changes within a distinctly integrated animal. It is true that he envisioned the coat color gene as a “handle,” a tool for manipulating whole assemblies of related traits. This rhetoric is more indicative of his regard for the tangibility of the pleiotropy idea (its ability to be experienced) and his sociological (i.e. eugenical) goals than an endorsement of the engineering ideal in biology. Keeler placed a premium on experiential knowledge in his defense of pigment-gene-pleiotropy, especially the testimony of those most familiar with the animals’ heredity and behavior, their breeders and caretakers. His reliance on this manner of evidence was spurred in part by necessity, but also by a genuine appreciation for the knowledge of experience.

Curt P. Richter displayed few qualms about manipulating the terms of his experiments, either animals or apparatus, to produce a desired result. Indeed, his ingenuity in eliciting desired behaviors within the laboratory was a hallmark of his career as a physiological psychologist. He imbibed his faith in the power of the laboratory from

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John B. Watson, but never the latter’s views on the nature of animal behavior. Richter’s interest in innate behaviors remained steadfast throughout his career; they were more biologically relevant, he thought, than the conditioned responses emphasized in Watson’s

Behaviorism. His construct of domestication-as-degeneration was predicated, in fact, on the notion that the demands of civilization ill fit the innate physiological and psychological traits that adapted pre-civilized humans to the exigencies of life in the wild.

He found inspiration for this view outside of the laboratory, from a range of social and scientific precedents and in the political ideologies on display in the Baltimore social circles in which he travelled at midcentury.

While Watson exerted the greatest influence on Richter’s belief in the power of the laboratory, it was his longtime mentor, Adolf Meyer, who more fundamentally shaped his biological philosophy. If we seek concrete connections between the scattered centers of domestication research, we may find no better candidate than Meyer. In the first half of the 20th century, Meyer became the most famous explicator of psychobiology, and perhaps the least understood (like the pragmatic philosophers he admired and emulated, he was often accused of being abstruse). His influence in psychiatry was profound, establishing a view of the patient as an integrated, adapted whole, the sum, not only of biological mechanisms, but also of individual experience. True to the spirit of his pragmatist idols- Pierce, James, and Dewey- Meyer sought to undermine the dichotomies that traditionally distinguished these factors.

Meyer came closer to this philosophical position at the University of Chicago in the early 1890s, where he was hired as a part-time lecturer in Donaldson’s Department of

Neurology. At Chicago, Meyer drank heartily from the well of one of the great centers of

372 pragmatic philosophy, which saw wide application in biology, psychology, and social reform. He worked initially under Donaldson and maintained close ties to the latter’s laboratory even after he accepted a position at the Illinois Eastern Hospital for the Insane at Kankanee, some 90 miles away. In 1895 Meyer reversed the course of many of his

Chicago colleagues, including Donaldson and Charles Otis Whitman, by taking an appointment as pathologist at the Worcester State Hospital for the Insane and, jointly, as docent at Clark.

It was at Worcester, according to Cheryl Logan, that Meyer became convinced of the usefulness of the Norway rat for studying neural and psychological development.

Meyer became close friends with Clark experimental psychologist C. F. Hodge, who had employed rats in experiments on activity and fatigue prior to Meyer’s arrival. By 1896

Meyer was engaged in research on wild caught gray rats (perhaps in association with

Hodge’s student, Colin C. Stewart, who was then performing physiological experiments with gray rats), and in the following year he maintained a breeding colony of albino rats.

Logan suggests this colony may have been the first in the United States, and that Meyer likely supplied Stewart with the albino animals he used in the later rounds of his research.

In the same year Meyer convinced Donaldson of the appropriateness of the albino rat as a subject for experiments in brain growth and provided him a shipment of rats from the

Worcester colony (a second shipment followed in 1900, after the first arrived disastrously, and with this group Donaldson began his intensive study of brain growth in the rat). Meyer’s colony was also likely the source of the rats Linus Kline and Willard

Small used in their graduate research at Clark. It is possible that, in addition to providing

373 animals, he may have helped to promote a particular perspective of appropriate biobehavioral research.

Indeed, Meyer was an inveterate campaigner for his notion of psychobiology, one

“anchored in pragmatism, progressivism, and holism.”774 His greatest influence came in the field of psychiatry, but his vision and ambitions had always been much broader. He wished, in a Deweyan phrase, to do away with the “useless contrast of mental and physical,” to unite biology and psychology along with their relevant medical and social applications. Meyer’s influence on basic science was greatly extended by the administrative efforts of Alan Gregg, whose program of “psychobiology” at the

Rockefeller Foundation was largely informed by Meyer’s articulation of the subject.

Charles Stockard, whose studies of domesticated dog breeds and their hybrids promised to unite genetics, endocrinology, and behavior, benefitted from Gregg’s Meyer-inspired psychobiology.775 And it was a sense that Stockard’s research had failed to achieve its aims that inspired Gregg to fund work such as Clyde Keeler’s pleiotropic interpretation of domestication.

The point of discussing Meyer’s influence at the border of American psychology and biology is not to imagine him to be a philosophical Johnny Appleseed, a single source for psychobiological perspectives in domestication research. Individual researchers formed a true plurality. They arrived at the subject of domestication’s effects by different means, developed unique modes of understanding the problem, and interpreted its significance in different, even contradictory, ways. Domestication

774 Quotation taken from Logan, C.A. The Altered Rationale. p.11. 775 Stockard may have engaged with Meyer’s views years before becoming associated with Gregg or the Rockefeller Foundation. His first 3 years at Cornell Medical College overlapped with Meyer’s tenure there as professor of psychiatry (his position upon leaving Worcester, Massachusetts).

374 generated no formal school, community, or tradition. It remained a problem, not a creed.

However, domestication was also, unmistakably a cultural entity. Culture and cultivation share obvious etymological origins, and it was a common assumption prior to the 20th century that domestication was a prerequisite to “higher” civilization.776 The individuals who subjected domestication to experimental scrutiny in the new century continued to draw, necessarily, from culture, understood here as historian Louis Menand describes it,

“the name for a set of products, practices, and perspectives of which individuals can avail themselves.”777 This set included not only wide cultural notions of domestication, but views more closely circumscribed by the activities of well connected individuals such as

Meyer, Gregg, and Donaldson, by the minor solidarity afforded by antipathy toward overtly reductive approaches such as Behaviorism, and by the perseverance of an alternative perspective that held knowledge to be relational, not univocal. The inducement to study domestication was not simply its ability to highlight particular biological or psychological problems; this was always paired with its perceived ability to clarify, by analogy and relation, more obscure contemporary social dilemmas.

This was the legacy of turn of the 20th century biobehavioral research Meyer chose to remember at the 50th anniversary of the founding of the Journal of Comparative

Neurology in 1941. He recounted wistfully of those years surrounding 1900: “It was a period that brought together man and nature and culture as never had happened before- presenting a new and more vital perspective within science in general, a perspective

776 For an excellent demonstration of this argument immediately prior to the turn of the century, see Nathaniel Shaler’s Domesticated Animals: Their Relation to Man and to His Achievement in Civilization. New York: Scribners, 1895. 777 Menand, L. The Metaphysical Club: A Story of Ideas in America. New York: Farrar, Strous, and Giroux, 2001. p.407.

375 essential for a true science of man, more and more needed, and more and more significant and helpful. Such a perspective is our chief hope especially today in a world of enormous possibilities but also of enormous confusion and disaster.”778 Similar perspectives and hopes can be traced in the work of most domestication researchers, although they always found unique expression. Those investigators did not imagine the sole function of experiment to diminish the influence of externalities (i.e. of culture) and the idiosyncrasies of the investigator on scientific interpretation; they demonstrated, by design or by effect, that the evidence derived from experiment was to be understood through its complex relations with other frames of knowledge.

Knowledge of domestication changed dramatically over the course of the 20th century, thanks in part to its experimental investigation. Animal domestication has grown from a cautiously attended subject, to one of recognized biological and psychological import, albeit still not a central theme in either discipline. Animals have been granted greater agency in the domesticative relationship, which has been reimagined as more symmetrical and cooperative than the traditional notion of master and subject.

Domestications, especially those occurring prior to the 18th century, are increasingly explained by spontaneous modifications and unforeseen consequences rather than the careful deliberations of breeders. Unilateral explanations, invoking property and control, have largely given way to more complex and contextualized understandings of the human-animal relationship in domestication. These new interpretations appeared mostly after the middle of the 20th century, not as the result of greater experimental rigor, but rather by recognizing and challenging prior assumptions about domestication.

778 Meyer, A. The Contemporary Setting of the Pioneer. Journal of Comparative Neurology. 1941. Vol.74(1):2-3.

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The most consistent role of the domesticated animal in 20th century biobehavioral research was as a mirror unto ourselves, as it had been in countless folktales, fables, and allegories. It is in the spirit of this reciprocal understanding that domestication researchers to midcentury derived their conclusions about human will, agency, and consequence from experimental results on domesticated rats, dogs, and other animals.

After midcentury, domestication persisted as a means of inward reflection, though the focus had shifted to the field of comparative psychology. Critics argued that the ostensibly degraded nature of domesticated creatures made them especially poor candidates upon which to pin so much of the credibility of the field. Their primary interest, however, was not in denigrating the animals, but in diminishing the hegemony of

Behaviorism in comparative psychology, the assumption of generality from limited animal models, and the narrow focus on conditioned learning. Throughout the 1970s

Robert Boice guided a further introspection, this into the use of “domestication” in comparative psychology and its association with the idea of degeneracy. The importance of Boice’s inquiry was not in finally setting domestication research on a firm, objective, and truly experimental footing, but in the self-conscious reflection on the utility of domestication as a concept in biobehavioral science and its relations to the wider culture in which that science was done.

The examination of animal domestication connotes the examination of the human self. Perhaps it is impossible, as Willard Small suggested, to interpret what has happened to the animal except in the terms of human experience. To the extent that domestication researchers devoted themselves to uncovering the human implications of their work on domestic animals and brought the authority of experimental science to bear on such

377 interpretations, they ensured Small’s statement would appear true. The recognition that experiment did not divorce scientific knowledge of the effects of domestication from human experience, either in the form of cultural notions or the experience of individual investigators, does not diminish the validity of experiment in 20th century biology and psychology. Rather, it situates experiment alongside other ways of knowing that remained vital in the interpretation of domestication. It is hoped that the historical perspective provided in this work will add to the ways of knowing domestication, specifically through the experience of 20th century American biobehavioral research.

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