The Cuvier-Geoffrey Debate: French Biology in the Decades before Darwin

The Cuvier-Geoffroy Debate

Monographs on the History and Philosophy of Biology RICHARD BURIAN, RICHARD BURKHARDT, JR., RICHARD LEWONTIN, JOHN MAYNARD SMITH EDITORS

The Cuvier-Geoffroy Debate: French Biology in the Decades Before Darwin TOBY A. APPEL

Controlling Life: Jacques Loeb and the Engineering Ideal in Biology PHILIP J. PAULY Beyond the Gene: Cytoplasmic Inheritance and the Struggle for Authority in Genetics JAN SAPP

The Cuvier-Geoffroy Debate French Biology in the Decades Before Darwin

TOBY A. APPEL

New York Oxford OXFORD UNIVERSITY PRESS 1987

Oxford University Press Oxford New York Toronto Delhi Bombay Calcutta Madras Karachi Petaling Jaya Singapore Hong Kong Tokyo Nairobi Dar cs Salaam Cape Town Melbourne Auckland and associated companies in Beirut Berlin Ibadan Nicosia

Copyright © 1987 by Oxford University Press, Inc. Published by Oxford University Press, Inc., 200 Madison Avenue, New York, New York 10016 Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press.

ISBN 0-19-504138-0

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To my family

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Acknowledgments

Many individuals and institutions have contributed over the years to the making of this book. Theodore M. Brown introduced me to Cuvier and Geoffroy in his course on evolutionary biology over fifteen years ago, and has remained ever since a warm friend and advisor. John C. Greene and Frederic L. Holmes, by their strong encouragement, provided the initial impetus to undertake the project of expanding my earlier research into a monograph. With the assistance of a grant from the Secretary's Fund of the Smithsonian Institution, I was able to return to France to research manuscript material. I am especially grateful to the Hannah Institute for the History of Medicine for providing the opportunity as a Hannah post-doctoral fellow to devote a full year to research and writing, and to Toby Gelfand at the University of Ottawa for his hospitality and friendship. Several people have read the manuscript at various stages in its evolution and provided valuable advice: Richard Burian, Richard Burkhardt, William Coleman, Robert Friedel, Gerald Geison, John C. Greene, Frederic L. Holmes, David Hull, Andrew Lugg, Dov Ospovat, Philip Pauly, and Jean Chandler Smith. Paul Israel, my former colleague at the Edison Papers, helped with my computer files until they could be transferred to Washington. Silvio Bedini, Kathryn Olesko, and Pauline Mazumdar assisted with languages. Although this book relied upon the resources of numerous libraries and the generous assistance of their staffs, I am particularly grateful to the History of Medicine Division of the National Library of Medicine where I was able to do most of the final checking of sources and quotations, often during lunch break from my work at the American Physiological Society. Jean Chandler Smith, who has been traveling to Paris to complete a bibliography of the writings of Cuvier, very kindly researched a few final sources and acted as intermediary to obtain microfilm and illustrations from the Bibliotheque Centrale of the Museum d'Histoire Naturelle. Working with Oxford University Press and Biology Editor William F. Curtis has been a pleasure throughout. I am most thankful for Jonathan Harrington's thorough copyediting and for his much-welcomed expertise in anatomy and zoology. Finally, there are two people whom I have relied upon from the beginning to the end of this project. I am greatly indebted to Gerald Geison for providing guidance and encouragement at every step of the way, and to my husband Robert Friedel for his patience and support, and his critical skills and editing ability.

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Contents

Chapter 1. Introduction: A Classic Confrontation and Its Interpretations

1

Chapter 2. Cuvier and Geoffroy: Collaborators on a New Science

11

Chapter 3. "Le Legislateur de la Science": Cuvier and Functionalist Anatomy

40

Chapter 4. Geoffroy and the Emergence of Philosophical Anatomy

69

Chapter 5. The Battle Lines Are Drawn: 1820-1829

105

Chapter 6. The Debate before the Academic

143

Chapter 7. Beyond the Academie: The Many Uses of the Debate

175

Chapter 8. Teleology, Morphology, and Evolution: The Debate and the Future of Zoology Appendixes Notes Index

202 238 245 293

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The Cuvier-Geoffroy Debate

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CHAPTER 1

Introduction: A Classic Confrontation and Its Interpretations

On 2 August 1830, shortly after the conclusion of the Revolution of 1830, resulting in the fall of Charles X from the French throne, Frederic Soret of Geneva recorded a memorable conversation with his friend, the naturalistpoet Johann Wolfgang von Goethe: The news of the Revolution of July, which had already commenced, reached Weimar today, and set everyone in a commotion. I went in the course of the afternoon to Goethe's. "Now," he exclaimed as I entered, "what do you think of this great event? The volcano has come to an eruption; everything is in flames, and we no longer have a transaction behind closed doors!" "A frightful story," I replied. "But what else could be expected under such notorious circumstances and with such a ministry, than that matters would end with the expulsion of the royal family?" "We do not appear to understand each other, my good friend," replied Goethe. "I am not speaking of those people at all, but of something entirely different. I am speaking of the contest, of the highest importance for science, between Cuvier and Geoffroy Saint-Hilaire, which has come to an open rupture in the Academy." This expression of Goethe's was so unexpected that I did not know what to say, and for some minutes felt my thoughts completely at a standstill.1

The chief participants in the "great event" which so aroused the aged Goethe in Weimar were the two most noted comparative anatomists and zoologists of the day, Georges Cuvier and Etienne Geoffroy Saint-Hilaire. Their scientific rivalry, one of the most celebrated of the century, had erupted into an acrimonious public debate before the most august of scientific tribunals, the Academie des Sciences in Paris. Each Monday afternoon, at the weekly meeting of the Academie, Cuvier and Geoffroy took turns defending their views before a large and noisy audience of scientists and nonscientists, until the arguments became so heated that they agreed to call a halt. Although the formal debate before the Academie lasted little more than two months, it was the culmination of a divergence that had its roots in the closing years of the previous century, and that had been aired before the scientific community since 1820. Scientific disputes have long been an attractive subject for study by histori1

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ans because of the insights they can provide into the nature of scientific explanations and their social context. Approaches to debates have covered the full spectrum from the narrow history of scientific ideas to the purely sociological. Controversies allow the intellectual historian to clarify scientific theories and the evidence supporting them, to better appreciate the magnitude of transformations of thought, to discover the part played in science by philosophical and religious beliefs, and to make evident the assumptions and metaphysical commitments that are usually taken for granted in the normal pursuit of science. Those who take a somewhat broader purview can almost always identify personal and institutional factors that are nearly as important as the intellectual in understanding the impact of a controversy on contemporaries. And in many recent sociological studies of controversies, it has become fashionable to show that intellectual differences reflect personal and institutional loyalties, vested interests in certain ideas and methods, attempts to control patronage and resources, and other social and political interests. As Goethe's analogy of erupting volcanoes suggests, scientific controversies are valuable to the historian because they bring to light what is usually hidden beneath the surface.2 The Cuvier-Geoffroy debate provides an especially rich subject for historical study. While disagreements in science are commonplace, in every century there are a few scientific controversies that, in a critical moment, crystallize the key issues of an entire discipline. Brought to a head in the Cuvier-Geoffroy debate of 1830 was a fundamental division in the biological sciences: whether animal structure ought to be explained primarily by reference to function or by morphological laws. This controversy, by calling attention to the morphological approach to animal structure known in France and Britain as "philosophical anatomy," had a major place in the ferment of biological theorizing that preceded the advent of the doctrine of Darwinian evolution. But the debate's meaning for biology is only a part of its significance for the history of science. In the past this controversy has been interpreted primarily from the standpoint of the internal history of biology. When traditional analysis of Cuvier's and Geoffroy's scientific ideas is extended and when it is enriched by a consideration of personalities and scientific styles, of institutions and politics of science, a much fuller and more dramatic picture emerges. Not only is one led to revise prevalent interpretations of the debate's outcome and its place in the history of biological ideas, but one discovers a host of other issues, both scientific and extra-scientific, that were also at stake in the conflict. The story of the debate, when it is more fully told, has much to say about the national style and structure of French biology, and more generally, about the dynamic interplay of people, nature, and social context that constitutes the scientific enterprise. The Cuvier-Geoffroy debate was first studied by scientists and historians for its direct bearing on the controversy over the evolution of species. Geoffroy followed Jean-Baptiste Lamarck in a line of French speculative evolutionists, while Cuvier has come down in history as the archfoe of transformism and Lamarck's chief opponent. Historians of the late nineteenth century, those who accepted evolution as well as those who did not, interpreted the debate as a

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3

pre-Darwinian testing of the doctrine of transmutation of species. Some historians, such as Franck Bourdier, still contend that the dispute was primarily concerned with evolution.3 Most recent studies, however, have minimized the relevance of the evolution controversy, for evolution was scarcely mentioned in the proceedings of 1830. Following the lead of the biologist/historian Edward Stuart Russell, most recent historians prefer to regard the conflict as a confrontation between a teleological and a morphological approach to nature. Russell, author of the now classic Form and Function (1916), saw the Cuvier-Geoffroy debate as the climax of a struggle in biology which could be traced back to the idealist Plato and the teleologist Aristotle: "The contrast between the teleological attitude, with its insistence on the priority of function to structure, and the morphological attitude, with its conviction of the priority of structure to function, is one of the most fundamental in biology. Cuvier and Geoffroy are the greatest representatives of these opposing views." As a Lamarckian and an opponent of reductionism in biology, Russell did not take a neutral stand. Despite Cuvier's antievolutionism, Russell placed him on a pedestal because Cuvier had given precedence to holistic, functional considerations rather than strictly morphological considerations. Geoffroy, by contrast, had tried to create a science of pure morphology, and his failure, amply revealed by the controversy of 1830, was proof that pure morphology was unworkable. The debate was for Russell "a struggle between 'comparative anatomy' and 'morphology,' between the commonsense teleological view of nature and the abstract transcendental." As a practicing biologist, Russell was intent upon demonstrating by an appeal to history that the teleological approach of Aristotle and Cuvier was the right course for the future of biology.4 In recent studies, Cuvier and Geoffroy are portrayed as more nearly complementary, but important aspects of the conflict of their ideas are still misunderstood. Historians often interpret Cuvier as more of an adherent of abstract types than he in fact was, and hence they underestimate the extent of his disagreement with Geoffroy. Geoffroy, in turn, is often depicted as a French Naturphilosoph whose fanciful morphological doctrines were quite rightly put to rest by the common-sense arguments of his opponent.5 Most modern historians still agree with Russell that Cuvier was able to "score a decisive victory."6 A closer study of the debate, however, shows that both Cuvier and Geoffroy defended extreme positions. The significance of the debate for the future of biology is that it laid bare the untenability of both Cuvier's and Geoffroy's views, and challenged naturalists to come to a creative resolution of them. While Geoffroy's more speculative doctrine of the "unity of composition" in the animal kingdom may have been rejected in 1830, his morphological program, known as "philosophical anatomy," eventually became an integral part of biological science in both France and Britain. Only in the last few years have historians come to appreciate the role of morphology, and hence of Geoffroy, in early nineteenth-century biology.7 From his arrival in Paris in 1795 and his installation as acting professor in

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the chair of anatomy of animals at the Museum d'Histoire Naturelle, Cuvier presided over a renaissance of comparative anatomy. Up to this time, anatomy had been largely an adjunct of medicine. By treating anatomy in a comparative manner, Cuvier hoped to make of it a science with close ties to natural history. Through studying the various structural means by which animals carried out the major processes of life, Cuvier believed he could establish rational criteria for a "natural system" of classification. On the basis of the functioning of the nervous system, Cuvier posited four basic plans of organization, or embranchements—the vertebrate, articulate, mollusk, and radiate plans. He insisted that there were no transitions between plans, and that even within an embranchement, the number and arrangement of pieces varied to suit functional requirements. Every part of an animal, Cuvier believed, was designed to contribute to the functional integrity of the animal or to adapt it to its environment. Whenever a new part was required for functional purposes, the Creator was free to fashion a new and appropriate organ. For Cuvier, function was paramount: the animal's needs sufficed to determine its structure. Geoffroy, professor of zoology at the Museum, challenged this traditional functionalist view of nature with a new set of doctrines, "philosophical" or "transcendental anatomy," which were at the heart of the science of morphology. In the first decade of the nineteenth century, he proposed to take comparative anatomy beyond description and classification and, by the process of abstraction, to make it truly "philosophical." Instead of describing each class of animals separately, using different names for different structures in each class, Geoffroy believed one could discover a generalized vertebrate anatomy, a single structural plan that could be traced throughout the vertebrates. The key to determining the ideal plan, Geoffroy showed, was to ignore the form and the function of the parts and concentrate instead on the connections between parts. Geoffroy's "principle of connections" became, in fact, the main guide in the nineteenth century to determining homological relationships. In 1820 Geoffroy extended "unity of composition" from the vertebrates to the invertebrates by assuming that the latter respresented stages in the embryological development of higher animals. Thus, invertebrates shared the same basic plan of organization, though they lacked many of the structural elements of higher animals in fully developed form. For Geoffroy, then, the unity of plan in the animal kingdom preceded particular modifications of the plan to suit functional requirements. Philosophical anatomy had as its aim the discovery of resemblances, often obscured from the naturalist by changes of form and function, between elements of animal structure. Geoffroy became the chief spokesman for philosophical anatomy in France. Its key concept was that of homology. Naturalists sought homologies between the parts of different animals, homologies between the parts in a single animal, homologies between the structures in the fetus of higher animals and the adult form of lower animals, and homologies between structures in so-called monsters and those in normal animals. For the philosophical anatomists, animal organization appeared to have a constancy in the

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5

number and arrangement of parts that was independent of the form of the parts and the uses to which they were put. Although Cuvier exhibited a better command of the details of anatomy during the 1830 debate, and while his arguments were presented in a more logical fashion, Geoffroy and his followers were by no means routed. Geoffroy's morphology, in a modified form, became an integral part of the mainstream of French zoology. No longer could animal structure be explained by the "conditions of existence" alone. Zoologists after Cuvier no longer accepted his dictum that morphological laws unduly restricted the sphere of action of the Creator. Comparative anatomists came to recognize the many evidences of unity of plan that transcended similarities of function. The leaders of French zoology after 1830—in particular, Henri de Blainville, Isidore Geoffroy Saint-Hilaire, Pierre Flourens, and Henri Milne Edwards—reconciled the extremes presented by Geoffroy and Cuvier within a nonevolutionary framework. In England, in a different cultural context, the comparative anatomist Richard Owen also formulated a compromise between the views of Cuvier and Geoffroy, one that played an important and, until recently, overlooked role in the development of Darwin's thought. Dov Ospovat has argued that the rejection of strict teleological functionalism was instrumental in preparing naturalists for the acceptance of Darwinian evolution. For instead of referring each structure to adaptations provided by the Creator to supply the specific needs of each species, the naturalist was free to search for general laws of animal organization. The morphological approach as it was developed in France, Germany, and England could therefore be said to be an innovative force in early nineteenth-century biology. Morphology not only provided Darwin with some of his strongest evidence for evolution, but it also, as Ospovat has suggested, led him to rethink his theory of natural selection in the two decades before he published On the Origin of Species.8 Viewed from a later perspective, the Cuvier-Geoffroy debate was therefore an important chapter in the history of evolutionary biology. The prevailing interpretation of the debate as teleology versus morphology, while it captures the principal anatomical disagreement, is nonetheless partial and misleading because it conceives of the debate from the post-1859 vantage point, and solely as a contest of scientific ideas. Russell, in particular, wrote as though the confrontation might have arisen at any time and in any place. Goethe and other contemporaries were well aware that much more was at stake than a point of anatomy. Considerations of time and place, far from being irrelevant, are essential to understanding the controversy. As Goethe intimated by his allusions to volcanoes and transactions behind closed doors, the CuvierGeoffroy debate, like all great scientific controversies, was not limited to a difference of scientific ideas or even of scientific philosophies broadly conceived. To understand what the debate meant for Cuvier, Geoffroy, and their contemporaries in France, it is necessary to go beyond the anatomical arguments. The controversy must be viewed in its widest context, taking into account not only ideas, but also personalities, institutions, the careers of the participants and

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THE CUVIER-GEOFFROY DEBATE

their colleagues and would-be colleagues, the relations of science and religion, the relations of science and government, the micropolitics of science, and the macropolitics of nineteenth-century France. Though it certainly had implications for biology elsewhere, the debate was above all a peculiarly French scientific controversy. This has not always been sufficiently appreciated. Cuvier has been assimilated by Ospovat to British natural theology, while Geoffroy has been alternately depicted as a precursor of Darwin and a follower of German Naturphilosophie. The primary context for the Cuvier-Geoffroy debate was neither Britain with its individualism and Protestant natural theology, nor Germany with its philosophical idealism and competing centers of university learning, but the centralized, public scientific institutions of Catholic France. The interpretation of the debate as teleology versus morphology, although we will use it for convenience, derives, not from the French, but from the British context of the 1830s and 1840s. In some ways this interpretation says more about Owen and Darwin than about Cuvier and Geoffroy. In Britain, Owen and others had to find a compromise between Geoffroy's extreme version of philosophical anatomy and the prevailing dogma of Paleyan natural theology that all structure was adaptation. Both Owen and Darwin thought of themselves as reconciling teleology and morphology. In France, where natural theology was not part of the common culture, the reconciliation took on a different character and was effected in a much less explicit manner. French contemporaries were more likely to interpret the debate as a contest between one plan of nature and four (if they favored Cuvier), or as a conflict between facts and ideas or analysis and synthesis (if they favored Geoffroy). The Cuvier-Geoffroy debate, like many celebrated scientific disagreements, was a highly personal affair. The antagonists were wholly unlike in temperament. Cuvier was cold in manner, crisply logical in his writings, and handled himself with an ingrained sense of his authority, while Geoffroy was intuitive and impetuous, and wrote in an obscure and diffuse style. Both, however, were overly sensitive and quick to take offense. The controversy was sustained for more than a decade because both Cuvier and Geoffroy wished it to be sustained. The fireworks died down as soon as the two leading actors left the scene. But although personalities were essential in maintaining the passions of debate, the controversy was by no means a simple personality conflict. On a broad plane, the future of biology was affected by the outcome. On a narrower level, other aspiring French naturalists were caught in the middle of the battle, and their careers were shaped by their responses to both the men and the issues. Differences of scientific methodology played a large role in the controversy. Cuvier, for both professional and political reasons, championed an extreme empirical methodology. The naturalist's role, as he saw it, was to gather "positive facts." Hypotheses could lead only to unfounded and perhaps dangerous generalizations. Cuvier thus sought to circumscribe the questions a naturalist might ask and the methods that he might employ to answer them. Geoffroy, from the outset of his career, saw science in different terms. Facts were only

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7

the building blocks of science; the essence of science was ideas. No question ought to be placed outside the realm of scientific inquiry. Because of Cuvier's and Geoffroy's divergent positions on scientific methodology, many contemporaries, Goethe among them, interpreted the debate not as teleology versus morphology but as facts versus ideas or analysis versus synthesis. An understanding of the differing approaches of Cuvier and Geoffroy to the relationship between God and nature is also indispensable for making sense of the debate. Scientific ideas in this period, especially in the life sciences, cannot neatly be separated from religious belief. Cuvier, a Protestant in a Catholic milieu, held the traditional belief that the Creator could not be constrained in His activity by any such pretended laws as the chain of being or the unity of plan in the animal kingdom. The Creator foresaw the needs of each species and created just those organs that were necessary to carry them out. Although Cuvier wrote in the Aristotelian language of "conditions of existence" or "final causes," these concepts in fact had a religious basis. Geoffroy, too, was in part motivated by religious concerns. A child of the Enlightenment, he was attracted to a Deist view of nature similar to that of Buffon and Lamarck. God had in the beginning established laws, and nature was then left to unfold in accordance with them. Philosophical anatomy, with its emphasis on transcendent laws of organization, appeared to Cuvier religiously suspect because it hinted of pantheism or, worse, materialism. No less central to the controversy was the major transformation that had taken place since the Revolution in the institutional context of French science. Cuvier's and Geoffroy's scientific doctrines became increasingly polarized against the background of a transition from the broad, philosophical life science of the eighteenth century to the more specialized, institutionalized, and professional science of the nineteenth century. While Geoffroy remained sympathetic to Buffon and the eighteenth-century philosophes, Cuvier quickly emerged as the undisputed leader of a new generation of professionally conscious zoologists and comparative anatomists. Founded in 1793, the amply endowed Museum d'Histoire Naturelle, where Cuvier and Geoffroy lived, taught, and worked together for thirty-seven years, quickly became the European center for research in zoology. The Academie des Sciences, reorganized in 1795, became the supreme arbiter of science in France, its weekly meetings attended by all the leading scientists of the capital. Other institutional changes, providing educational opportunities and new avenues of employment, enabled a small but active community of zoologists and comparative anatomists to emerge and flourish. The transformation in the institutions and content of zoology from 1793 to 1815 was accompanied by an increase of professionalism, characterized by the setting of community standards as to what constituted appropriate research. Cuvier's Lectures on Comparative Anatomy, Researches on Fossil Bones, and his compendium of systematic zoology, The Animal Kingdom, were especially instrumental in providing a framework within which other naturalists might work. This restriction of the field was not accomplished without some tension and resistance, particularly from zoologists, among them Geoffroy, trained before

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the founding of the Museum. When the Museum was established in 1793, no one in France was adequately prepared to teach zoology. By a fortuitous set of circumstances, the first permanent chairs of the subject in France were given to Geoffroy, aged twenty-one, who had studied a bit of mineralogy, and to Lamarck, then over fifty, who was known as a botanist. Both refused to limit their work to the memoirs of taxonomy and descriptive anatomy that were expected of them. Lamarck, Richard Burkhardt has shown, was partly to blame for the poor reception of his evolutionary theories, for he refused to present his arguments in the form that had come to be insisted upon by the community of naturalists. He wrote, instead, in a deductive manner, as though he had given up all hope of convincing his colleagues. When his theories were not wholly ignored, they were ridiculed by Cuvier and his followers.9 Geoffroy adapted better to the professional demands of his colleagues, but he too had difficulty controlling his desire, present from the outset of his career, to reach beyond zoology and comparative anatomy in order to search for the most fundamental laws of nature. As his career progressed, he developed an increasingly elaborate and speculative synthesis. He attacked vitalism and preformation, and advocated the unity of physiological and psychological processes, the unity of plan of organization in the animal kingdom, a theory of evolution based on the recapitulation theory of the German Naturphilosophen, and, finally, a theory on the action of subtle fluids in the universe. At several points in his career, Geoffroy stifled his speculative impulses in order to maintain the approval of his colleagues, but in his old age he gave full vent to his natural inclinations. Contemporaries' responses to the biological issues brought out in the debate were thus colored by their reactions to Geoffroy's materialistic philosophy, his romanticized notion of science, and his at times nonprofessional style. The controversy among naturalists over philosophical anatomy was fought out not just in terms of ideas, but also in the arena of scientific politics. With the founding of a new set of institutions of science during the Revolution and the Empire, French science became establishment science. French science, unlike English science, was centralized and largely confined to government institutions. Access to a career was controlled by the patronage of a small number of individuals. An aspiring naturalist was thus not free to come to an impartial decision on the merits of the controversy, for his evaluation of the arguments might have an important bearing on the course of his career. Cuvier, in particular, exploited all the authority of his multiple positions— permanent secretary of the Academie des Sciences, professor at the Museum and College de France, member of the Council of the University, and friend to those in political power—to combat the morphological approach to anatomy, which he condemned as false, speculative, and potentially dangerous. A number of savants who were initially disposed toward philosophical anatomy moderated their views in order to be assured of Cuvier's patronage. Critics of the nineteenth-century scientific establishment accounted for Cuvier's apparent success in the debate by pointing to the usurping of science by an elite that misused its power and authority to suppress new ideas. Geof-

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froy's supporters in the 1830s and 1840s, many of them outsiders to the scientific community, looked beyond the main anatomical conflict and regarded Geoffroy as the leader of an entire movement of synthetic and progressive science. Many of these advocates—literary figures, would-be scientists, and scientific popularizers—resented what had happened to science in France since the eighteenth century. In some sense, nature had been taken out of their hands and had become the property of specialists, who no longer seemed to be asking relevant questions. The sciences were developed under the aegis of the government in centralized public institutions that seemed to exert an autocratic influence over the dissemination of ideas. It was natural to blame these institutions and their supposed leader, Cuvier, for the decline of true science. Geoffroy was depicted by some of these critics as a philosopher dedicated to unraveling the mysteries of nature for the common man, while Cuvier was seen as an elitist fact collector, upholder of Biblical orthodoxy, manipulator of patronage, and suppressor of the ideas of men like Lamarck and Geoffroy. The social uses of the controversy by outsiders bring us to the relationship of the debate to the broader context of French society and culture. To fully explore the many points of contact would require an extended study of French society, which is not the intent of this work, but a few suggestions on the relationship between the debate and politics can be offered. A political elitist who advocated government by expert administrators, Cuvier feared above all the return of revolution and rule by the uncontrolled mob. Scientific ideas, if unrestrained, might be exploited by unscrupulous men to weaken the social order. Thus, it was partly for political reasons that Cuvier insisted so strongly on limiting science to "positive facts." The buildup of the controversy in the 1820s can be tied to the political and religious tensions of this period, and Cuvier's willingness to descend into the arena in 1830 can be linked to his anxiety in the months preceding the Revolution of 1830. The considerable support Geoffroy received at that time can be attributed in part to the association in the public mind between Geoffroy's view of nature and the liberal ideology of the 1830 Revolution. Like most important scientific controversies, the Cuvier-Geoffroy debate functioned on many levels at once, and it functioned in different ways for different individuals. There is no single "right" set of labels to apply to the principals in the debate, for the debate had different meanings to Cuvier and to Geoffroy, and still other meanings to contemporaries. There was not one interpretation, but many of the Cuvier-Geoffroy debate. In this book we shall examine the debate mainly at the levels of ideas, scientific methodology, and the personalities, institutions, and micropolitics of science. Above all we shall be concerned with the Cuvier-Geoffroy controversy as it relates to the development and integration into biology of the morphological approach to nature. The primary thread of the narrative will treat the rise of the major concepts of philosophical anatomy in the work of Geoffroy Saint-Hilaire and their elaboration in the work of Blainville, Jules-Cesar Savigny, Victor Audouin, E.R.A. Serres, Milne Edwards, and a number of lesser-known figures. We shall trace the increasing tension between the promoters of the morphological view of

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anatomy and the advocates of the more orthodox school of comparative anatomy led by Cuvier, from the first public break between Cuvier and Geoffroy in 1820, through the eruption of the public confrontation in 1830, to the crescendo of polemics from 1830 until Cuvier's death in 1832. We shall explore the many faces of the controversy as it appeared to contemporaries, both scientific and nonscientific. Finally we shall examine how naturalists after 1830, both in France and in Britain, came to terms with the critical issues brought to a clear focus by the debate before the Academie, and how, by appropriating a modified form of philosophical anatomy, they brought new direction to the life sciences.

CHAPTER 2

Cuvier and Geoffroy: Collaborators on a New Science

When Etienne Geoffroy Saint-Hilaire at the age of twenty-one was offered a chair of zoology at the newly created Museum National d'Histoire Naturelle, he at first declined for fear that he would not be able to handle the task. LouisJean-Marie Daubenton, confident of his protege's abilities, was said to have insisted that he not refuse: You will not do it. I have the authority of a father over you, and I will take upon myself the responsibility for the event. No one has yet taught zoology in Paris. Guideposts scarcely exist to make of it science. Everything is still to be done. Dare to undertake the task so that in twenty years one can say: Zoology is a French science.1

Daubenton's assessment of the state of zoology, exaggerated for the effect on his young disciple, was nevertheless by and large just. Although one could find the ingredients of a science of zoology in eighteenth-century France, it was still an unformed, underdeveloped field conducted by semiprofessionals or amateurs with meager resources. The founding of the Museum d'Histoire Naturelle by the Decree of the Revolutionary Convention of 10 June 1793, it was agreed by both contemporaries and historians, marked a watershed in the history of zoology.2 Zoology did in fact become for a time, as Daubenton hoped, a French science. The Museum with its ever-increasing collections and magnificent facilities for research became the world center for the study of the life sciences. It was in France from 1793 to 1830 that the foundations were laid for the study of comparative anatomy, paleontology, and morphology, and that the major outlines of modern zoological taxonomy were established.3 Thrust, almost by accident, into leading roles in the creation of a new science were three men: Geoffroy, Lamarck, and Cuvier. Geoffroy and Lamarck, both unprepared to teach zoology in 1793, received their professorships because they had held minor appointments at the Jardin du Roi, the predecessor of the Museum. Two years later Cuvier was installed at the Museum as acting professor of comparative anatomy, ironically, as it turned out, with support chiefly 11

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from Geoffroy and Lamarck. An outsider, educated in Germany, Cuvier could scarcely have hoped to establish himself in Paris in circumstances other than those created by the flux of the Revolutionary period. It was, however, Cuvier, the interloper, rather than the two charter professors, who took most heed of Daubenton's prophetic advice. With his vastly superior knowledge of zoology and his skill in making contacts, he quickly surpassed Lamarck and Geoffroy in scientific prominence. When the Academie des Sciences was reinstituted in 1796, Cuvier was made a member, while Geoffroy had to wait until 1807 for a seat. The future antagonists began as the closest of friends. When, with Geoffroy's help, Cuvier became substitute lecturer (suppleant) for A.-L.-F. Mertrud, professor of animal anatomy at the Museum, he moved from his hotel into Geoffroy's lodgings. They lived together for several months, and in the course of two years published five papers together. Both were engaged in the great task of increasing the collections of the Museum and in creating a foundation for the future study of zoology. In a scientific community dominated by the physical sciences, both Cuvier and Geoffroy clearly perceived the need to improve the relatively low status of the life sciences, especially zoology. Their common goal was to make zoology a "philosophical" rather than a merely descriptive science, to raise it above the task of Linnaean classification. Only if zoology were based on principles could it hope to attain the respect accorded to such sciences as astronomy, experimental physics, chemistry, and, most recently, mineralogy. Although Cuvier and Geoffroy then appeared to be working in harmony on a common program, to a perceptive observer the sources of their future disagreement might already be seen. These differences were most clearly evident in their divergent attitudes to classification, the chief activity of the naturalist. From his first scientific papers, Geoffroy assumed, with his mentors Buffon and Daubenton, that classification, while necessary, was ultimately arbitrary. No classification, however complicated, could adequately mirror nature. In a 1795 paper, he presented the first hints of "philosophical anatomy" when he suggested that an underlying unity of plan could be traced throughout the animal kingdom. Cuvier, in contrast, was animated by the goal of creating a unique natural system of classification based on function. In his early papers on the orders of mammals (with Geoffroy) and the classes of invertebrates, Cuvier based his divisions on the principle of "subordination of characters," determined by functional considerations. There might also be detected in these early years their divergent attitudes toward theorizing. Geoffroy's desire to discourse on broad theories of nature was deep-seated, although in his first years as a professor he kept his speculations under restraint, probably on the advice of Daubenton. Cuvier, however, was sufficiently chastened by the experience of the French Revolution to begin identifying unbridled scientific theorizing with social and moral chaos. In their years of collaboration and close friendship, these differences were scarcely perceptible, for both men were absorbed by the challenges in laying the groundwork of a new science. In hindsight, however, there can be seen even at this time the contrast in backgrounds, training, and

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inclinations that would drive the careers and scientific outlooks of Cuvier and Geoffroy in opposite directions. Zoology in France before 1793

Before the founding of the Museum, zoology did not exist as a formal discipline in France. There were no courses, no adequate collections, no successful specialized journals, and no organized community of practitioners. In contrast, botany was already a well-established science. Not only were plants relatively easy to collect and preserve, but their study was considered of utilitarian value to both medicine and agriculture. Thus the French government had supported the teaching of botany at the faculties of medicine in Paris and the provinces as well as at the Jardin du Roi. Botany, moreover, was recognized as one of the sections of the Academie des Sciences, the members of which received state subsidies. There was as yet no section of zoology. Besides the medical and agricultural constituencies, botany had a popular constituency. Nobles, physicians, lawyers, and government functionaries maintained cabinets of curiosities, purchased handsomely illustrated books and keys to identification, attended courses, and went on local field trips.4 Zoology shared to some extent in the tremendous popular vogue for the other branches of natural history, botany and mineralogy, but zoological collections were limited by the difficulty of gathering and preserving them. Hunting for animals was a far more complex undertaking than collecting plants or rocks, one that required greater patience and dedication and more extensive equipment. As the animals had to be killed, zoology field trips lacked the gentility of botanical and mineralogical forays. Once captured, animals either had to be preserved in bottles of alcohol or dissected, stuffed, and mounted. It was generally known in the eighteenth century how to prevent immediate putrefaction of animal tissues; however, an effective means of preventing subsequent destruction of animal skins by insects was still lacking. It was only with the discovery of arsenic as a preservative, which was known to the staff of the Museum and became generally known about 1800, that the formation of large and permanent collections of natural history was made possible.5 Zoological cabinets of the eighteenth century thus tended to be limited to corals, shells, fossils, and insects, specimens which could be readily stored and easily preserved. In eighteenth-century France, instead of a recognizable group of zoologists working on shared problems, we find instead isolated individuals writing about animals from varying viewpoints. There were popular writers on nature such as the abbe Noel-Antoine Pluche, whose Spectacle de la nature (1732), written to illustrate the pervasiveness of God's Providence, was one of the most widely read books of the century. There were those, like the polymath Rene-Antoine Ferchault de Reaumur, who focused on the curious habits of animals, particularly insects. He, too, pointed to evidence of God's design in nature. There were the philosophes, a loose group of writers of diverse backgrounds. Several

14

THE CUVIER-GEOFFROY DEBATE

of them—DeMaillet, Robinet, Maupertuis, Diderot, and La Mettrie—wrote about animals, especially generation and animal faculties, from a philosophical standpoint in order to combat the design argument. Naturalist-voyagers such as Pierre Sonnerat described new and exotic animals in their literary travel accounts written for a popular audience. Comparative anatomy, not yet linked with taxonomy, was represented by Louis-Jean-Marie Daubenton, who described the internal anatomy of animals for Buffon's Histoire naturelle, and Felix Vicq d'Azyr, who treated the various means by which functions were carried out in his volume on comparative anatomy for the Encyclopedie methodique. Finally, in a class by himself was Georges-Louis Leclerc, comte de Buffon, author of the extraordinarily popular multivolumed Histoire naturelle generale et particuliere, which began publication in 1749 and took all nature for its province.6 As a Deist and philosophe, Buffon sought theories by which the phenomena of the universe could be accounted for by natural laws, and he pioneered in introducing a true historicity into natural history. As several historians have recently suggested, Buffon epitomized a major conceptual shift from natural history (the description of nature) to the history of nature (a truly historical account of the natural world).7 Unhampered by the restrictions of scientific professionalism, he speculated freely on such matters as the formation of the globe, the history of the earth, the nature of living matter, spontaneous generation, and the possible formation of animal families by "degeneration" from an original stock. His volumes on mammals and birds, written in a lively style for a cultured audience, neglected taxonomy in favor of discoursing on animal habits, generation, faculties, and their relation to their environment. Buffon's disciple Bernard-Germain-Etienne de Lacepede, who wrote the volumes of the Histoire naturelle on reptiles and fishes, endeavored to imitate the literary style of his mentor, and earned the reputation of "successor of Buffon." More interested in taxonomy than Buffon, he nonetheless preferred his own highly artificial system of classification to that of Linnaeus.8 Compared with the nineteenth century, there was little official support for the study of zoology and comparative anatomy. Buffon, his collaborator Daubenton, and Lacepede held posts at the Jardin du Roi as intendant, keeper of the cabinet, and assistant keeper of the cabinet, respectively. Daubenton also taught mineralogy at the College de France (after 1778) and "rural economy" at the Ecole Veterinaire d'Alfort (after 1782). There were no official professorships of zoology; the nearest to a professorship of comparative anatomy was Vicq d'Azyr's appointment as a professor of the anatomy of animals at the Ecole Veterinaire d'Alfort. As this was only a part-time post, Vicq d'Azyr was also dependent for his income on medicine, in particular on his position as permanent secretary of the official Societe Royale de Medecine. Through sponsoring expeditions, the French government supported the careers of a handful of nonspecialized naturalist-voyagers. And some zoologists could supplement their income by satisfying the public appetite for handsomely illustrated works of natural history such as the volumes on zoology and comparative anatomy of the Encyclopedie methodique.9 Otherwise researchers in zoology were depen-

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dent upon positions in botany, medicine, or unrelated sciences, or upon private wealth. It was only toward the end of the century that there emerged a small group of naturalists with a common program of research based on Linnaean taxonomy.10 The members of this group were trained as botanists, but all wrote on zoological subjects as well. Their attempt to institutionalize natural history was thwarted first by the opposition of the Academie des Sciences and then by the Revolution. During his ascendancy, Buffon had resisted the spread in Paris of the use of Latin binomial names and of Linnaeus's hierarchy of classes, orders, genera, and species. The leading Parisian botanists, while adopting the Linnaean nomenclature, agreed with Buffon in rejecting the artificial Linnaean system of classification. This was a system for the identification of plants based on external characters (numbers, relative sizes, and positions of stamens and pistils) that was convenient to use but not intended to express the true relations of the plants to each other. The French botanists proposed instead to create a "natural" system of classification based on the real affinities of plants. The most successful effort was that of Antoine-Laurent de Jussieu, who founded the families in Genera plantarum (1789) on a rational "subordination of characters" according to their functional significance.11 At the Faculte de Medecine at Montpellier, however, the Linnaean system received a more favorable hearing. Antoine Gouan, professor of botany from 1767 to 1803, acquainted several generations of students with the doctrine and method of Linnaeus. About the time of Buffon's death, three of his students, Pierre-Marie-Auguste Broussonet, Jean-Guillaume Bruguiere and Guillaume-Antoine Olivier, were instrumental in bringing Linnaean taxonomy to Paris. The Societe Linneene, founded in Paris in 1787, and its successor, the Societe d'Histoire Naturelle, marked the decline of Buffon's reputation among French naturalists. At the meetings of the Societe Linneene, the members presented to each other descriptions of the external characteristics of new species in Latin written according to the rules of Linnaeus and accompanied by exact drawings. An independent organization, the Societe Linneene collapsed in 1789, its downfall brought about by the uncertainties of the Revolutionary era as well as by the jealous opposition of the official society, the Academie des Sciences. Having revived the society in 1790 under the less provocative name of the Societe d'Histoire Naturelle, the naturalists had the satisfaction of ceremoniously inaugurating a bust of "the immortal Linnaeus" in Buffon's very sanctuary, the Jardin du Roi. The organization's secretary, Aubin-Louis Millin, one of the most ardent supporters of Linnaeus in France, pronounced what was soon to become a common verdict on Buffon. Although his literary talent was deserving of praise, Buffon had nonetheless "retarded the progress of true knowledge in Natural History by the scorn he has shown and instilled toward systems and methods, without which this science can offer only confusion, and can be nothing but an inextricable labyrinth."12 In the early 1790s, naturalists began the task of giving organizational structure to French natural history, through the meetings of the Societe d'Histoire Naturelle and through the establishment of courses and journals. The members

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THE CUVIER-GEOFFROY DEBATE

of the Societe d'Histoire Naturelle instituted a series of informal courses—Millin and Alexandre Brongniart taught various branches of zoology—and published a single volume of Actes. But by 1793 the political upheaval in Paris forced the society to disband. A second attempt to found a professional journal, the Journal d'histoire naturelle, edited by Lamarck, Olivier, Bruguiere, the mineralogist Rene-Just Hauy, and the chemist Bertrand Pelletier, did not continue beyond two volumes appearing in 1792. Although the society and the journals failed, mainly because of the distractions of the Revolutionary period, they were evidence that zoology was on the verge of becoming a recognized field of study when in 1793 the Jardin du Roi of the ancien regime was suddenly transformed into the Museum National d'Histoire Naturelle. From Jardin du Roi to Museum d'Histoire Naturelle

Established in 1626 under Louis XIII, the Jardin du Roi or Jardin des Plantes was originally intended to provide an alternative source of medical education to the rigid and antiquated curriculum of the Paris Faculte de Medecine. By the end of the seventeenth century, three areas of teaching had been established: botany, chemistry, and anatomy. Oriented toward medicine, for the first century of its existence the Jardin was administered by the chief physician to the king. In the eighteenth century, however, the Jardin began to lose its ties with the medical profession and to become associated with the natural sciences. This tendency was reinforced in 1739 by the appointment of Buffon, a scientist without a medical degree, as intendant. During Buffon's long reign of fifty years the Jardin du Roi flourished as never before. Buffon doubled the land of the Jardin, reorganized and tripled the "school of botany," greatly enriched the cabinet of natural history, and sent seeds and plants to the far corners of the world. It was through the Histoire naturelle, which began publication in 1749, that zoology and comparative anatomy came to be permanently linked with the Jardin.13 When Buffon died in 1788, a crisis in the affairs of the institution was reached. The professors and other principal employees of the Jardin who had chafed under Buffon's autocratic administration turned for leadership to Daubenton, the "patriarch of natural history" of the 1790s. Brought to Paris in 1742 to be Buffon's collaborator for the Histoire naturelle, Daubenton was given in 1745 an official post as keeper and demonstrator of the Cabinet du Roi attached to the Jardin. Until he began to work with the Cabinet, it contained little more than medicinal herbs and shells, collected by Joseph Pitton de Tournefort, a former professor of botany. Daubenton made of it a research tool for the study of natural history. Minerals, fruit, woods, shells, stuffed birds and mammals, and his own anatomical preparations were collected and arranged in good order in four large halls which were open to the public twice a week.14 The transformation of the Jardin into the Museum during the Revolutionary period was initiated not by the government but by the professors and principal

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17

employees of the Jardin, led by Daubenton.15 Daubenton opposed the position of intendant, given after Buffon's death to the courtier, the marquis de la Billarderie, because he regarded the post as an unnecessary patronage award at the disposal of the crown. Instead he wished to see the Jardin administered democratically by the major employees, or "officers" as they came to be called. The Jardin of 1788 had three professorships in the same subjects that had been taught since the seventeenth century—botany, chemistry, and anatomy—and three demonstrators to act as assistants. Rene Desfontaines, professor of botany, lectured at the Jardin itself, while Antoine-Laurent de Jussieu, demonstrator of botany, taught rural botany through a series of botanical excursions in the vicinity of Paris. Antoine Portal lectured on human anatomy, while the demonstrator Antoine-Louis-Francois Mertrud gave occasional independent lecture-demonstrations of animal anatomy. Chemistry was taught by AntoineFrancois Fourcroy, with the aid of preparations made by the demonstrator Antoine-Louis Brongniart. All the professors and demonstrators of 1788 had medical degrees, with the exception of Brongniart, who had been apothecary to Louis XVI. The remainder of the "professional" staff, except for Daubenton, were not trained as physicians. They included Andre Thouin, the chief gardener; Daubenton, keeper of the cabinet; Lacepede and Barthelemy Faujas de Saint-Fond, both assistant keepers of the Cabinet; and Gerard van Spaendonck, artist. Jean-Baptiste Lamarck, a former protege of Buffon, was given a post of botanist to the king attached to the Cabinet (sometimes called keeper of the herbarium) in 1789. These twelve men were to become the professors of the new Museum. The officers were already dissatisfied with the organization of the Jardin when the coming of the Revolution brought new pressures. The Jardin was first affected when, in 1790, the Committee of Finances sought to reduce the budget and eliminate the two positions held by Lamarck and Faujas de Saint-Fond.16 In protest, the officers of the Jardin prepared an address which was presented to the National Assembly on 20 August 1790. Instead of stressing research and professional science, they carefully worded their proposal so that it would be appealing to the members of the Assembly, by dwelling on the public character of the Jardin and its enormous utility to the nation. They praised unreservedly the grand philosophical views of Buffon, popular with members of the Assembly, which the officers claimed had "so well prepared minds for the great ideas of liberty and regeneration." In conclusion, the officers requested the opportunity to draw up a proposal for a new organization of the Jardin that would at the same time suit their needs and be in keeping with revolutionary ideals. "Shall the tree of liberty," they asked, "be the only tree that cannot be naturalized at the Jardin des Plantes?"17 This proposal was to form the basis for the creation of the Museum in 1793. In response to the address, the National Assembly postponed the decision of the Committee of Finances for a month while the officers drafted their plan. In a deliberate affront to la Billarderie, the officers chose Daubenton to preside at the meetings of the officers. The resulting document, presented to the Assembly on 9 September, called for an establishment with twelve chairs, to

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THE CUVIER-GEOFFROY DEBATE

be administered democratically by an assembly of professors headed by a director elected each year and eligible for only one term. Salaries were to be made equal, rather than based on seniority. The post of intendant was deliberately left unmentioned. Recalling to mind the ideals of equality, fraternity, and liberty that would prevail in the new establishment, the officers concluded: "It is thus, Messieurs, that the Jardin des Plantes, transformed into a veritable Museum d'histoire naturelle, could soon become a sort of metropolis for all the sciences useful to agriculture, commerce, and the arts."18 A sample budget showed how all these improvements could be accomplished within the funds allotted by the Committee of Finances. Despite the attraction of the proposal, nothing was done, for both the Commitee of Finances and the first Committee of Public Instruction, created in October 1791, were too preoccupied with other matters. Only in 1793 was the proposal resurrected and acted upon. In the midst of the Reign of Terror, when all other scientific institutions of the ancien regime were being suppressed as elitist, the Jardin was not only spared, but expanded. With the demise of the Universite de Paris and other educational establishments on 18 August 1792, and with the eradication of all academies, including the Academie des Sciences, the Academie de Chirurgie, and the Societe Royale de Medecine on 8 August 1793, the Jardin too may have been in danger of dissolution, although accounts of this matter differ. The immediate events leading to the founding of the Museum dated from a visit of the politician Joseph Lakanal to the Jardin on 9 June 1793. It is not clear whether his purpose was to warn the Jardin of peril or simply to discuss arrangements for housing the confiscated cabinet of the prince of Conde.19 In any case, Daubenton took this occasion to discuss the needs of the officers and to give him a copy of the proposal of 1790, which Lakanal revised through the night and presented to the Convention on the next day. It was adopted immediately without discussion. In the turmoil following the recent removal of the Girondins from the Convention, the decree of 10 June establishing the Museum passed almost unnoticed. Just two months before the Academie des Sciences was closed, the officers of the Jardin were suddenly granted their wish that the Jardin be reorganized into a museum of natural history. It has been suggested that the Jardin des Plantes did not share the fate of its sister scientific institutions because natural history, unlike the physical sciences, was seen by the Jacobins and the populace as a science of the common man, one that did not divorce nature from moral ends.20 As a result of the decree of 10 June, the Museum was endowed with twelve chairs, which were to be filled by the twelve incumbent officers. Abolishing the post of intendant, the decree placed the Museum under the control of the officers and gave it an annual budget of 100,000 livres.21 The officers immediately went to work on a code of internal regulations, adopted by the Convention on 21 June, which stated that the twelve courses of the Museum would be taught by the twelve current officers of the establishment. They were to decide among themselves who would fill each chair. The new institution was to have three chairs of botany, two of zoology, two of chemistry, two of anatomy, and chairs of mineralogy, geology, and iconography or illustration. It was relatively easy

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to assign chairs to the three former professors and their demonstrators, but more difficult to apportion the remaining chairs. Given his choice, Daubenton selected the new chair of mineralogy. Since both the botany positions were filled, Lamarck, who owned a shell collection and who had been a close friend of Bruguiere and Olivier, both experts on invertebrates, was chosen to teach the course on "insects and worms." Lacepede, popularly known as Buffon's successor, would have been the obvious choice for professor of vertebrate zoology, but in March 1793 he had been forced, as a noble, to flee Paris for political reasons. Ostensibly on the grounds of ill health, he resigned his place as assistant keeper and demonstrator of the Cabinet. Daubenton had awarded this post, suitably scaled down, to his young protege Etienne Geoffroy SaintHilaire, who had held it for only a few months when the Jardin became the Museum. By this wholly fortuitous chain of events, Geoffroy, only twenty-one and entirely unprepared, became one of France's first official professors of zoology. Geoffroy's Early Career

Etienne Geoffroy Saint-Hilaire was born at Etampes, a small town to the south of Paris, on 15 April 1772, the seventh of fourteen children (of whom seven died very young).22 His father, Jean-Gerard Geoffroy, was a barrister who was elected to the court of Etampes at the beginning of the Revolution. The family was distantly related to three former members of the Academie des Sciences, one of them the eighteenth-century chemist Etienne-Francois Geoffroy, known for his theory of chemical affinities. Like most naturalists raised in the ancien regime, Geoffroy owed his successful start in a scientific career more to sheer luck than to training. He was to prepare for three other careers before turning to science, and even then, he planned to become a mineralogist rather than a zoologist. Originally Geoffroy was intended by his parents to enter the priesthood, an appropriate vocation for a child of frail constitution coming from a large family of modest means. When he was still quite young, his father employed his connections to have conferred upon him an advantageous benefice by the abbe de Tressan, a friend of the family. After his studies at the local College d'Etampes, Geoffroy was sent in 1788 to the College de Navarre in Paris as a scholarship student (boursier). There he received a broad philosophical education typical of clerically run schools in the ancien regime. At Navarre, Geoffroy was first attracted to the sciences through the course of physique experimentale given by MathurinJacques Brisson, a member of the Academie des Sciences who had published in natural history, particularly ornithology, before turning to physics in the 1760s. In 1790, eager to study the sciences, Geoffroy begged his parents to be allowed to remain in Paris after receiving his philosophy degree, in order that he might attend courses at the Jardin des Plantes. His father, who doubted that science was a viable career, assented only on condition that he enter as a boar-

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THE CUVIER-GEOFFROY DEBATE

der at the College du Cardinal Lemoine and prepare to become a lawyer. Law agreed with the young Geoffroy no more than had theology. He dutifully obtained a bachelor's degree in law before the end of 1790, but finding the subject arid, he again requested permission for a change of career plans. Having no intention of practicing medicine, he convinced his parents that he ought to enter medical school. They agreed because medicine was a respectable and secure profession, and Geoffroy was hence free, at least while a student of medicine, to take courses in the natural sciences. Before Geoffroy could obtain a medical degree, the Revolution intervened and closed down the old Faculte de Medecine. By the time a new medical school was established, Geoffroy had already embarked on a scientific career. At the College du Cardinal Lemoine, Geoffroy met the man who was not long afterwards instrumental in bringing him to the Museum. The abbe ReneJust Hauy, who resided at the college as a regent emeritus, was teaching Latin while continuing to pursue an active scientific career. A mild and retiring man, he had been introduced by Daubenton to the study of mineralogy, and had transformed the science by demonstrating that all crystals could be constructed from a combination of a small number of basic types. For this achievement, he was elected to the Academie des Sciences in 1783, and after the death of Daubenton, in 1800, he was to become professor of mineralogy at the Museum. Geoffroy and Hauy had a common bond, for Hauy had also been a boursier at the College de Navarre and a student of Brisson. The young Geoffroy approached Hauy and his friend, the elderly grammarian the abbe Charles-Francois Lhomond, with a naive admiration, and despite the difference in age, he became their companion, sharing in their scientific conversations and accompanying them on their promenades in the Jardin des Plantes. Enthusiastically applying himself to learn the fundamentals of Hatty's specialty, he attended Fourcroy's course in chemistry at the Jardin and Daubenton's course in mineralogy at the College de France, all the while neglecting his medical courses. Through Hauy, he was introduced to the leaders of the physical sciences in Paris. In 1792 he was attending meetings at Hatty's lodgings along with such distinguished savants as Lagrange, Lavoisier, Guyton de Morveau, and Berthollet.23 At age twenty his only ambition was to learn mineralogy. This ambition was only abandoned because of a fateful series of events that occurred on the eve of the September massacres in 1792. While Geoffroy was occupied studying the sciences, the successive Revolutionary governments were becoming increasingly radical. Regardless of his probable Revolutionary sympathies,24 Geoffroy was concerned for his teachers, most of whom were priests and hence suspect. Following the journee of 10 August 1792 and the deposition of Louis XVI, Hatty and several other priests from the College de Navarre were imprisoned at the Seminaire Saint-Firmin. Geoffroy was successful in bringing about Hatty's release by alerting Daubenton and several other savants, who then intervened with the authorities in the name of the Academie des Sciences. Isidore Geoffroy dramatically recounted in his biography of his father the spine-chilling tale of Geoffroy's attempt to rescue his

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other priestly friends from prison with the aid of a disguise and a ladder at 2 o'clock in the morning of 2 September, just before the September massacres. Exhausted by the excitement of this episode and fearful that he would be punished for aiding priests to escape, Geoffroy returned to Etampes two days after the massacres. There he fell gravely ill with what Isidore Geoffroy described as a "nervous fever" and did not return to Paris until winter. In response to Geoffroy's generous efforts on his behalf, Hauy was said to have urged his friend Daubenton to "love, aid, adopt my young liberator."25 When, in early March 1793, Lacepede found it expedient to abandon his post as keeper and subdemonstrator of the Cabinet, the opportunity arose for Daubenton to honor Hauy's request. On Hauy's recommendation, Daubenton obtained the approval of the intendant, Jacques-Henri Bernardin Saint-Pierre, and only a few days after Lacepede's resignation, had Geoffroy nominated for the post. The letters in support of Geoffroy emphasized that he was by no means intended to replace Lacepede, a naturalist of wide reputation. Instead the title of the position was deliberately reduced to the modest "subkeeper and subdemonstrator of the Cabinet," a post clearly subordinate to that of Daubenton. It was argued that as a student of Daubenton, Geoffroy was ideally suited to assist him.26 Hardly had Geoffroy installed himself at the Jardin in lodgings near those of his benefactor, when the Jardin was transformed into the Museum and Geoffroy found himself professor of zoology, a subject about which he knew next to nothing. Neither Geoffroy nor Lamarck was qualified to be a professor of zoology in 1793. They were given the posts because they were already employed by the Jardin des Plantes, but also because there was little alternative in this period of revolutionary upheaval. Several of the more accomplished French zoologists, such as Olivier, Broussonet, Bruguiere, and Lacepede, had left Paris. Nonetheless, there was considerable opposition to Geoffroy's appointment. The future mineralogist Alexandre Brongniart, only two years Geoffroy's senior, was incredulous at the sudden elevation of "the young unknown of twenty-one years" to so choice an honor. First, a position left vacant by Lacepede had been filled on the whim of Daubenton without announcement of the opening or competition. This despotic action was then compounded by giving Geoffroy a professorship that by all rights should have gone to a naturalist with experience and reputation. If a more senior savant could not be found, then Brongniart felt that he was more deserving of the prize than Geoffroy. A member of the Societe Philomatique, a society founded in 1792 for young scientists hoping to get into the Academie, and of the Societe d'Histoire Naturelle, Brongniart had already studied zoology for several years. In the spring of 1793 he had given courses on three of the Linnaean classes: "amphibious animals" at the Lycee and "insects" at the Lycee des Arts, both private institutions for popular education, and "worms" at the Societe d'Histoire Naturelle. "What is the use of having worked, and having achieved a bit of renown?" he complained in private. "This injustice is so crying, that it disgusts me more than it hurts me." Upon making inquiries, Brongniart learned that a number of the professors of the Museum—among them Fourcroy and Desfon-

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THE CUVIER-GEOFFROY DEBATE

taines—were equally angered by Daubenton's high-handedness.27 Fourcroy was said to have wanted to offer the post to the German naturalist and anatomist, Peter-Simon Pallas. But the backing of Daubenton and the politician Lakanal, together with the prejudice against importing a foreigner, weighed the balance in favor of Geoffroy.28 Geoffroy himself did not feel competent to the task. He accepted only after obtaining the reassurance of Daubenton and the approval of Lacepede, whose place he feared he was appropriating. Indeed, as soon as it was safe for Lacepede to return, after 9 Thermidor, a thirteenth chair was created for him, a chair of reptiles and fishes, leaving Geoffroy the mammals and birds. Compared with the professors of botany, Geoffroy began with meager resources: a few skeletons used by Daubenton and a pitiful collection of poorly stuffed animals. Initially only 600 francs was budgeted for expenses for both professors of zoology.29 Geoffroy's assignment to increase the collection of vertebrates was soon supplemented by that of managing a menagerie of live animals, begun in 1794. As a professor, he had to give a public course of forty lectures, for which he had less than a year to prepare. By his own account, he learned taxonomy by organizing the collections confided to his care. His first course, taught in the spring and summer of 1794, treated the natural history of man and the mammals according to the doctrines and classificatory scheme of his mentor Daubenton, and concluded with a few lectures on the birds. The twenty-two-year-old professor began his first lecture on 6 May while the Jacobins were still in power, full of faith that he was presiding over the beginning of a new era for zoology: Citizens, the place where we find ourselves gathered together, the object for which we are assembled, all prove that a new order of things is established in this monument of the sciences. . . Whereas our brothers in arms are going to repel with sinewy might the futile attempts of the coalition of kings and cement with their blood the foundations of our republic, we in the quiet of study . . . are going to acquire new knowledge in order to add new luster to the glory of the nation.30

Geoffroy's Mentors: Buffon and Daubenton

When Geoffroy began his career as professor of zoology in 1793, he was dependent most of all on the guidance of two predecessors. The first was Daubenton, to whose patronage Geoffroy was indebted. But beyond Daubenton, there was a second mentor, one whom Geoffroy had probably never met, but who nonetheless through his writings inspired Geoffroy with his all-encompassing vision of natural history. This spiritual mentor was of course Buffon. Buffon and Daubenton struck Cuvier, Goethe, and many other nineteenth-century naturalists as a study in contrasts: Buffon, proud, aristocratic, and autocratic, a bold and imaginative thinker who tended to isolate himself from the savants of his day, and Daubenton, a modest, patient man, a republican citizen, reluctant to generalize, and a friend to all.31 The Histoire naturelle and Daubenton's volume

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on mammals for the Encyclopedie methodique were no doubt Geoffroy's first texts of zoology. His beliefs concerning the role of classification in natural history derived from both Daubenton and Buffon, but his philosophical approach to natural history came largely from Buffon. Buffon, in the earlier volumes of the Histoire naturelle, affected to scorn taxonomy, especially the Linnaean system of classification, as a dry, dull, and futile exercise, for he believed it was impossible to draw boundaries between groups of organisms. Only species had a real existence in nature; genera, orders, and classes were inventions of man. At best, classificatory "methods" were conveniences, useful but arbitrary. This criticism applied not just to artificial systems but to any system based on a hierarchy of characters. Whatever characters a naturalist chose for dividing animals, his method was sure to place together animals that were dissimilar and to separate animals that belonged together. The worst classification in this regard, according to Buffon, was that of Linnaeus who relied on such artificial characters as teeth and digits to divide mammals. Moreover, Buffon saw no value whatever in Linnaean nomenclature, with its bizarre Latin and Greek vocabulary. Far better, he believed, to employ common French names. In his own volumes on mammals, Buffon chose, with a certain degree of perversity, to arrange the animals in their order of importance to man, beginning with the horse, dog, and cow, continuing with native animals, and concluding with exotic animals.32 Later Buffon adopted a more liberal view of the utility of grouping animals, which was, however, based on the very radical premise that true natural groups were based on common descent. In later volumes of the Histoire naturelle, he proposed classifications of mammals and birds into natural families, but in contrast to the abstract groups of other taxonomists, Buffon believed his familes to have a physical existence in nature. For the mature Buffon, a family consisted of those animals that had descended by way of "degeneration" from a single parent stock. Modifications in the parent stock took place through adaptations to changes in the environment. Buffon was henceforth concerned with demonstrating by any means available that the environment did have the power to modify significantly animal structure.33 Geoffroy adopted Buffon's skepticism of taxonomy, for the most part shared by Daubenton, and was intrigued by the questions raised by Buffon's theory of degeneration. Early in his career he became actively engaged in investigating the role of the environment in zoology. Geoffroy probably took his initial idea of a unity of plan in the animal kingdom from Buffon. Buffon's history of quadrupeds was full of scattered references to gradations in nature and to the single plan of organization that could be followed from man on down through the fishes. For example, in his famous article on the ass, Buffon, in discussing the controversial question of whether the ass was a degenerate horse, adduced evidence of a common plan of structure, a certain "primitive and general design that one can trace very far" in the animal kingdom. He marveled at "the singular and almost perfect resemblance" between the skeleton of a man and that of a horse. The foot of the horse, which at first glance appeared so different from the hand of a man, was

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THE CUVIER-GEOFFROY DEBATE

nonetheless composed of the very same bones. The horse's hoof was nothing but a modified fingernail: One will judge if this hidden resemblance is not more remarkable than the apparent differences, if this constant conformity and this plan [dessein] followed from man to the quadrupeds, from quadrupeds to cetaceans, from cetaceans to birds, from birds to reptiles, from reptiles to fishes, etc. . . . does not imply that in creating animals, the Supreme Being desired to employ only one idea, and at the same time vary it in all possible manners, so that man might equally admire the magnificence of the execution and the simplicity of the plan.34

We shall see that Geoffroy's 1795 paper on the makis (lemurs) began with this same presentiment of a structural unity among the vertebrates. Indeed, many of Geoffroy's general ideas concerning natural history, evident well before he announced the program of "philosophical anatomy," can be attributed to Buffon. Geoffroy shared with Buffon his distrust of final causes, his Deism, and his tendency to materialism. Although there is no reason to suspect that Geoffroy was an evolutionist early in his career, he was led by Buffon's historicism to consider seriously the role of the environment in modifying animal structure. Following Buffon, Geoffroy rejected the doctrine of preformation from the outset of his career, instead assuming that generation was an epigenetic process subject to environmental modification. Perhaps the most important part of Buffon's legacy to Geoffroy was his unwillingness to establish divisions between the sciences and to confine himself to any single science. Geoffroy, like his colleagues Lamarck and Lacepede, saw himself as a naturalist-philosopher, not simply as a vertebrate or invertebrate zoologist. Throughout his life, Buffon remained Geoffroy's ideal investigator of nature and a model worthy of emulation. Daubenton, in contrast, exemplifies the transition to the more restrained and specialized research that became the norm after the founding of the Museum. Although he shared Buffon's skepticism of taxonomy and refused to limit natural history to classification, he had no sympathy with his employer's penchant for generalization nor with his advocacy of literary style. Daubenton's was a static world in which the power of the environment was limited to the formation of varieties, and the primary task of the naturalist was to describe what existed. A physician and a childhood acquaintance of Buffon, Daubenton was deliberately sought out by Buffon to serve as a collaborator on the Histoire naturelle, complementing his own particular strengths. Having little interest in the dry details of the external and internal description of quadrupeds, Buffon delegated this task to Daubenton, who had a natural inclination for this type of careful work. As Buffon's collaborator for the volumes on mammals, Daubenton furnished a detailed description of the external structure, skeleton, and viscera of each animal (illustrated by plates) and a description accompanied by exact measurements of all relevant specimens in the Cabinet. All his descriptions were made on a common plan so that one might easily compare the anatomy of some 200 species of quadrupeds. Daubenton's anatomies thus also contributed to the notion of a unity of plan among the mammals. Despite a tendency

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to lose himself in detail, Daubenton became, with Vicq d'Azyr, one of the major revivers of the science of comparative anatomy in the generation before Cuvier. Daubenton's collaboration with Buffon lasted only through the first fifteen volumes of the Histoire naturelle. According to Cuvier, whose sympathy lay with Daubenton's careful dissections rather than with Buffon's broad insights, flatterers convinced Buffon that his work would be more attractive to the general public without the tedious additions of his assistant. One might readily understand why the precise dimensions of a horse's stomach might not appeal to Buffon's cultured audience. Buffon therefore arranged, without informing Daubenton, to publish a new edition of the Histoire naturelle in thirteen volumes, retaining his own discursive essays on the habits of animals but omitting Daubenton's anatomies and external descriptions. The later volumes of Histoire naturelle—those on birds and minerals—were written without the assistance of Daubenton, much to the detriment of these volumes as science, in Cuvier's opinion.35 Daubenton generally shared Buffon's views on classification—that all classifications were arbitrary, and that only species existed in nature—yet he was more willing than Buffon to recognize the practical necessity of taxonomy. Therefore, while he was not enamored of the Linnaean system, and would have preferred French names to Latin binomials, he nevertheless encouraged the careers of several of the Linnaean naturalists. In contrast to the Linnaean naturalists, however, he believed that recognizing and naming species formed only a small part of the subject matter of natural history, which also encompassed the study of the internal and external structure of organisms, their habits, their useful and harmful properties, and all the changes that they underwent from birth to death.36 Daubenton judged some classifications to be more natural than others because they took more affinities into account, but in the end he believed that no simple scheme of classification could fully capture the multiple affinities found in nature. Particularly reprehensible because of its gross distortion of animal relationships was the doctrine of the chain of being, according to which all animals could be placed in a single sequence. As early as 1782, Daubenton took the leadership in refuting this celebrated and attractive theory, popularized by the Swiss naturalist Charles Bonnet. Although Daubenton believed with Buffon that there were no gaps in nature, he denied that all animals could be arranged linearly. Instead he asserted that affinities between animals followed a network of radiating lines. In 1795, in his lectures at the Ecole Normale, he declared that in his study of the mammals, he "discovered no continuous sequence from one species of quadruped to the others, no direct order, no progression of characters that might indicate the place that each species ought to occupy relative to the others in order to be arranged along the same line. But I have seen clearly several lines joined to each other at different points."37 These same notions of imperceptible differences and multiple affinities appeared, as we shall see, in Geoffroy's first published paper. Geoffroy's first paper may in one sense be regarded as an application of Dau-

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benton's ideas of classification, but in another sense, it signaled a potential conflict with Daubenton's strictures on the proper role of the naturalist. The Museum and other new scientific institutions created in the 1790s gave impetus to the trend, already well underway in the physical sciences, toward a more specialized research with results directed to other scientists rather than to the educated public. As the "patriarch of natural history" of the 1790s, Daubenton took a leading role in the replacement of the nature writer of the eighteenth century by the professional naturalist of the nineteenth. For many years, Daubenton had chafed under Buffon's domination, unappreciated by the majority of the readers of the Histoire naturelle. Once in a position of authority, however, he enjoined both naturalists and the public to reject the Buffonian concept of natural history, in favor of a more mundane and narrowly conceived view. Especially revealing of the trend toward specialization were Daubenton's lectures given at the short-lived Ecole Normale in 1795. His immediate audience consisted of prospective elementary school teachers in the provinces, but his lectures in published form reached a wide public. In his introductory lecture, "On the definition and the limits of natural history," Daubenton defined natural history as a science which ought be distinguished from all other sciences, especially from chemistry, anatomy, veterinary medicine, and agricultural science. While these fields were evidently related to natural history, they should never be confounded with it. Daubenton emphasized that it was difficult to learn all the principles and details of a single science, and impossible for an individual to acquire an in-depth knowledge of all the sciences. "Embracing several sciences at once is an act of indiscreet curiosity," he warned his audience, "a sort of libertinage of the mind, which leads one to be unable to study them thoroughly or teach them well." One must therefore recognize the limits of the various sciences and become a specialist in order to make an effective contribution to the progress of knowledge.38 In other lectures of the course, Daubenton attacked facile generalizations of eighteenth-century science, as exemplified by Bonnet's chain of being, and the use of literary forms, typified by Buffon. Theories, like methods of classification, Daubenton held to be inevitably faulty. Some might serve as conveniences, provided they were based on facts and accepted provisionally rather than as dogma, but others, such as the chain of being, had no place in natural history, for they were simply untrue. Although Daubenton admitted that theories might have their practical advantages in terms of summing up knowledge and encouraging research, he viewed literary style in uncompromising terms as a detriment to the progress of natural history because it obscured truth and led the naturalist into "exaggerated expressions which produce false ideas." Readers of literary works of natural history might at first find themselves attracted to "pompous style," but when they examined the matter more carefully, they would find that nature had been "decked out and disguised." In Daubenton's opinion, the true style of the naturalist should be "as simple as nature."39 Daubenton thus viewed the naturalist not as a wide-ranging philosopher of

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nature, but rather as an expert who had mastered a carefully delimited segment of human knowledge, and who endeavored to communicate that branch of knowledge to others in as clear and unadorned a manner as possible. The expertise of the naturalist could be gained not from books alone, nor from traveling to exotic lands, but only through long hours spent in the quiet of a methodically arranged cabinet of specimens. Daubenton had a sobering influence on the impressionable Geoffroy, but he was only partially successful in countering the attractions of Buffon. Geoffroy's Debut

Even on the occasion of his first memoir, read on 1 December 1794 before the recently revived Societe d'Histoire Naturelle, the conflict between Geoffroy's generalizing proclivities and his need for professional recognition was evident. The paper was to deal with a limited subject, the creation of a new genus of quadrupeds to accommodate the unusual mammal, the aye-aye of Madagascar, first described by the voyager Sonnerat. In the original version of the memoir, Geoffroy insisted on beginning with a long discourse on the doctrine of the chain of being and on the philosophy of classification. Taking the part of the "indefatigable observer" against that of the superficial philosopher who was carried away with "hazardous conjectures," Geoffroy dismissed the "universal chain" as enticing but nevertheless "a true chimera." Although he believed an ideal system of classification to be impossible, Geoffroy did not think that naturalists had therefore to resort to arbitrary but convenient classifications based on a single character. The naturalist could still hope to construct a practical system of classification using as many characters as necessary to preserve as many natural affinities as possible. Not until page nine of the manuscript did the aye-aye of Madagascar, the ostensible subject of the paper, make its appearance. Although Daubenton no doubt agreed with Geoffroy's generalities, which were essentially derivative of his own work, he appears to have thought his young protege's rambling philosophical introduction to be out of place in a first memoir, and urged him to omit it from the published paper. Geoffroy complied, having decided upon rereading the pages in question that they were "a bit too declamatory."40 The published memoir, which appeared in the scientific and literary journal La decade philosophique, dived immediately into the problem at hand, that of classifying the aye-aye, which is now considered a primate but which Linnaeus had placed among the rodents (Glires) and the German naturalist Johann Georg Gmelin had placed in the genus of squirrels. Geoffroy showed that the conformation of the animal was sufficiently "bizarre" to merit a genus to itself. Accordingly, he christened it, after his mentor, Daubentonia madagascariensis, the name it still bears today. Although continuing to classify the aye-aye in the order of rodents, he admitted that it did not fit any of the orders of mammals established by Daubenton, for the aye-aye had multiple affinities with the monkeys, the makis, the didelphs (marsupials), and the rodents.41

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Such multiple affinities were not at all disturbing to Geoffroy, for they only confirmed what he already knew from Buffon and Daubenton: that no system of classification could be perfect. Any attempt to place animals in categories, he held, was sure to emphasize some natural affinities at the expense of neglecting others. Even the categories themselves could not always be clearly circumscribed. For, while Geoffroy rejected Bonnet's chain of being, he continued to believe, with both his mentors, that nature made no leaps but proceeded by nearly imperceptible steps. From the beginning of his career, then, Geoffroy appeared to have renounced the ideal of the one true natural system of classification. In a second preface, this one published, Geoffroy presented in an undeveloped form the doctrine which eventually became the focal point of his debate with Cuvier. In the opening remarks of a taxonomic memoir on the makis (lemurs) written at the end of 1795, he put forward the proposition that one could trace, at least within each class of animals, a common plan of organization. "It seems that nature has enclosed herself within certain limits," he wrote, "and has formed all living beings on only one unique plan, essentially the same in its principle, but which she has varied in a thousand ways in all its accessory parts." As examples of hidden resemblances, Geoffroy cited the osseous pouch of the howler monkey, which was an enlargement of the base of the hyoid bone; the pouch of didelphs (marsupials), which was a fold of the skin; the trunk of the elephant, which derived from a prolongation of the nostrils; and the horn of the rhinoceros, which was a mass of hair adhering together. Thus, he concluded: The forms in each class of animals, however varied, all result in the end from organs common to all. Nature refuses to employ new ones. Thus, all the most essential differences which affect each family within the same class come only from another arrangement, from another complication, in short, from a modification of these same organs.42

Clarifying the web of affinities of these little-known primates presented a challenging problem for Geoffroy. He broke up the single genus Lemur of Linnaeus into five genera: Maki Lemur, Indri (a kind of tailless lemur), Loris, Galago (the bushbabies), and Tarsius (the tarsiers of the East Indies). The "flying lemur" or galeopithecus of the East Indies, he showed, was not a lemur at all. As in his paper on the aye-aye, Geoffroy found evidence to support his conviction that there were no gaps in nature. The indris formed a genus intermediary between the monkeys and the makis, while the galago, which he was first to name and describe in this paper, was a "mixed species" connecting the loris and the tarsiers. "It seems that nature has formed it so that there would not be a sudden leap [saut brusque] in her work, and in order to arrive at the extraordinary form of the tarsier, until then unique in all creation, by an insensible gradation."43 Such speculations on the unity of plan among the mammals and on the links between separate groups of animals were Buffonian in character, and Buffon was their obvious source. But Buffon had not undertaken the research neces-

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sary to establish specific homologies of parts. Unity of plan was rather a vague insight which found expression from time to time in the course of his rambling discourses on mammals and birds. Likewise, Geoffroy's first pronouncements on unity of plan were of the nature of pleasant introductory remarks and did not yet constitute a program of research. The examples that Geoffroy cited in the preamble to his paper on the makis were common knowledge and bore on organs as a whole rather than skeletal pieces. Geoffroy had yet to establish any new homologies, and he had yet to single out the skeletal system as he would in his later works. How far unity of plan extended was still vague in his mind. It seemed to him evident mainly at the level of class, in this case the mammals. Geoffroy's ideas underwent considerable development between 1795 and 1806, as these first speculations were transformed into a systematic attempt to establish "unity of composition" among the four classes of vertebrates. Cuvier and Geoffroy: Colleagues and Collaborators

Geoffroy had scarcely begun his career as a zoologist when, in late 1794, he was placed in contact with Cuvier, then a young naturalist employed as a tutor in Normandy. A friend of Geoffroy's family, the abbe Henri-Alexandre Tessier, who "discovered" Cuvier in Normandy, wrote in enthusiastic terms to Geoffroy on his behalf. When, at Geoffroy's request, Cuvier forwarded samples of his work, the new professor was easily impressed. Given Geoffroy's poor training in zoology, it is not to be wondered that he saw in Cuvier, though only three years his senior, a fund of helpful expertise. Despite the warnings of colleagues, who feared he would create a rival for himself, Geoffroy undertook to bring the young man to Paris and establish him as a colleague at the Museum. Georges Cuvier was born on 23 August 1769 into a Protestant family of modest circumstances in Montbeliard, a French-speaking region in the Duchy of Wurttemberg, which became part of France in 1793.44 A precocious child, Cuvier was brought to the notice of the duke of Wurttemberg, who had founded the Karlsschule, a school at Stuttgart to prepare young men for ducal service. At this militarized German academy Cuvier spent the years from 1784 to 1788, obtaining a solid, career-oriented education quite different from the philosophical and classical training of Geoffroy. He enrolled in the faculty of administration, which included the study of law, finance, agriculture, and technology. Years later, he claimed that he chose this program because it was there that one could study natural history.45 Cuvier began his scientific career as primarily a Linnaean naturalist, but one with a more philosophical outlook. The Karlsschule's professor of natural history, Johann Simon von Kerner, an accomplished botanist, presented him with a tenth edition of Linnaeus's Systema naturae, which served as Cuvier's main guide in his early researches. There was no formal training in anatomy, but Carl Friedrich Kielmeyer, a fellow student and one of the early Naturphilosophen, taught him to dissect, and also, Cuvier recalled, gave him his "first

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ideas of philosophical natural history."46 The precocious young student began a herbarium, sketched more than a thousand insects, and founded with Christoph Heinrich Pfaff, Johann von Autenrieth, E.G.F. Hartmann, and others a natural history society which met once a week to read and discuss papers. The authors of the best memoirs were decorated with an order for which Cuvier furnished the design: a picture of Linnaeus surrounded by representatives of the three kingdoms of nature.47 Cuvier thus from the beginning saw taxonomy as the chief function of the naturalist. Although an excellent student, Cuvier was unable to obtain a position in the ducal service when he left the Karlsschule in 1788. In order to provide a living for himself, he took a position as tutor for the son of a wealthy Protestant family named d'Hericy, who were living in France. From the relative safety of Normandy he watched the Revolution unfold, describing his reactions to these events in a series of letters to Pfaff, his friend and fellow student at the Karlsschule. Although well-disposed to the initial phases of the Revolution, Cuvier soon became disheartened and disgusted by senseless violence committed in the name of liberty. His belief that ideas were at the root of the Revolution led him to fear the consequences of unbridled generalizations in the sciences, as well as in political and social thought. Even as early as his letters to Pfaff, Cuvier had little sympathy for "systems" or hypotheses in natural history.48 Cuvier's letters to his friend reveal him as an exceptionally gifted young man, who at the age of nineteen had already read most of the classics of natural history. As soon as new works were published, he obtained copies and read them, commenting upon them to his German correspondent. Not only did he become familiar with the writings of Brisson, Olivier, Lacepede, and A.-L. de Jussieu, but he also read Lavoisier and other chemists and physicists. In diaries which he kept for botany and zoology, he entered Latin species descriptions in the manner of Linnaeus. Already, however, he saw the need to broaden natural history to something more than Linnaean taxonomy. Following Daubenton's methods of comparative anatomy, he dissected birds, fishes, and invertebrates, sketching parts and noting measurements. One is astonished at the seriousness with which the young tutor pursued his avocation; in far-off Normandy, he became as experienced a naturalist as anyone in Paris. Cuvier had already begun to contemplate turning his hobby into a Paris career and had tried making contacts with Paris naturalists to that end, when an opportunity was opened for him by a chance encounter with the abbe Henri-Alexandre Tessier, an agronomist and member of the Academie des Sciences who was hiding from the Terror under an assumed name in Normandy. Recognized by Cuvier at a meeting of the local Society of Agriculture, Tessier was so surprised by the young unknown's talent that he sent glowing letters to Jussieu and to Geoffroy. To Jussieu he was said to have written on 10 February 1795: "At the sight of this young man, I experienced the ecstasy of that philosopher who, thrown on an unknown shore, sees traced there figures of geometry."49 Cuvier forwarded to Geoffroy memoirs on the anatomy of the octopus and the snail, illustrated by sketches, and some reflections on the classification of quadrupeds, and sent other work to Millin, secretary of the Societe d'His-

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toire Naturelle. Millin had Cuvier named a member of the society and offered him a minor post on the temporary commission of arts, but it was Geoffroy who took most of the initiative in bringing Cuvier to Paris. In prophetic terms, Geoffroy claimed to have written, "Come to Paris. Come play among us the role of another legislator of natural history."50 Enthusiastic and generous by nature as well as politically naive, Geoffroy was eager to have Cuvier come to Paris immediately and work with him. There was a possibility that a position would be available for Cuvier. Mertrud, the demonstrator of anatomy in the ancien regime Jardin des Plantes, had been named professor of the anatomy of animals after its reorganization. A man of whom almost nothing is known, wholly untrained in the systematic study of comparative anatomy, Mertrud was incapable of teaching the proposed course. By a peculiar and controversial feature of the French educational system, professors at institutions such as the Museum who could not or would not teach did not have to resign their posts. Instead they might sublet the teaching to a suppleant for a portion of their salary, while still maintaining the privileges and status of titulary professor. With Vicq d'Azyr dead, no one else in Paris was trained to teach comparative anatomy, except for Claude Richard, named professor of medical natural history at the newly founded Ecole de Medecine in 1795, who antagonized others by his quarrelsome personality. Cuvier suggested that it was to avoid bringing Richard to the Museum that he as an outsider was given consideration. Hoping to convince Mertrud to offer him the suppleance of his chair, he arrived in Paris with his pupil early in 1795.51 Geoffroy's colleagues were distinctly less inclined to promote the careers of outsiders. Only Lamarck showed any positive reaction at all, and that was slight. Cuvier was unknown; all he had published were two notes on Crustacea and a memoir on the anatomy of the limpet (Patella), a mollusk, in the shortlived Journal d'histoire naturelle. Educated in a foreign country, he had only recently become a French citizen. Moreover, he, with his superior knowledge, might become a rival to Geoffroy. Isidore Geoffroy recounted that one day Daubenton invited his young colleague to dinner and placed before him a copy of Lafontaine's Fables, opened to the poem "The Bitch and Her Friend." The fable of the bitch giving in to the successive importunities of her companion until her home was taken over concluded, "Yield but a foot to their appeal/ And find they have usurped four feet."52 According to Cuvier, Hauy, who had influence over Geoffroy, warned him that Cuvier would take all the credit for their joint work. Cuvier, well aware of the resistance to his establishment in Paris, recalled that Geoffroy stood firm in his friendship: He [Hauy] tried to make him [Geoffroy] believe that in associating with me, I [alone] would have all the glory of our work and he engaged him not to favor me in any way; but this excellent young man, after bearing for a week the unhappiness that this advice caused him, freely confided in me, and assured me that his conduct with respect to me would never change.53

Heedless to the protective arguments of his advisors, Geoffroy assisted Cuvier in his rapid rise to prominence. On 20 March 1795 and more definitively on 2 January 1796, Cuvier was named professor at the newly created

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secondary school, the Ecole Centrale du Pantheon. Soon Geoffroy and Cuvier were on intimate terms; they worked together and dined together, and planned to collaborate on an extended work in mammalogy. When the position at the Museum was secured in July 1795 and the pupil sent home to Normandy, Cuvier moved into Geoffroy's lodgings at the Museum. They lived together for some months until Cuvier was given rooms of his own in the apartment of Mertrud.54 The melodramatic quality of the two years of collaboration has been sensed by all of Geoffroy's biographers: the most celebrated of scientific rivals began as the closest of friends. To what extent there was true intimacy or a community of ideas even at the beginning of their collaboration is open to question. One wonders if the intimacy might not have been a product of Geoffroy's wishful thinking. Despite their later scientific falling out, Geoffroy always counted it among his greatest achievements to have brought Cuvier to Paris, and looked back upon their youthful friendship with a romantic nostalgia.55 As the two young men embarked upon a new science, they had much in common. In a scientific community dominated by the physical scientists, they both were acutely aware of the need to improve the image of their field. Chemistry in France had recently become systematized through the work of Lavoisier, while physics had undergone rapid development through the mathematical astronomy of Laplace and the application of mathematics to electricity by Coulomb, Poisson, and others. Even mineralogy had been endowed with a theoretical character by the work of Hauy. Natural history, by comparison, seemed a puerile exercise in collecting and describing. As early as the letters to Pfaff, Cuvier had recognized the importance of placing natural history on a more philosophical basis than Linnaeus's Systema naturae.56 Together the two young naturalists wrote five memoirs, dealing with the classification of mammals, the two-horned (African) rhinoceros, the species of elephants, the natural affinities of the tarsier, and the natural history of orangutans. In these papers, they were concerned not merely with the details of classification and description, but with the establishment of principles.57 Cuvier's earliest comments on classification, as revealed in his letters to Pfaff and in his first two papers, published in 1792, show that he did not at that time differ greatly from Geoffroy. In 1790 Cuvier still agreed with Daubenton and many other naturalists that all taxonomic divisions above the level of the species "are simple abstractions of man."58 Cuvier had already rejected the possibility of arranging all species on a linear scale, but, like Geoffroy, he continued in the 1790s to believe in continuity in the animal kingdom. He wrote of passages between families, and of the "degeneration" of the nervous system as one passed from man to the zoophyte. In his first published paper in 1792 on woodlice (terrestrial Crustacea), for example, he traced a gradual chain from woodlice to crayfish, and concluded with the familiar adage, "Here, as elsewhere, nature makes no leaps."59 Once in Paris, however, Cuvier began to modify his early view of taxonomy. Even in 1795, one can perceive the beginning of a divergence in the approaches taken by Geoffroy and his new collaborator. The theoretical basis of their most famous memoir written together, "Memoir on a New Division

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of Mammals and on the Principles Which Ought to Serve as a Foundation in This Type of Study," has always been attributed to Cuvier, and it reflects his growing conviction that a classificatory scheme could be something more than a mere convenience. Cuvier's reading of Jussieu's Genera plantarum (1789) and his own research in comparative anatomy gradually convinced him that a natural system in zoology was possible. One might create natural genera, orders, and classes, Cuvier came to believe, if the choice of characters employed to delimit groups was reformed according to rational principles. Animals would be classified not according to a few external characters chosen arbitrarily, but according to their true affinities, which would be revealed to zoologists by comparative anatomy. Cuvier's innovation, which initiated a major reform in natural history and led to the modern delineation of zoological classes, was the introduction of comparative anatomy into zoology.60 While the actual reform of taxonomy may have depended more on empirical technique than on Cuvier's rational principles of classification, these principles were central to Cuvier's program for natural history. His first task was to determine criteria for forming natural groups—criteria based on functional considerations. In the memoir on mammals presented with Geoffroy to the Societe d'Histoire Naturelle in April 1795, shortly after his arrival in Paris, Cuvier applied the principle of "subordination of characters," introduced by Jussieu into botany, in order to establish a natural classification of Geoffroy's bailiwick, the mammals. This principle, which implicitly assumed the later-enunciated principle of correlation of parts, stated that the primary divisions in taxonomy should be those functions that exercised the greatest influence over the life of the animal, while secondary divisions ought to derive from subordinate functions. Reasoning that generation, respiration, and circulation were, in that order, the three primary functions, he divided mammals into fourteen natural orders, on the basis of characters associated with these functions.61 Cuvier's most important early paper, read on 5 May 1795, a month after his first paper with Geoffroy, continued the application of subordination of characters to the subject of his own previous researches, the "white-blooded animals." Relying in large part on his own anatomical preparations, since invertebrates had previously not been systematically dissected, Cuvier established, by comparison of internal organs, six classes of invertebrates—mollusks, crustacea, insects, worms, echinoderms, and zoophytes—in place of Linnaeus's two classes of "insects" and "worms."62 While the most significant of the joint papers primarily reflected Cuvier's concerns, Geoffroy's input in these papers was also visible. These collaborative efforts contained several provocative, speculative passages on continuity in the animal kingdom. In the tarsier paper, for example, the authors indicated that "this genus can be considered as the link which unites the quadrumana to the chiroptera or bats."63 And in the orangutan paper, they offered in passing the following remark, which has appeared to some historians as favorable to evolution: But, it is a rather general law of nature that the number of species in a genus is approximately proportional to the fecundity of each of them, whether what we

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call species are only the various degenerations of a single type, . . . or whether many of them have arisen from the coupling of neighboring species.64

Such remarks, probably originating from Geoffroy (by way of Buffon) were not taken further, and in no way can be interpreted as "evolutionary." We can be sure, however, that through his reading of Buffon, Geoffroy was already attuned to the problems of distinguishing species from varieties, and of determining the extent to which modification of the environment could cause a species to vary. In a letter written in English in 1796 to Charles Willson Peale, proprietor of a natural history museum in Philadelphia, Geoffroy requested Peale to send him, among other things, animals, such as rats, mice, and shrews, that had been transported from Europe to America. He then mused: "How many analogous forms are taken for Species. It would be interesting to know what degeneration their transplantation has produced on their economy. They would lead us to a more exact knowledge on the nature of the species and even of the species in general."65 It would be a mistake to conclude from passages such as these, as at least one historian has done, that Geoffroy was an evolutionist from the beginning of his career or that Cuvier actually at one time supported the chain of being or entertained the notion of evolution of the species.66 Whether or not the two naturalists agreed on all points of doctrine, it is safe to say that the lines of their future division were not yet clearly drawn. Geoffroy was still interested in pursuing the natural system (although he envisioned it in practical rather than ideal terms). Establishing the rational bases for a classificatory system was an intriguing problem for Geoffroy, since the principles of classification bore on the nature of "animality," a topic of high philosophical import. Cuvier, for his part, was at this point in his career still feeling his way. He was not yet opposed to noting links between separated groups of animals, nor had he as yet fully developed his later aversion to speculation. It is only in retrospect that one can locate the seeds of Cuvier and Geoffroy's later divergence. Cuvier, Geoffroy, and the Museum Environment

Cuvier and Geoffroy, as professors at the Museum, found themselves in an environment that was unique in the world.67 Their future work cannot be understood without some reference to this institutional setting and how it functioned. For it was the resources of the Museum that enabled Cuvier and Geoffroy to do the research and writing on which the debate was based. Almost all of Geoffroy's papers, as well as his masterwork of philosophical anatomy, Philosophie anatomique, were dependent on the collections of the Museum. One could not conceive of Cuvier's Leqons d'anatomie comparee, his Recherches sur les ossemens fossiles, and especially his Le regne animal without the facilities offered to him as the Museum's professor of comparative anatomy. First and not least among these resources was an assured income. Neither

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Cuvier nor Geoffroy was independently wealthy. Both could rely for their sole support upon their income as scientists. Professors at the Museum earned an annual salary of 5000 francs. This was not a sufficient income to live in high style, but because of the way in which the French system of institutions functioned, many of the professors were able to increase it by additional posts. Geoffroy was able to make do with his Museum income alone, but Cuvier, as we shall see, supplemented his salary as lecturer at the Museum with funds from several other public sources. In addition to salary, professors were provided with housing on the Museum premises, and most professors took advantage of the offer. Both Cuvier and Geoffroy resided on the Museum grounds for the remainder of their lives. Thus was created the possibility of a very close-knit group of researchers living together amidst all the facilities necessary for research. The reality was somewhat different, as the story of the Cuvier-Geoffroy debate makes clear, for the Museum community was rent by personal and family rivalries.68 The duties of the professor were not unduly onerous, and they provided ample time for research or other activities. Each professor taught a single course of forty lectures a year in the subject of his chair. As in the Jardin du Roi, the courses were free and open to the public; no degrees were conferred, but the professors signed certificates of attendance. In addition to teaching and sharing in the administration of the Museum, each professor was in charge of the Museum's collections in his area, and it was his task to augment them, arrange them methodically, and publish catalogues. Geoffroy was in charge of the galleries of mammals and birds, while Cuvier maintained the Museum's collection of comparative anatomy. The collections grew with extraordinary rapidity. During the Revolution, the cabinets of emigres as well as cabinets confiscated from the lands invaded by France were added to the collections. Particularly valuable was the cabinet of the Stadholder of Holland, obtained in 1795, which supplied material for several of Cuvier and Geoffroy's early papers.69 From an original four rooms in the ancien regime, the Cabinet of the former Jardin du Roi soon spilled over into several buildings. In the Restoration period, the old Cabinet building, which had been enlarged several times, housed tools and agricultural machinery, the library, fossils, minerals, and the galleries of zoology. A second building, acquired by the Museum in 1795, contained the galleries of botany and the zoological laboratory where skins were prepared, stuffed, and mounted. In a single year (1805), we are told, some 100 quadrupeds, 500 birds, and 500 reptiles and fishes were made ready for display. The collections of comparative anatomy, first opened to the public in 1806, were kept in yet another building that was enlarged and attached directly onto Cuvier's lodging in 1817. The edifice also contained the anatomical laboratory, under Cuvier's control, in which the work of dissecting and preparing specimens took place. Aide-naturalistes and preparateurs were hired to assist the professors and carry on the day-to-day tasks of dissecting, mounting, preserving, and recording.70 It was not long before the Museum became the largest zoological repository in the world. When its first official history was written in 1822, the Cabinet

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contained 1500 mammals belonging to over 500 species, 6000 birds representing over 2000 species, 1800 reptiles belonging to over 700 species, 5000 fishes from over 2000 species, 25,000 species of arthropods (insects, spiders, crustacea), and an unspecified number of mollusks and zoophytes. Cuvier could expound the text of Le regne animal through the very halls of the Museum, as the vertebrates were arranged according to his classificatory scheme.71 Cuvier's Cabinet of Comparative Anatomy in 1822 contained a total of 11,486 preparations, 6231 of them dried and 5255 preserved in alcohol. There were human skeletons and skulls of different ages and races, 1500 skeletons and 1041 skulls of vertebrates, 300 crania showing the bones separated, several series of individual bones comparing the same bone in different animals, 870 preparations of teeth (considered essential for the classification of mammals), as well as collections of dried and injected intestines, feathers, nails, scales, branchial arches of fishes, sterna of birds (important for classification), and hyoid bones. Preserved in alcohol were 172 preparations of muscles, 216 brains, 327 eyes, 220 hearts, 915 preparations of all the viscera of various animals, 80 fetuses in their envelopes, 881 dissected mollusks, and 1097 dissections of other invertebrates. Access to materials such as these soon became essential to do acceptable research in natural history.72 Besides the collections, the Museum possessed several other facilities for research. First of all, there was the library, provided for in the decree establishing the Museum. Open to the public, by 1822 it contained 15,000 volumes. A second research facility was the menagerie of live animals begun in 1794 by Geoffroy. It was eagerly visited by the public, no doubt contributing to popular support for the Museum, and it piqued the imagination of many a popular writer. But the animals of the Museum served a larger function than the education and amusement of the public. Since little was then known about the habits of exotic animals, the residents of the menagerie served as valuable research tools. Both Geoffroy and his assistant, Cuvier's younger brother Frederic, wrote technical research papers based on their study of living animals at the Museum. Together they published a four volume illustrated folio work (1824-1842), intended to appeal to a wide cultured audience, but nonetheless containing original observations on animal behavior.73 The inhabitants of the menagerie provided the raw material for Frederic Cuvier's significant and controversial research on the nature of animal instinct and intelligence. In death the animals continued to serve the needs of research as subjects for dissection in the laboratory of comparative anatomy and for mounting in the laboratory of zoology. A third and especially important facility was the house journal of the Museum, the Annales, first published in 1802. Before that time there were no permanent journals in France for publishing research in natural history. The Memoires of the Institut, which in any case had little space for long memoirs on taxonomy, were at least two years behind schedule, while the Bulletin of the Societe Philomatique published only in extract form. Most of Cuvier's and Geoffroy's early papers appeared in La decade philosophique, litteraire et politique or the Magasin encydopedique, ou journal des sciences, des lettres et des

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arts, edited by the Linnaean naturalist Millin. These were general periodicals which contained allegories in prose and verse, politics, history, archeology, and theatrical and literary reviews, in addition to all branches of science, medicine, and agriculture. Their readers might be expected to appreciate Geoffroy's discussions of unusual new animals such as the aye-aye or the tarsier, or Cuvier's general revision of invertebrate classification, but they could hardly be expected to wade through Geoffroy's routine classifications of the species composing the order of primates or Cuvier's detailed anatomies of mollusks. Thus the founding of the Museum Annales was a major event, for it gave the professors of the Museum a fast outlet for technical research. As the Annales was not dependent on subscribers for turning a profit, it could afford to publish articles of interest to only a very select audience. After 1802 Cuvier and Geoffroy published most of their work in the Annales and its successors, which were the major journals for zoology in France until the founding of the Annales des sciences naturelles in 1824.74 The Museum, defended during the most radical phase of the Revolution for its utility to the public, soon became the center of an elitist and technical science. Naturalists from all over the world came to attend the courses, work with the collections, and envy the professors who were supported by government largesse to live, teach, and do research in the midst of such unparalleled resources. With its secure professorships, its immense collections, its zoo, its ample library, and its house journal, the Museum encouraged specialization and professionalism. Daubenton's and later Cuvier's proselytizing also contributed to this same end. Increasingly the professors directed their writings not to the educated public, but primarily to other naturalists and physicians. Instead of writing for general-interest periodicals, zoologists and comparative anatomists began to publish most of their work in specialized scientific and medical journals. Although the public retained throughout this period an active interest in the affairs of science, it was no longer accorded the right, as it had in the eighteenth century, to judge scientific works. Even those more popular journals which reviewed scientific books and scientific meetings for the public hired scientists and physicians to write the reviews. Not all the professors of the Museum were willing to make the adjustment to the specialized research that came to be expected of them. This was especially true of Buffon's two former disciples, Lamarck and Lacepede. Although Lamarck pleased his colleagues by the publication of a seven-volume Histoire naturelle des animaux sans vertebres (1815-1822), which laid the basis for the classification of invertebrates, he was never content to limit himself to taxonomy. Instead, he saw himself as a "naturalist-philosopher" in the Buffonian mold. In the 1790s he alienated many academicians by writing attacks on Lavoisier's chemistry and speculative works on meteorology and hydrogeology. By the time he published his major statement on evolution, Philosophie zoologique, in 1809, he had almost entirely lost effective contact with his colleagues.75 Instead of marshaling evidence to support his thesis, he wrote in a speculative and deductive manner, seemingly oblivious to the methodological qualms of his peers. Hardly anyone in France took his grandiose theories seri-

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ously. As Henri de Blainville, one of the few naturalists who openly admired him, recalled, Lamarck had become so wrapped up in his own thought that it was no longer possible to have a rational exchange of ideas with him.76 Lacepede taught his course at the Museum only long enough to complete the volumes of the Histoire naturelle that he had promised to Buffon. For these works, especially the Histoire naturelle des poissons (1798-1803), he composed literary discourses in which he gave full reign to speculation on evolution and other subjects. Then, in 1804, he let Constant Dumeril teach his course, and practically dropped out of the scientific community. The remaining twenty years of his life he devoted to the type of unspecialized writing he truly enjoyed: a natural history of man, romantic novels, and a history of civilization.77 Geoffroy, only beginning his career when he became professor at the Museum, was able to adapt better to the new surroundings, but all his life he retained a vision of science that was shaped by his eighteenth-century training and his admiration of Buffon. Never content with description, and skeptical of creating a natural system of classification, Geoffroy did not write the systematic treatises of mammalogy and ornithology that might have been expected of a man in his position. He lacked the patience for writing textbooks that might have served as a guide to others. What attracted him from the start was the enterprise of reducing nature to universal laws primarily through the use of reason rather than by the painstaking collection of data. This vision brought him into sporadic conflict with his colleagues in the course of his career, and led in his old age to an increasing alienation from them. Cuvier, on the other hand, flourished in the new environment. From a modest beginning, his rise to fame was nothing short of meteoric. Not only was he possessed of unusual scientific abilities, he also had an exceptional capacity to seek out scientists of importance and gain their support. He cultivated PierreSimon Laplace, leader of the physical scientists, whom he met at the Ecole Normale in 1795; he frequented the weekly meetings of the Societe Philomatique, where he came in close contact with Brongniart and the mathematician Sylvestre Francois Lacroix, and he quickly presented impressive memoirs at the Societe d'Histoire Naturelle. His most important paper of 1795 had aimed at no less than a revision of the general classification of the invertebrates. In 1796 he began his famous series of paleontological papers destined to prove to the scientific world the reality of extinction. Geoffroy had as yet published very little on his own. "It was through working with me on some memoirs on animals," Cuvier recalled, "that he truly began to make himself known."78 When in December 1795 the ancien regime Academie des Sciences was recreated in a new form as the First Class of the Institut de France, Cuvier was so far advanced in his career that he was chosen one of the six original members of the new section of anatomy and zoology, whereas Geoffroy, already established in Paris as professor of zoology for two years before Cuvier arrived on the scene, was unsuccessful. At 26, Cuvier was the youngest member of the Institut, but he was not at all surprised by his good fortune. "I found myself well enough known that everyone supposed in advance that I would be named,

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and that in fact happened on 17 December 1795," he recalled in his memoirs.79 According to Cuvier, it was supposed at the time that Geoffroy would have been the sixth member of the section, but the place instead went to Richard. The botanists did not want him and insisted that he be selected by the zoologists. "It was with true chagrin," he wrote, whether sincerely or not, "that I saw myself thus pass before Geoffroy who had been the principal author of my advancement."80 Geoffroy did not enter the Academic until twelve years later, in 1807. Cuvier's quick success was in part due to his philosophical treatment of natural history and the utilization of comparative anatomy in zoology.81 In part it was due to his ability to identify patrons and to publicize his talents. However, the disruption caused by the Revolution must also be taken into account. Of the few workers in zoology that France possessed in the period before the Revolution, many died, left Paris, or turned to other pursuits. In Germany, anatomy and zoology were flourishing. Cuvier, abreast of the work of Kielmeyer, Jean Herman, Autenrieth, Pallas, and Johann Friedrich Blumenbach, had a tremendous advantage over his early competitors, who, partly because of the Revolution, were largely ignorant of German work.82 Having arrived at a propitious moment, within a few years Cuvier became the most noted naturalist in France. The colleagues who warned Geoffroy were indeed prescient, for Geoffroy was quickly eclipsed by the brilliant newcomer. In the end, it was Cuvier rather than Geoffroy who carried out Daubenton's mission of making zoology a "French science." Through his celebrated writings as well as through the multiple resources of patronage, Cuvier was able to establish a conceptual framework for future research in zoology and to gather disciples around him. Thus he occupied throughout his career a very different role from Geoffroy in the scientific community. The emergence of philosophical anatomy took place in the context of this framework for natural history erected by Cuvier and promoted through the exploitation of the resources at his command. To understand the conflict of ideas culminating in the Cuvier-Geoffroy debate of 1830, and the response of contemporaries to the arguments placed before them, it is necessary not only to examine Cuvier's functionalist anatomy and the intellectual assumptions behind it, but also Cuvier's predominant position in French science. Thus, before considering the further development of Geoffrey's morphological thought, it is to Cuvier, the "legislator of science," that we now turn.

Chapter 3

"Le Legislateur de la Science": Cuvier and Functionalist Anatomy

Known to all as Cuvier's chief scientific rival, Geoffroy nonetheless never ceased to speak of Cuvier as "the greatest authority in the natural sciences." At Cuvier's funeral in 1832, Geoffroy counted it as one of his greatest achievements "to have been the first to recognize and reveal to the learned world the scope of a genius who did not yet know it himself." "Throughout the land, each of those who cultivate the science of nature," Geoffroy declared, "owes especially to Cuvier what he knows and what he is in natural history. All have been molded by the inspired genius and immense knowledge of our great zoologist."1 Geoffroy was quite sincere in his sentiments. Whether or not they agreed with his scientific doctrine, Cuvier's contemporaries were in nearly unanimous accord that Cuvier was the greatest naturalist of the age. Georges Louis Duvernoy, disciple and author of a biography, acclaimed him as "the legislator of science, he who regulated all its progress, he to whom it owes its finest conquests." Pierre Flourens, disciple and successor to many of Cuvier's posts, lauded him in his eulogy before the Academie as "one of those men whose name alone would suffice to the glory of a nation and a century."2 Cuvier was indeed more than the dean of naturalists: he was a spokesman for the scientific enterprise, a much-honored public citizen, and a popular idol who had unveiled the past ages of the earth through his almost magical ability to reconstruct fossils. Nearly every newspaper and journal of the day published the speeches made at Cuvier's funeral. The title frequently accorded Cuvier—the legislator or arbiter of natural history—reflects the enormous influence Cuvier exerted on natural history in France and elsewhere throughout much of the nineteenth century. In France, his authority was felt in two ways, first through his courses and books, which erected a basic structure for natural history, and second through his pivotal position in the institutions of French science. Cuvier's work was of such breadth and stature as to define a program for natural history which other scientists could develop by filling in the details. His researches in paleontology, taxonomy, and comparative anatomy specified the questions that naturalists ought to ask as well as the methods to be used in seeking answers. More than 40

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any other individual, Cuvier set the standards for research and publication in zoology and comparative anatomy. The synthesis of paleontology, taxonomy, and comparative anatomy that Cuvier achieved was based on a teleological approach to nature, one that gave primacy to functional purpose over structural affinity. Underlying all Cuvier's work, as William Coleman and Henri Daudin have shown, was a commitment to the functional integrity of the organism.3 His Lecons d'anatomie comparee, organized by functional systems, described the various means whereby each function was carried out in the animal kingdom. His paleontological reconstructions were based on a related functional principle, that of correlation of parts. Likewise his natural system of classification was built on a hierarchy of characters determined by subordination of functions. The fundamental principle in Cuvier's biology was "the conditions of existence," which, he readily granted, was another term for final causes. Cuvier implicitly assumed that the Creator, having considered "the conditions of existence"—the conditions necessary for the animal to survive and reproduce in a given environment—had created just those organs which were needed. There was no place in Cuvier's thinking for useless organs or for organs created to maintain serial relationships, symmetry, or unity of plan. In fact, his 1812 division of animals into four absolutely distinct embranchements denied the possibility of any meaningful structural affinities across embranchements, and even within embranchements such structural unity as existed was subordinate to "the conditions of existence" or functional necessity. While not stated in explicitly religious terms, Cuvier's functionalism nonetheless gave support to the traditional argument from design. It is not surprising that Cuvier's works were enthusiastically received in Britain, where works written in the framework of natural theology were commonplace. Cuvier advanced his doctrine on methodological as well as on scientific grounds. Although his work was far from empirical, Cuvier adopted and promulgated an empiricist ideology in order to combat opposing points of view. In his writings from about 1800 on, he resolutely maintained that natural history ought to be restricted to "the positive facts." His campaign against "systems," especially those of the eighteenth-century philosophes, was in keeping with the emerging professionalism of French science, which was already much advanced in the physical sciences, and reinforced Cuvier's authority among the leaders of these sciences. Through his strategic positions in the French scientific community, and later on the Council of the University and the Council of State, Cuvier was able to promote his ideas with an unsurpassed effectiveness. Already admitted to the ranks of the leaders in French science by his election to the Academie des Sciences in 1795, Cuvier's star continued to rise. When Daubenton died in 1800, Cuvier was chosen to fill his chair of natural history at the College de France, an institution founded in the seventeenth century to provide free public lectures on a variety of humanistic and scientific subjects. This gave Cuvier a prestigious podium from which to reach a broad audience on a virtually unrestricted choice of scientific topics. In 1802, at Mertrud's death, Cuvier assumed

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official title to the chair of animal anatomy, which he renamed the chair of comparative anatomy. Finally, in 1803, when the regulations of the Institut were revised, Cuvier became one of the two permanent secretaries of the First Class of the Institut. This position, perhaps the most coveted in the sciences, gave Cuvier unparalleled opportunity to pass judgment on his predecessors and contemporaries and to legislate the progress of natural history through his reports on others' work, his annual analyses of the work of the Institut, and his eloges of deceased members. Thus to understand Cuvier's opposition to Geoffroy and the philosophical anatomists, we need to examine not only Cuvier's biological theories, but also his general views on the nature of science and his position in the French scientific community, for, as will become clear, Cuvier was motivated by a combination of science, religion, and politics. As Cuvier's scientific doctrines have been amply treated by Coleman, Daudin, and others, we need only to sketch them here. This chapter will instead concentrate on other aspects of his career that have only recently attracted the attention of historians, namely his pronouncements on the proper method to do science, and the forms of patronage by which he exercised so preponderant an authority over other naturalists.4 Cuvier's framework for zoology, we shall see, on the one hand provided fruitful stimulus to his numerous disciples in France and abroad, but on the other hand proved a multidimensional obstacle for those who found themselves in disagreement with it. Functionalism and the Natural System of Classification

Before 1820 Cuvier had written three classic works which together served to establish a new basis for the study of natural history in France. None of them could have been written without the extensive collections and facilities afforded by the Museum. These multivolumed texts—Lecons d'anatomie comparee (1800-1805), Recherches sur les ossemens fossiles (1812), and Le regne animal (1817)—went through numerous French editions in the nineteenth century, not to mention translations into a host of foreign languages.5 Contemporaries were well aware of the epoch-making character of each of these books. They defined the problems of natural history and served as models on which to base future research.6 Lecons d'anatomie comparee, based on Cuvier's courses at the Museum, reawakened interest in comparative anatomy, for it was the first systematic work to compare the basic functional apparatuses in all groups of animals, including the invertebrates.7 The first volume, for example, treated the organs of locomotion, that is, the extremities of mammals, birds, and reptiles and the fins of fishes, as well as the varied means of locomotion in cephalopods, gastropods, acephalate mollusks, Crustacea, insect larvae, adult insects, worms, and zoophytes. In the remaining four volumes, Cuvier covered in a similar manner the organs of sensation, digestion, circulation, respiration, voice, generation, and excretion. The work was aimed at zoologists as well as at anatomists, physiologists, and medical students, for Cuvier insisted that zoology henceforth be based on a detailed knowledge of the internal parts of animals.

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Central to the Lecons was Cuvier's belief in the mutual dependency of organs in the animal economy. Only certain organs could coexist, and the "laws" that regulated which organs could be found together were, Cuvier claimed, as definite as any laws in metaphysics or mathematics. It is evident, he wrote, that the harmony among organs in a given animal "is a necessary condition of existence of the being to which they belong, and that if one of its functions was modified in a manner incompatible with the modifications of the others, that being could not exist."8 Likewise, Recherches sur les ossemens fossiles served as a base for all subsequent work in vertebrate paleontology.9 Fossil vertebrate bones had been unearthed in the eighteenth century and earlier, but there had been no general agreement as to whether these bones came from elephants and rhinoceroses once living far north of their current habitat, or whether they were the remains of "lost species." From his first paleontological memoir on living and fossil species of elephants, read to the Institut in 1796, Cuvier took a stand. On the basis of fragments of only two individuals, he concluded that the mammoth of Siberia and the mastodon of North America (then also called "mammoth") were of the same genus as modern elephants, but belonged to different species. That is to say, the mammoth and mastodon were once-living species that had perished.10 With the resources at the Museum, and with the extensive fossil deposits in the quarries of Montmartre, Cuvier was able to reconstruct scores of extinct species of elephants, hippopotamuses, rhinoceroses, armadillos, deer, and cattle. In 1812 he republished all his paleontological memoirs to date in the four-volume Recherches sur les ossemens fossiles, preceded by his famous and widely read "Discours preliminaire" and by a stratigraphical study of the Paris region made by Cuvier and his friend Alexandre Brongniart. He claimed to have classified the remains of seventy-eight quadrupeds, both viviparous and oviparous, of which at least forty-nine represented species unknown to zoologists, while twenty-seven of the forty-nine belonged to seven hitherto unknown genera.11 It was Cuvier, more than any other single individual, who proved to the world that extinction was a fact of nature—that many animals living in past eras had since become "lost species." Only someone with a thorough knowledge of comparative anatomy could have succeeded at so difficult a task. Cuvier's near-magical unveiling of past ages in the history of the earth had a momentous impact upon his contemporaries and earned for him the status of a popular hero.12 Paleontological reconstruction, Cuvier claimed, was dependent on the same notion of functional integrity as informed comparative anatomy. "Every organized being forms a whole, a unique and closed system, whose parts mutually correspond and concur to the same definite action by a reciprocal reaction. None of its parts can change without the others also changing; and consequently each of them taken separately, indicates and determines all the others."13 Cuvier himself propagated the legend that he could recreate the entire animal from a fragment of a single bone.14 With Cuvier's work, paleontology became a part of natural history. Recherches was intimately linked to Lecons and to Le regne animal, for both a detailed knowledge of comparative anatomy

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and extensive familiarity with the natural system of classification were necessary to reconstruct a fossil species and place it within the system of classification. His paleontological and stratigraphical research led Cuvier to propose his controversial theory of "revolutions of the globe," or catastrophes by which all the forms of animal life in entire geographical areas were wiped out by sudden deluges and replaced by other forms of life, either created anew or migrating in from elsewhere. However hypothetical such catastrophes may appear today, to Cuvier they appeared based on solid fact, for how else might one explain the clear demarcations separating the various strata, the overturning of strata, and the specter of the Siberian mammoth found frozen in the ice with food still in its mouth?15 He rejected the notion that extinction could be accounted for by assuming that fossil species were the ancestors of living species. Although he noted a progression in fossils from the earliest to the latest rocks, which seemed to indicate the radical hypothesis that God had renewed His creation in different periods of history, Cuvier backed away from a clear commitment to multiple creations. Leaving his options open on so controversial and potentially troublesome an issue, he suggested that the fossil record might also be accounted for by a complex series of migrations.16 The task of the paleontologist, as Cuvier defined it, was not to meditate upon such murky questions, but to reconstruct and describe fossils, carefully identifying the strata in which they were found, and to classify these fossil species within the framework of the natural system. The keystone of Cuvier's work as a naturalist was his natural system of classification, displayed in the monumental Le regne animal.17 Not since Linnaeus had the entire animal kingdom been classified in one systematic work. From very early in his career, Cuvier boldly took for himself the task of reforming on the basis of new principles the entire classification of the animal kingdom. The result would be the creation for the first time of a truly natural system of animals, one that reflected the real affinities of animals to one another. In his paper on mammals with Geoffroy and in his 1795 classification of invertebrates, he applied the principle of subordination of characters as used by the botanists, to construct rational bases for a hierarchy of groups within groups.18 In Tableau elementaire de l'histoire naturelle des animaux (1798), a text published for students in the ecoles centrales, and again in Lecons d'anatomie comparee, Cuvier offered a natural classification of the entire animal kingdom.19 In 1800 his classes were still based on characters drawn from respiration and circulation. By 1812, however, he came to view the nervous system as the fundamental system of the animal economy, and the key to establishing the major divisions among animals. The four embranchements, which formed the basis of zoological classification throughout most of the nineteenth century, were introduced in a pathbreaking memoir of 1812.20 Previously naturalists had placed the four traditional classes of vertebrates (mammals, birds, reptiles, and fishes) on the same taxonomic level as the major groups of invertebrates (mollusks, Crustacea, etc.). Ever since Aristotle, a bipartite division had been recognized between

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animals with backbones and those without backbones. The latter had been christened "invertebrates" by Lamarck. Cuvier's brilliant innovation, followed ever since, was to reject the primary division of animals into vertebrates and invertebrates, and instead consider the vertebrates as a whole to be the equivalent of each of the basic groups of invertebrates. Based on the general form of the nervous system, Cuvier distinguished four general plans of organization— the vertebrate, mollusk, articulate (insects, Crustacea, etc.), and radiate (starfish, jellyfish, etc.)—according to which all animals were formed. Subsequent divisions of the "four principal forms" were "only rather slight modifications founded on the development or the addition of some parts, but which change nothing of the essence of the plan."21 Because of such statements Cuvier is often interpreted as someone who believed in abstract morphological types. His use of such expressions as "plans of organization" has led to much confusion and has muddied the distinction between his views and those of Geoffroy. In the midnineteenth century, after "philosophical anatomy" was accepted as legitimate, disciples of Cuvier sometimes attributed the principles of philosophical anatomy to Cuvier himself. However, Cuvier's four embranchements were not the abstract types of the pure morphologist. Cuvier did not even use the term "type." It is significant that "type," as a taxonomic category, was instead introduced by Blainville, who was deeply influenced by philosophical anatomy.22 For Cuvier, animals shared similar basic plans only because they carried out a similar combination of interrelated functions. In effect, the Creator freely chose to form all animals on the basis of four distinct plans of nervous system. The unity within an embranchement came not from a comprehensive unity of plan, but from a common arrangement of the nervous system, functionally the most important system of the animal. The forms of the other major systems remained constant throughout an embranchement because the other systems— respiration, circulation, etc.—were functionally subordinate to the nervous system and determined by the requirements of the nervous system. Animals within an embranchement could vary almost arbitrarily in their accessory parts, precisely because accessory parts were not necessitated by the choice of a nervous system. Because the fundamental plan was completely different in each embranchement, there were and could be no transitional forms leading from one embranchement to the next. In describing the mollusks in 1812, for example, Cuvier emphasized, "There is no nuance between this second embranchement and the first, nor any resemblance in the general disposition of parts."23 The embranchements were absolutely distinct from one another because the functional requirements of the animals in each embranchement were radically different. Cuvier believed, and his disciples agreed, that he had accomplished far more than a revision of previous classifications; he had, according to Flourens, established "the true method in natural history."24 It was through this method that natural history deserved to be respected as a mature science. The method as elaborated by Cuvier had the status of absolute applicability, and the result it aimed at was a, or rather the, unique natural system of classification. As Cuvier

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insisted, "There can be only one perfect method, which is the natural method." The natural method alone permitted generalizations in natural history, and therefore "this unique natural method . . . ought to form the goal of all efforts of naturalists."25 In essence, Cuvier wished to subsume under the natural method the whole of zoology. When the natural system is fully articulated, he proclaimed, we will have "an exact and complete expression of all of nature. . . . In a word, the natural method would be the entire science, and each step that one takes toward it will help the science approach its goal."26 Cuvier's system of thought entailed certain metaphysical commitments which were of great importance in his future battle with Geoffroy. Because of the environment in which he worked, Cuvier expressed these commitments in the more secular terms of Aristotelian teleology rather than in the language of natural theology.27 Like Aristotle and the British natural theologians, Cuvier stressed the functional integrity of the organism. Although he avoided such terms as "natural theology," "design," or "contrivance," he did employ the Aristotelian "final cause." His primary principle of natural history, "the conditions of existence"—which, as he admitted, were "commonly called final causes"—stated that since no animal could exist without the conditions which rendered its existence possible, the parts of an animal were necessarily correlated to assure internal harmony as well as harmony with its environment.28 Thus, to take Cuvier's standard example, the carnivore, if it were to survive, required sharp teeth and claws to catch its prey, a stomach and intestines suitable for digesting flesh, etc. If one then discovered by the form of a tooth that an animal was carnivorous, one could then, Cuvier claimed, deduce all the other parts. One could in fact calculate which parts belonged together, "just as the equation of a curve determines all its properties."29 In paleontology, a corollary of the conditions of existence, known as the principle of correlation of parts, enabled one to recreate the entire animal from a single bone. Although he rarely mentioned the Creator directly, Cuvier frequently employed teleological reasoning in his writings, deducing structures from needs in order to account for why animals had the properties they did. To give an example, Cuvier would reason that because mammals and birds needed more energy than fishes, they were provided with a pulmonary circulation. Because birds required a great amount of movement to sustain themselves in air, their lungs were cellular and provided with large appendages. For Cuvier, therefore, functional requirements were always primary, and structure secondary.30 Empiricism and the Battle Against Speculation

The natural system was not simply a convenient means of ordering animals, but the embodiment of correct methodology in the biological sciences. As a self-proclaimed standard-bearer of the life sciences, Cuvier had much to say about norms of scientific discourse. For both scientific and tactical purposes, Cuvier advocated an analytic and empirical methodology that contrasted

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sharply with the synthetic methodology favored by Buffon and Lamarck and admired by Geoffroy. What Lavoisier had accomplished for chemistry by attention to methodology, Cuvier's natural method supposedly achieved for natural history. Since the latter part of the eighteenth century, the physical scientists at the Academie had been conducting a campaign against "systems" in the sciences. Instead they put forward as a model the analytic method of Etienne Condillac, to which the great success of Lavoisier's chemistry was attributed.31 This method consisted of breaking a problem into components, reforming nomenclature so that the name clearly signified and classified the thing, weighing and measuring, and employing, where possible, mathematics. The natural method of classification was Cuvier's application of the analytic method, so prized by the physical sciences, to the natural sciences. If, and only if, animals were arranged according to "the method," could the naturalist hope to group facts into laws: If the subdivisions have not been established arbitrarily, but made to depend on the true fundamental relationships, on the essential resemblances of beings, the method is the most sure means of reducing the properties of these beings to general rules, expressing them in the fewest terms, and engraving them easily in the memory.32

Such "laws," Cuvier claimed, were comparable to the laws of chemistry and physics. Comparison in the natural sciences, Cuvier contended, took the place of experiment and calculation in the physical sciences. It was impossible, he thought, to analyze the properties of an animal through experiments and calculation, because the researcher could not isolate single phenomena. Instead the naturalist was forced to deal with the entire organism at once with all its properties inextricably linked. To separate one part from the whole was to bring that part into the realm of dead substances and to change completely its essence. The method of comparison provided therefore a means of compensating for the inability to experiment, for nature presented in the various organisms almost all possible combinations of organs.33 Indeed, comparative anatomy was often justified to medical students as a means of studying physiology. If one examined all the organs throughout the animal kingdom which performed a certain function, one would be able to reduce that function to its essentials. Although he praised the experimental research of Laplace on combustion and respiration and encouraged the physiological experiments of Flourens and Francois Magendie, Cuvier continued to view both natural history and physiology as basically sciences of observation. Although his own work, in retrospect, can be seen as highly theoretical, Cuvier affected an exaggerated empiricism in his public pronouncements. In his 1808 report to the Emperor on the progress of science, he insisted that Our natural sciences are only the facts brought together, our theories only formulas which embrace a great number of them [facts]. And by a necessary consequence, the smallest well-observed fact ought to be gathered, if it is new, since it can modify our best accredited theories.34

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Generalizations were essentially comparisons and classifications of observed facts. One of Cuvier's favorite expressions was "the positive facts." Those research projects which added to the store of "positive fact" were deemed valuable, while other research—which he labeled metaphysical or hypothetical— was at best judged to be useless, if not downright detrimental to the progress of science. Cuvier's "positive facts" had little to do with the positivism of Auguste Comte, and little in common with modern positivism.35 His was rather a Baconian faith that if facts were gathered and classified, generalizations would somehow be self-evident. Despite the obvious metaphysical assumptions of Cuvier's functionalist zoology, Cuvier resolutely maintained that his "principles" of the conditions of existence and of correlation stemmed from an empirical methodology. It may never have occurred to Cuvier that his support of final causes and his insistence on empiricism should be in contradiction.36 In his role as watchdog of the natural sciences, Cuvier continually admonished naturalists to refrain from speculation and to direct their efforts instead to collecting more facts. Time and again he would argue the fruitlessness of formulating broad hypotheses or of setting up a priori categories and then finding facts to fit them. "Indeed," he warned in his report to the Emperor, most of those who are devoted to these speculative researches, ignoring the positive facts, have arrived at results that are so far from the truth, that they lead one to suspect their method of demonstration to be quite at fault.37

Through much of his career, he waged a battle against a host of theories which he condemned as speculative and metaphysical, and hence unfit subjects for research: spontaneous generation, epigenesis, the chain of being, evolution, the nature of subtle fluids in the body, phrenology, etc. His tactics for combating these theories are important to note, for they reappeared in the controversy with Geoffrey. Through his powerful positions in French science, Cuvier had an unrivaled opportunity to pronounce judgment upon contemporary and historical scientific theories. Openly and shamelessly he exploited the media of the eloge, the annual analyse of the work done at the Academie, and his courses in history of science at the College de France to present object lessons on the right and wrong ways to approach science. One might suppose that an eloge of a recently deceased member of the Academie ought to provide the occasion for a flattering eulogy concentrating on the man's scientific accomplishments, but for Cuvier an eloge was a golden opportunity to moralize. He confessed in the Advertissement to his collected eloges that perhaps the principal utility of this genre of writing was "to mark the false routes on which so many superior men have not passed up [the temptation] to engage themselves, seduced by their imagination or by the desire to gather too promptly the applause of the multitude."38 "False routes" held a singular fascination for Cuvier; he seems to have read with perverse pleasure every speculative theory he could find. He never tired of discoursing on the extremes to which reckless thinking might lead. In the

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Recherches, for example, Cuvier recounted with not a little exaggeration the entire history of speculative theories of the earth. The hypotheses of Burnet, Woodward, Leibniz, De Maillet, and Buffon were each in turn made to appear absurd by his combination of witty style, deliberate oversimplification, and juxtaposition of conflicting claims. All of this served as a prelude to one of Cuvier's rare references to Lamarck (although his name was mentioned only in a footnote), who was soundly ridiculed by association with these earlier discredited theories: In our day, minds more free than ever have also wished to exert themselves on this grand subject. Some writers have reproduced and prodigiously extended the ideas of Demaillet. They say that all was fluid in the beginning; that the fluid will engender very simple animals at first, such as the monads or other infusorian and microscopic species; that in the course of time and in taking on diverse habits, the races of these animals will become complicated and diversified to the point where we see them today.39

Eighteenth-century naturalist-philosophers, especially Bonnet and Buffon, were chided for having left aside the patient investigation of nature for the approval of the public, despite the public's ignorance of true science. In his 1800 eloge of Daubenton, Cuvier took Buffon to task for having spurned taxonomy and for having terminated his collaboration with Daubenton to assure greater appeal for the Histoire naturelle. The work of Buffon, attractive to "litterateurs" and found on the "toilette des femmes," was deftly contrasted to the superior work of Daubenton, of interest to only a small circle of naturalists. Buffon, in Cuvier's estimation, "seemed to want to divine the truth, and not observe it," while Daubenton "affirmed only that which he saw and touched; [and] far from wishing to persuade by means other than the evidence itself, he discarded with care from his writings all image, all expression likely to seduce."40 The Swiss naturalist Charles Bonnet, the chief proponent of the great chain of being, was likewise the target of a biting eloge. After admiring Bonnet's experimental work on plant nutrition for its "severe logic" and the "solidity of its results," Cuvier proceeded to reproach him for turning to works of "speculative philosophy" when his eyes became too weak for detailed observations. These works, on which Cuvier dwelt at length, were characterized by "physical hypotheses. . . by which he seemed to want to offer seizable objects to the mind even when the senses refused to present them." Bonnet's fatal error was his rejection of taxonomy. If he had applied himself to the detailed study of species, Cuvier moralized, he would never have fallen into the illusion of the great chain of being. But, like so many men of his century, Bonnet scorned classification, not recognizing "that in natural history, it is the necessary basis of all other research."41 On several of the great controversies of the eighteenth century, Cuvier tried to close debate by declaring the issue outside the realm of positive science. For example, in the age-old battle between epigenesis and preformation, Cuvier sided with the preformationists, but he avoided all discussion of preformation in his works. Rather, he argued that the subject of generation, "the most pro-

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found mystery of the natural sciences," ought to be considered outside the domain of scientific inquiry. Germs were only to be found in nature, already formed and already possessing that "mouvement de tourbillon" that constituted life. It was impossible for the savant to go back to the first origin of living bodies. As far as one could see, "life is only born from life." Although the impossibility of spontaneous generation could never be absolutely demonstrated, speculations on epigenesis and spontaneous generation (which he tended to link together) were bound to be unproductive. It would be foolishly bold, Cuvier warned, for anyone to attempt to deduce from a single principle both the original formation of a germ and the phenomena it manifested. How could one possibly explain the perfect correlation of parts in the developing embryo by means of a formative principle or plastic virtue? The only remaining alternative was preformation, which, while admittedly incomprehensible, was yet necessary.42 In the vitalist-mechanist debate, much discussed in the eighteenth century, Cuvier was a vitalist who refrained from elaborating on the nature of the vital force, since it could not be observed. Although he often spoke of functions as "animal machines" and praised the work of Laplace on animal heat, regarded by historians as a major example of mechanist biology, he nonetheless disavowed reductionism. Like the physiologist Xavier Bichat, he believed that life consisted of a tourbillon, a vital movement, which opposed itself to the forces of physics and chemistry. When the organism died, physical and chemical forces gained command and dissolved the organization of the body, but while the body was still alive, the "vital force" counteracted "the laws which govern brute bodies."43 Cuvier had little to say regarding the properties of the vital force. Its existence was a necessity, since the movements of animals could not be accounted for by mechanics alone. Its operation, however, was a mystery, one that could not and should not be penetrated. Of all speculative theories, the one that most aroused Cuvier's passions was the eighteenth-century doctrine of the chain of being. It became in effect his bete noire. Although he had rejected the chain of being as early as his letters to Pfaff, in the 1790s Cuvier still believed that "nature makes no leaps."44 In his earliest classifications of the animal kingdom, he continued to arrange classes in serial order. However, from 1800 he began to reject all attempts to find graded nuances, and to insist on gaps in nature. This change of heart may be connected with the appearance in the same year of Lamarck's theory of evolution, first presented in his Museum course. Perhaps Cuvier came to see that the assumption of continuity in nature's productions left the way open to evolutionary speculation. Although Lamarck explicitly rejected the chain, by which he meant the possibility of arranging all beings along a single linear scale, the chain of being played a central role in his evolutionary theorizing. Lamarck accounted for the hierarchy of organisms by the interaction of two factors, the first being an inherent tendency toward increasing perfection of organization. If only this tendency were at work, animals might indeed be arranged along a linear scale of increasing complexity. Anomalies in the scale were the result of the second

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factor, the use and disuse of organs. Modifications of the environment, creating new needs, led to new habits, and ultimately, through use and disuse, to a modified structure. Thus, as the historian Daudin long ago noted, Lamarck's evolution can be regarded as a means of saving the chain of being by explaining its apparent deviations.45 Cuvier's main stated objection to the chain was that it was a speculative a priori scheme that went beyond the facts. In combating the doctrine, he deliberately interpreted it in a more literal and analytic manner than any of its supporters would have done. If one could not actually arrange all species in a line, then the chain of being was without foundation. In 1800 he argued that although one could trace the degradation of a single organ, especially within a class, the different organs did not follow the same order of degradation, and thus there would have to be nearly as many different series as organs. Between animals with different arrangements of the principal organs, "there is no longer resemblance in anything, and one cannot mistake the most marked interval or leap."46 By 1812 Cuvier had already renounced even the possibility of arranging classes along a scale of perfection. For example, among the articulates, the insects might be said to be higher because they have greater movement, but the other three classes (arachnids, Crustacea, and annelids) surpassed the insects by their circulation and respiration. Indeed, one could not even definitely place the reptiles between birds and fishes, rather than between mammals and fishes. Cuvier thus maintained that in both cases "there are . . . four classes which belong to the same embranchement, but without forming a series, or enjoying an incontestable rank."47 In Le regne animal he asserted, "The pretended scale of beings is only an erroneous application of partial observations to the totality of creation . . . and this application has, in my opinion, harmed to a degree that can scarcely be imagined the progress of natural history in our day."48 Not only did the chain of being seem to nullify the work of taxonomists, it also conflicted with Cuvier's teleological functionalism. Animals might vary in insignificant properties, he contended, but once a major organ was modified, all the organs must be modified so that the animal maintained its integrity. It would be impossible to pass from one combination of organs to another by insensible gradations, because the intermediaries would simply not be able to function as animals. This same argument of the nonviability of intermediate forms was used by Cuvier to combat evolution. Cuvier publicly rejected evolution well before Lamarck announced his theory in 1800. In his first paleontological paper, delivered in 1796, he explicitly denied the possibility that the anatomical differences between the Indian and African elephants might be owing to climate. To believe otherwise, he argued, would be tantamount to proposing that "all quadrupeds could derive from a single species; that the differences they manifest are only successive degenerations. In a word, this would reduce all natural history to nothing, since its object would consist only of variable forms and fugitive types."49 Given Cuvier's historical role as the main defender of zoological orthodoxy in the early nineteenth century,50 it may at first seem surprising that he rarely

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discussed evolution, and never presented a serious and detailed refutation of Lamarck's work, as Charles Lyell was to do in England. Instead, he preferred to combat Lamarck by ignoring him: Philosophie zoologique rated not even a mention in Cuvier's annual reports on scientific work of the Institut. The few times Cuvier referred to Lamarck, he did so by innuendo only. Cuvier exaggerated Lamarck's examples, oversimplified his theory, and disingenuously conflated him with discredited evolutionists of the past. The combination of public silence and private ridicule, Lamarck was well aware, destroyed his work while leaving him no means of defense.51 Only at the end of his life did Cuvier attack Lamarck directly, in his eloge of Lamarck, and then the attack was indirectly aimed at Geoffroy, Blainville, and the German Naturphilosophenofthe 1820s. Though Cuvier failed to give Lamarck the consideration he merited, he did offer several empirical arguments against evolution, which, in the context of available evidence, were more persuasive than Lamarck's speculative theorizing. If species had been transformed into one another, Cuvier reasoned, we should expect to find intermediate forms among living animals. Instead we find clearly delimited groups. Paleontology revealed no intermediate forms linking ancient and modern animals, as would be predicted if ancient animals had been the ancestors of modern animals. Furthermore, such links could not exist, for the organisms would lack the harmony of parts necessary to survive. No one had ever observed the transformation of one species into another. The 3000-year-old animal mummies brought back from Egypt by Geoffroy in 1802 proved that species had remained unchanged over lengthy periods of time. Variation in nature, as far as could be seen, was strictly limited. Climate and domestication were powerless to modify an organism beyond fixed limits, and if the organism were returned to its original environment, it would soon regain its original state. Even the species that had exhibited the greatest variability— the domestic dog—had not been modified in any essential point. Arguments such as these were strong enough to combat effectively all theories of evolution before that of Charles Darwin in 1859.52 Cuvier thus constituted himself a public authority on the proper way to advance the life sciences and at the same time retain the support of political and religious authorities. He freely legislated for other naturalists in this matter with the assurance that he knew what was best for them. As we shall see, religious and social concerns as well as strictly scientific and professional ones motivated him in his battle against eighteenth-century natural philosophy and nineteenth-century speculative theorizing. In a politically volatile country which had recently experienced traumatic revolution, Cuvier justly feared that speculative theories, most of which had a materialist tinge, would be exploited in the name of science to undermine religion and promote social unrest. If science could be limited to experts and restricted to accumulating "positive facts," then it might achieve a measure of autonomy, while at the same time the questioning that might lead to heretical theories would be eliminated.53 As Cuvier became increasingly concerned about the danger posed by certain biological theories, he became increasingly insistent on the restraints imposed by

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proper scientific method. Cuvier's methodological strategy thus became one of the major issues of dissension in the Cuvier-Geoffroy debate. Personality, Politics, and Religion

Cuvier's battle against speculative theories was in part motivated by his convictions about the nature of science and the proper place of natural history among the sciences. But Cuvier was an intensely political man, and his scientific battles were always fought with a view not only to promoting the best science but also to defending deeply held political and religious views. He thus did not hesitate to use political tools and political power in his scientific conflicts, and no scientist in France was better equipped to do so. Cuvier's three interrelated careers—as a scientist, educator, and administrator—gave him control over considerable patronage. His career in education began in 1802 when Napoleon appointed him one of the six inspectors general of public instruction, and in this capacity he went on a mission to organize lycees (secondary schools) in Nice, Marseilles, and Bordeaux. When he was chosen permanent secretary of the First Class of the Institut, Cuvier resigned his post as inspector, but in 1808 he was called to serve on the Council of the newly formed University of France. The seat on the Council, which he held for the remainder of his life, gave him immense power in obtaining jobs for his supporters in the University hierarchy. In large part he was responsible for choosing the first professors of the Faculte des Sciences at Paris.54 As an administrator, Cuvier served on the Council of State, an executive and quasi-judicial body, from 1813 until his death, with the exception of the Hundred Days (Napoleon's brief return to power after his first exile).55 Cuvier was able to remain on the Council through the Empire, three kings, and several ministries because he held no extreme opinions and was willing to support whatever regime was in power. He served on the Committee of the Interior (the committee charged with public education) and in 1827 became director of non-Catholic religions, also under the supervision of the Committee of the Interior. Few men of his day received so many official honors. Each new regime added to his renown. Louis XVIII made him a baron in 1820, and Louis-Philippe made him a peer of France in 1831. Even the august Academie Francaise opened its doors to him. His detractors claimed that no one man should hold so many posts, while his supporters replied that he filled each position more than competently. In fact, however, administrative functions did intrude on scientific functions, and after 1814 he rarely taught his courses at the Museum and the College de France.56 Politically Cuvier was a constitutional monarchist who worked for various enlightened reforms, yet because of his distrust of the masses, he found himself at times allied with conservative interests. Order was the guiding principle of his thought in politics as in biology. Above all he feared a return of the social chaos that marked the Terror.57 The Revolution had been a sobering experience for him, and the lesson he drew from it was that subordination and obe-

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dience to authority were even more important than liberty. At first, as a Protestant and outsider, Cuvier welcomed the Revolution as putting an end to the despotism of the king and the antiquated privileges of the aristocracy and clergy. The original bourgeois reforms of the Revolution as embodied in the declaration of 4 August 1789 appeared to Cuvier "a model of philosophical justice." "Liberty and equality are engraved in the heart of every enlightened man," he wrote to Pfaff in December 1790.58 But as radicalism gained the upper hand, Cuvier quickly became disillusioned with all parties. The idealistic attempt to create a new order, he now saw, had led only to political assassinations and mob violence. By 1792 he declared to Pfaff: "My manner of seeing the revolution has changed . . . In any case, remember always that for honest men, liberty exists under all forms of government." Greatly distressed by mob executions in the name of liberty, Cuvier came to the conviction that no liberty could exist without subordination.59 Long after the Revolution, in his eloges of scientists who had lived through it, he spoke with fear and loathing of those days of mob rule.60 Given the unstable conditions of the Empire and Restoration, Cuvier's fear of social unrest was understandable, and it is important for understanding his views of science. Cuvier, like many men of his day, sincerely believed that ideas had been at the root of the Revolution. Writers such as Rousseau and Voltaire had been responsible for arousing the revolutionary fervor of the populace. During the Revolution, Cuvier read with relish all the current religious and scientific "systems," ridiculing their contents in letters to Pfaff. He justified to Pfaff his interest in such obscure works as the mystical and "crypto-Catholic" doctrines of the Martinists because such doctrines often have the greatest influence on the course of politics, and thus they are excessively interesting to the observer. For proof, I could show you what influence the philosophy of Rousseau and that of Voltaire have had on the current events in France.61

Shaken by the Revolution, Cuvier came to fear the effects of "systems," whether strictly political or supposedly scientific, on the uneducated multitude. His anxiety concerning the possible abuse of science, although exaggerated, was in some measure legitimate because many times before the Revolution and many times afterward, scientific theories had indeed been politicized in France. During the Enlightenment, the philosophes had attacked the authority of the Catholic Church through their biological theories. Just before the Revolution, the subtle fluid theories of Mesmer, initially presented as pure science, became joined to revolutionary ideas taken from Rousseau and others. Later, in the nineteenth century, Gall's phrenology and Felix Pouchet's spontaneous generation became liberal causes, and still later Darwin was translated into French, not just in the name of science, but also as part of a deliberate assault on the Church.62 Cuvier regarded education as providing the foundation of civilization and the main means of securing the social order. He thus promoted centralization, for he believed that a centralized system could better inculcate respect for laws

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and the duties of citizenship. According to his biographer John Vienot, he was often heard to say: "Give [us] schools before giving political rights. Make the citizens understand the duties that the state of society imposes upon them. Teach them what political rights are before letting them enjoy them."63 The liberals reproached Cuvier for working with the minister Guizot, on censorship laws and for acquiescing in the electoral law of 1820, which restricted the liberties accorded by the king in 1816. Though often accused by detractors of servility to whatever regime was in power,64 he acted out of something beyond mere opportunism. Any lawful regime, he sincerely believed, was preferable to disorder. Baron Pasquier, a member of the Chamber of Peers, expressed well this multidimensional concern with order in his eulogy of Cuvier: The study of nature—the continual admiration of the order which is found in its smallest components—of this order which produces, which animates, which conserves all, instilled in him the need to find everywhere the same principle in political and social organization, to care for it and to defend it. And as governments are everywhere the natural guardians of order, they were, for that reason alone, the object[s] of his particular interest.65

In short, in both politics and science, Cuvier believed in the necessity for expert and elitist authority. Cuvier's attitude towards religion can best be understood as a reflection of his political views. Religion was important to him chiefly for its value in maintaining social stability. Though he attended church regularly, and was for many years vice-president of the Bible Society of Paris, one could not call him devout. There is little evidence that he had any interest in exploring the metaphysical, emotional, or mystical sides of religion and theology. As religious doctrines were not amenable to human reason, he held, they should be accepted on faith alone. Religious belief was a private matter and not a subject for public debate. It is not surprising that as a Protestant in a predominantly Catholic country undergoing a religious revival, Cuvier was particularly concerned with avoiding doctrinal clashes.66 Cuvier's deep-seated aversion to religious controversy emerges most clearly in his eloge of the English chemist Joseph Priestley, delivered in 1805. Employing a typical rhetorical strategy, Cuvier separated and contrasted two Priestleys: one, a "circumspect physicist," and the other, a rash Unitarian theologian who "takes on with audacity the most mysterious questions" and "scorns the beliefs of centuries." Since to Cuvier, an eloge was not a eulogy but a moral lesson, he found it perfectly legitimate to discourse twice as long on Priestley the theologian and politician as on Priestley the scientist. Priestley, according to Cuvier, had committed the unforgivable sin of becoming embroiled in religious polemics, recklessly attacking settled beliefs, and forcing upon others his opinions on matters which were forever hidden from the understanding of man. By arousing religious controversy and by promoting "a politics of dissidence," Priestley brought upon himself the violence of the angry mob which razed his home and forced him to emigrate to America. The lesson to be learned by this great man gone astray was that one paid dearly for political and

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religious immoderation. Priestley's "greatest fault," Cuvier concluded, "was to have mistaken his vocation and attached too much importance to his particular opinions on matters in which the most important of all sentiments ought to be the love of peace."67 Unlike his British counterparts, Cuvier rarely mentioned God or Providence in his work, nor did he attempt to correlate Genesis with the evidence of geology. This is not, as has sometimes been suggested, because Cuvier, or French naturalists in general, were uninterested in the religious ramifications of their science, or because, as employees of the state, Cuvier and his colleagues did not feel compelled to deal with such issues.68 Natural theology in the early nineteenth century was largely a British phenomenon. As the differences in the relation of science and religion in France and England have never seriously been studied, one can only conjecture on the causes. They stem no doubt from differences in culture, both the culture of the French nation and the more particular subculture of French science. Natural theology in the form popularized by scores of writers from John Ray's The Wisdom of God Manifested in the Works of the Creation (1691) to Archdeacon Paley's Natural Theology (1802) and the Bridgewater Treatises (1833-1836) was a product of Protestant culture, and it made little headway in Catholic France. At the beginning of the eighteenth century there were a few French works written with the purpose of demonstrating God's power and wisdom through an examination of his works, among them the widely popular Spectacle of Nature by the abbe Pluche and Reaumur's volumes on the natural history of insects.69 But works written in this genre were effectively discredited by the Enlightenment writers and they never regained a foothold in France. Several reasons may be suggested for this. First, the Catholic Church did not place the same emphasis as the Protestant denominations on natural theology. What mattered in Catholicism was not the literal truth of the Bible but the accumulated doctrines of the Church, and those had to be accepted on the basis of authority. The demonstration of God's attributes through biology might lead to belief in God, but would not necessarily lead one to the doctrines of Roman Catholicism. Then too, the Catholic Church did not encourage its members to interpret the Bible for themselves. In the Roman Catholic tradition, the meaning of biblical passages was a matter for theologians, not laymen. Thus, works which purported to reconcile Genesis and science were suspect. Scriptural geology written by scientists in nineteenth-century France therefore would be probably have been perceived by the Church hierarchy as meddling in religion rather than as welcome support for religion. Second, British natural theology was linked to the particular economic and social context in Britain, and especially to the Industrial Revolution. The watchword of British natural theology was not merely utility, but contrivance, the minute correlation of means to ends. This display of contrivance, especially seen in such works as Charles Bell's Bridgewater Treatise The Hand, Its Mechanism and Vital Endowments, as Evincing Design (1833), was associated with a milieu in which man-made contrivances, epitomized by the machines of the Industrial Revolution, were familiar artifacts. In France, where indus-

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trialism took on a different form, the extended machine analogy was not as prevalent. French naturalists, even when they adopted the utilitarian argument from design as found in the works of Cuvier—that all structures were intended for predetermined functions—had none of the British obsession with contrivance. Cuvier, in his writings, stressed the overall correlation of functions within an organism, not the manner in which the smallest detail exhibited design. As a Protestant, Cuvier was, in fact, more concerned with adaptation than most other French naturalists, and because of this his work was easily assimilated by British naturalists, albeit with a certain amount of reworking to fit it into the British context.70 A third reason that British-style natural theology was not to be found in France concerns the structure of the French scientific community. French scientists had by the 1790s become more professionalized than their British counterparts. They were employed by secular public institutions and wrote mainly for their scientific colleagues and students, not for a general educated audience. With professionalization came a desire on the part of scientists for autonomy from other and older professions. In France a reconciliation of natural science with the Bible not only would have been discouraged for meddling with religion, but also would have been scorned as unwarranted popularization.71 Finally, Cuvier's caution in discussing the role of God in nature can be partly attributed to the dominance of the physical scientists in the French system of institutions. Because the Academie des Sciences took on the role of chief arbiter of science, natural scientists were placed in a position of being judged, not just by other naturalists, but by astronomers, mathematicians, mechanicians, and chemists. These latter sciences, influenced by Enlightenment thought and having less call for natural theology than the life sciences, had largely abandoned it. Laplace, a student of the Encyclopedist Jean Le Rond d'Alembert, and chief among the physical scientists, had accounted for the origin of the solar system in his mathematical system of the world by the nebular hypothesis, leaving out all reference to a Deity. When this omission was questioned by Napoleon, Laplace is said to have replied, "I have no need of that hypothesis." Even if not wholly authentic, the statement, as Hahn has suggested, was a fair representation of his position. It was not that Laplace intended to be irreligious. He simply did not believe religious doctrines had a place in scientific discourse. In fact, Laplace argued in a manner similar to Cuvier that, as first causes were inacessible in science, scientists should concern themselves only with positive knowledge.72 Cuvier, who came to occupy a position in the natural sciences parallel to that of Laplace in the physical sciences, was much concerned to impress Laplace, and even dedicated his Recherches to him. It is not to be wondered that Cuvier refused to inject the Creator overtly into the biological sciences when He had been so definitively ejected by Laplace from the physical sciences. The French attitude toward natural theology is illustrated by the case of Henri Marie Ducrotay de Blainville, professor of zoology at the Faculte des Sciences and Cuvier's successor at the Museum, one of the few examples of a French naturalist who argued explicitly that science and religion provided

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mutual support. In his course on history of science begun at the Faculte des Sciences in 1839, Blainville tried to demonstrate, on the one hand, that biology when correctly conceived (that is, according to Blainville's doctrine of the animal series, a much modified chain of being), lent support to Catholicism, and on the other hand that the animal series could be deduced a priori from philosophical (i.e., religious) considerations.73 Blainville's enterprise received little sympathy. All commentators, even his student and biographer Pol Nicard, felt he was out of line in mixing science and religion. Constant Prevost, a geologist and friend of Blainville, summed up the general reaction of French naturalists when he wrote to Blainville: "To make faith the basis of science—would not that submit the latter to despotism? To deliver faith to the uncertainties of science—would not that expose the former to anarchy?"74 Cuvier, in a delicate position as a Protestant in a Catholic country, was in accord with most of his colleagues that an overt attempt to demonstrate religious doctrine by science was improper. By failing to understand the cultural differences between French and British natural history, historians have often tended to downplay religious motivation in Cuvier's work.75 That Cuvier had legitimate scientific justification for his catastrophism and antievolution cannot be denied. One must, however, distinguish between motivation and the logic of justification. To say that religious motives affected his scientific doctrines is not equivalent to saying that he blindly rejected evolution on the basis of religious faith. Insofar as the evidence allowed, Cuvier's scientific theories upheld the traditional view of God's creation and providential concern for His creatures, the recent creation of man and the biblical Flood. It is no accident that Cuvier found geological evidence of a great inundation not so very long ago. Although he did not seek to verify Moses through geology, as a British naturalist might, he nonetheless did associate his last revolution of the globe with Noah's Flood. Comparing the account in the Bible with the accounts of sacred works of other cultures, he noted that all written records pointed to a flood about four thousand years ago.76 It is difficult to make the case that Cuvier supported the traditional age of mankind, about six thousand years, merely to argue against Lamarck that man could not have caused extinction. The teleological principles of conditions of existence and correlation of parts, although not presented in an explicitly religious framework, were, in fact, intended as a defense of God's providential concern for His creatures. Despite Cuvier's strategy of avoiding discussions of religion in his works, he was perceived by others, particularly by nonscientists, as a defender of traditional Christianity. The British geologist Charles Lyell accused Cuvier of "trading in humbug" by injecting religion into science. Novelists George Sand and Honore de Balzac pictured him as an upholder of the Bible, who by his insistence on a narrow science of facts, prevented man from questioning the creation. Cuvier himself noted with pride that the abbe de Frayssinous, grand master of the University, had cited his "Discours preliminaire" in his sermons "as if it were by a Father of the Church." And Cuvier's disciple Charles Leopold Laurillard felt the need to defend him from the charge "that his laborious his-

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torical and zoological works were conceived in a spirit of system, because they accord in part with the tradition of the Bible."77 In an era when reformers freely used scientific doctrines to support anticlericalism and social change, it was prudent to eliminate explicit religious themes from science and to defend traditional religion implicitly through scientific arguments. One could best maintain political and religious orthodoxy by banning controversial and potentially heretical topics such as spontaneous generation and evolution from discussion as outside the bounds of science. Though he generally refused to defend his scientific doctrine on religious and political grounds (at least until the 1820s), on occasion Cuvier publicly revealed his deeply held conviction that natural history rightly studied did have a beneficial effect on religion and politics. In the summary of his Historical Report on the Progress of the Natural Sciences, presented to the Emperor in 1808, Cuvier argued that the sciences were of value because they calmed the passions and inculcated the habit of rational thinking: To conduct the human mind to its noble destination—the knowledge of truth; to spread healthy ideas as far as the least educated classes of the people; to lead men away from the empire of prejudices and passions; to make reason the arbiter and supreme guide of public opinion—this is the essential object of the sciences. This is how they concur to advance civilization, and this is why they ought to merit the protection of Governments who wish to render their power unshakable by founding it on the common well-being.78

And in Le regne animal (1817), he asserted that the skill of classifying ideas that one obtains from natural history is conducive to public order and fosters an appreciation for God's handiwork: It consoles the unhappy, it calms hatreds. Once elevated to the contemplation of that harmony of Nature inexorably ruled by Providence, how feeble and petty are found to be those jurisdictions left to the free will of man . . . I avow it proudly: these ideas have never been alien to my works, and if I have sought with all my means to propagate this peaceful study, it is because in my opinion, it is more capable than any other of feeding the hunger for activity which has so greatly contributed to the troubles of our century.79 Institutions, Careers, and Patronage

In order to understand Cuvier as a "legislator of science," one must consider not only his ideas and scientific methodology but also the community in which he worked. The small and centralized community of naturalists that emerged in Paris in the three decades following the founding of the Museum was the primary audience for the controversy between Cuvier and Geoffroy. Cuvier's effectiveness in promoting his doctrines and his methodological prescriptions depended on the particular structure of this community and the key role of patronage politics. Although successful careers in science have always depended on patronage, it is with good reason that historians have especially pointed to the importance of patronage in early nineteenth-century French science.80 Cuvier, by virtue of his scientific stature and his key posts in the French

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system of scientific institutions, could more readily dispense patronage than any other naturalist, thus acquiring leverage for advancing his scientific ideas. It is thus essential to ask, who were the members of the community of naturalists over whom Cuvier assumed leadership, how did they acquire their training, and how did they secure financial rewards and recognition? For a community of zoologists and comparative anatomists to emerge, many more avenues of support were required than just four professorships at the Museum. If the posts held by Geoffroy, Lamarck, Lacepede, and Cuvier had been the only positions for zoologists and comparative anatomists in France, Cuvier's leadership would have been ineffective, to say the least. While by far the most important, the Museum represented only one of several institutional changes that took place in natural history during the Revolution and Empire that led to the creation of new positions and enhanced opportunities for remuneration. Although the four Museum professors continued to hold their chairs until they died (in Geoffroy's case until he went blind in 1841), they did not always teach their courses. The practice of hiring suppleants provided opportunities for several younger naturalists to teach. Lacepede, for example, did not teach after 1804 but continued as titulary professor until his death in 1825. Constant Dumeril, a disciple of Cuvier, taught the course for over twenty years before claiming title to the chair. When Cuvier was away on administrative assignments for the Napoleonic Empire, Blainville, then a disciple, took over the teaching. After 1814, since Cuvier rarely lectured, a succession of suppleants were given the opportunity to give the annual course. Lamarck also ceased teaching by 1818, and the courses and much of the curatorial work were taken over first by the entomologist Pierre-Andre Latreille and then by Jean-Victor Audouin. In 1830 and again in 1837 the faculty was enlarged by the division of Lamarck's chair and the creation of a new chair of comparative physiology. These new chairs, and vacancies caused by deaths, allowed Blainville, Latreille, Audouin, Achille Valenciennes, Frederic Cuvier, and Pierre Flourens all to become professors in the 1830s. (See Appendix A.)81 In addition to teaching chairs, the Museum provided a number of subordinate positions which could be used by beginning naturalists as stepping stones to the professorate. The internal regulations of the Museum called for four aide-naturalistes, with more to be added if necessary, who were to prepare chemicals, mount zoological specimens, make anatomical preparations, and arrange the galleries. By midcentury, there were sixteen aide-naturalistes and twenty-three preparateurs on the Museum's staff. Some of the holders of these positions, such as Cuvier's aide-naturaliste Laurillard, were good technicians, who never advanced beyond a subordinate role. However, the professors specifically recognized, in a committee report of 1803, that a primary function of these posts was to serve as "an honorable training ground [une pepiniere] where excellent naturalists will be produced and where professors can with advantage be recruited."82 Geoffroy's son, Isidore Geoffroy Saint-Hilaire, Latreille, Audouin, and Valenciennes were all able to advance from aide-naturaliste to professor of zoology at the Museum.

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Other teaching opportunities in public institutions were afforded by the College de France, the medical schools, the secondary schools, the Ecole Veterinaire d'Alfort, and later the Facultes des Sciences and the Ecole Normale. The impact of these opportunities, however, was to some extent limited by the practice of cumul, whereby the most significant posts outside the Museum were taken by men who also were or became professors at the Museum. The College de France, which owed its origin to the royal lectures created by Francis I in 1530 to teach humanist subjects not taught at the University of Paris, had by the late eighteenth century some twenty chairs in languages, literature, law, history, medicine, and the sciences. Like the courses at the Museum, those at the College de France were free and open to the public, and no degrees were conferred. Daubenton, the first occupant of the chair of natural history created in 1778, limited his lectures to his specialty, mineralogy. However, when Cuvier was appointed in 1800, the subject matter of the courses shifted to zoology. Initially the College de France simply gave Cuvier another podium, a more varied audience, and an additional salary, but after 1814 it provided employment to several more suppleants. Zoology, however, never became entrenched in the College de France. It was taught only by Cuvier and his disciple and successor Georges Louis Duvernoy. When Duvernoy died in 1855, the new titulary, Flourens, diverted the subject of the chair to physiology. (See Appendix B.)83 At the Ecole de Medecine in Paris, a chair of comparative anatomy was created in 1802 for Cuvier's disciple Dumeril. In Paris, as well as in the provincial medical schools, other naturalists, though usually botanists, were appointed to teach histoire naturelle medicale—a hybrid between natural history and materia medica.84 In the 1780s the Ecole Veterinaire d'Alfort, established in 1766, provided three positions in rural economy, held by Daubenton, the professor, Broussonet, his adjunct, and Olivier, also attached to the chair, and a professorship in animal anatomy held by Vicq d'Azyr. Zoology was taught there in the early nineteenth century by Olivier, Latreille, and after 1815, Anselme-Gaetan Desmarest, a former preparateur for Cuvier at the College de France.85 The public secondary schools, established in 1795 and later made part of the University system, offered another significant source of employment for zoologists. The ecoles centrales, founded by the Directory, were each provided with a professor of natural history. Early in his career, Cuvier taught at the Ecole Centrale du Pantheon, and Brongniart taught at the Ecole Centrale des Quatre Nations. Because they were based on the suspect philosophy of the ideologues, the ecoles centrales were suppressed by Napoleon in 1802 and replaced by the lycees, fewer in number and more traditional in curriculum. Except for the schools in Paris there were no longer professors of natural history, but some natural history continued to be taught in the course of sciences physiques. Chairs of natural history were gradually reintroduced into the secondary school curriculum after 1830, until by 1840 nearly forty positions were available.86 Rounding out the catalogue of public institutions was the Imperial University, organized in 1808, and including all public degree-granting institutions of

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higher education in the Empire—faculties, lycees, and communal colleges. Although private schools continued to exist in France, the state thenceforth assumed a monopoly on the granting of degrees. Faculties of theology, law, and medicine had already been established after the Revolution and were simply incorporated into the University. The other two types of faculties, letters and sciences, were true novelties. By 1815 there were ten faculties of sciences in operation, each with a chair of natural history, and some with separate chairs of botany, zoology, and mineralogy and geology.87 Zoology at the Paris Faculte des Sciences was provided with a titulary professor and an adjunct. By the decree setting up the University, titulary professors were to be chosen from among professors at existing institutions in Paris.88 Thus, Geoffroy became first professor of zoology, while still retaining his post at the Museum. But the adjunct professorship, first offered to Duvernoy, was given in 1812 to Blainville, then Cuvier's disciple, and provided him with status, income, and a podium for his views until he could also obtain a chair at the Museum in 1830. (See Appendix C.) The entire educational system was centrally administered during the Empire and much of the Restoration by a grand master, who after 1824 coincided with the minister of public instruction, and an Academic Council. Although not a professor at the Faculte des Sciences, Cuvier exercised considerable influence over appointments to the faculties in Paris and in the provinces through his seat on the Council. At first, even the Paris Faculte des Sciences was no more than another source of free, public courses, and its existence did little to transform the training of zoologists.89 The common practice for aspiring zoologists in this period was instead to obtain a medical degree and at the same time to audit the courses at the Museum, the College de France, and the Faculte des Sciences. Along the way, if they needed it, they would also submit a thesis and take the examination for the doctorat.90 There were few degree candidates at any of the Facultes des Sciences aside from candidates for the baccalaureate and, in Paris, students from the Ecole Normale. Only later in the century did the Faculte become a place where one might obtain a systematic formal education in the sciences. Geoffroy's successor, Henri Milne Edwards, was the first professor of zoology to take seriously his role as a master preparing future careerists.91 Also part of the University system was the Ecole Normale, established in 1808 to educate teachers. Originally the school did not have its own faculty, and students were expected to take courses at the other public institutions in Paris, especially the Faculte des Sciences and Faculte des Lettres, which were situated next door. When the school eventually instituted its own instruction in the sciences, the post of maitre de conferences in zoology was given in 1831 to Valenciennes, a disciple of Cuvier, who became professor of mollusks, worms, and zoophytes at the Museum in 1832 as well as professor of natural history at the College Rollin. Thus, by the 1840s there were an admittedly small but significant number of teaching positions related to zoology in public institutions, and as the century progressed the number of these posts continued to expand.92 In France, in contrast to the situation in England, almost all teaching of the

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sciences took place in public institutions. One of the few private teaching establishments in Paris was the Athenee, a school for adult education, founded in 1781 and in existence until 1847. It served as an early teaching post for several future professors of the Museum. Courses in natural history were taught by Alexandre Brongniart from 1797 to 1800, by Cuvier from 1801 to 1810, by Blainville from 1812 to 1828, by Isidore Geoffroy from 1831 to 1834, and by Audouin in 1835.93 Private enterprise, although it generated few teaching positions, did provide other means by which beginning naturalists could earn a living until such time as they could obtain a professorial chair. Naturalists trained as physicians could fall back on their medical skills to provide a living. Ties between physicians and naturalists in the early nineteenth century were still close. The biological disciplines were yet in the process of emerging out of the medical context. Almost all of the zoologists and comparative anatomists trained after 1795 took medical degrees. On the other hand, any medical student with an interest in research frequented the courses in natural history and comparative anatomy at the Museum, the College de France, and the Faculte des Sciences. Medical training was thus the chief avenue to a career in the life sciences, for it provided instruction in anatomy and dissection techniques, the opportunity to attend courses elsewhere, and a secure profession if needed. Publishing contracts offered perhaps the best means for aspiring naturalists to make ends meet, while at the same time enhancing their career prospects. Money could often be earned by editing a scientific journal or writing reviews of meetings of scientific societies for journals or newspapers. Audouin, Adolphe Brongniart and Jean-Baptiste Dumas founded the Annales des sciences naturelles in 1824 well before any of the three had an academic appointment. In the 1810s and 1820s, Blainville and Flourens, neither of whom were yet members of the Academie, wrote weekly reviews of the Academie meetings for scientific journals. One could likewise earn money by writing textbooks for secondary schools, as Milne Edwards was to do in the 1830s. An important source of income for naturalists came from the many contracts to write articles for the numerous multivolumed illustrated encyclopedias intended for the public, such as the Dictionnaire classique d'histoire naturelle (1822-1831) edited by the zoologist J.B.G.M. Bory de Saint-Vincent, and the sixty-volume Dictionnaire classique des sciences naturelles (1816-1830), edited by Frederic Cuvier. Besides providing employment for future professors of the Museum, publishing enterprises enabled a fringe group of physicians and others to keep their hand in zoology and comparative anatomy even though they were never able to enter the Museum or Academie. The institutions of French science created during the Revolution and Empire paved the way for a transformation of zoology by providing full-time careers for a small but critical mass of researchers. These were men who identified themselves as zoologists and comparative anatomists and came to see themselves as part of a professional elite.94 An unofficial but nonetheless recognized procedure was established for new recruits to enter this elite. A wouldbe zoologist was expected to take a medical degree, attend courses at the

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Museum, the College de France, and the Faculte de Sciences, and attend meetings of the Societe Philomatique and the Academie des Sciences. Having introduced himself to the professors, the young man would then gain access to the research facilities of the Museum, engage in independent research and present the results to the Academie des Sciences, and acquire a mentor who would help him to obtain his first paid post. Personal patronage was vitally important in such a system. It was helpful for publishing research and obtaining writing contracts, necessary for gaining access to the Museum collections, and essential for getting a teaching post. At the tribunal of French science, the Academie des Sciences, patronage was the order of the day. Although no longer the locus of original research that it had been in the eighteenth century, the Academie was no less important for the life of science, for it wielded an immense power over the direction of scientific research in France.95 Every Monday afternoon the leaders of science gathered together to hear and pass judgment on papers presented to them, to award prizes for competitions they sponsored, and to vote upon candidates to fill vacancies not only in their midst but also at the Museum, the College de France, and other major scientific institutions in Paris. Despite the great competition for seats, all the leaders of zoology eventually entered the Academie. They were elected not as old men, but at the height of their careers, usually in their late thirties or early forties. It was actually easier in the nineteenth century to become a member of the Academie than to become a professor at the Museum. Almost all the future professors of the Museum were elected to the Academie long before obtaining their professorial chair. (See Appendix D.)96 The two most powerful posts in all of French science were those of the permanent secretaries (secretaires perpetuelles) of the Academie des Sciences. In the 1790s the First Class of the Institut elected its secretaries for a limited period of time, but with Napoleon's reorganization of the Institut in 1803, a return was made to the permanent secretaries of the ancien regime. Two secretaries, each paid 6000 francs a year, were instituted, one for the "mathematical sciences" and one for the "physical sciences," and, as their title suggests, they held their posts for life. It was Cuvier's position as permanent secretary for the "physical sciences," coupled with his place on the University Council, that gave him such inordinate power in controlling the affairs of Restoration science. As permanent secretary, he acted as spokesman for the Academie in its relations with the government, maintained official correspondence, kept accounts of meetings, presented annual reports of the work of the Academie, and delivered eloges of deceased members. Through its reports on papers submitted, prize competitions, and the annual reports of the work of its members, the Academie passed judgment on what was appropriate or inappropriate research. If a young man desired recognition, he had to adapt his research to meet the standards set by the members of the Academie. Since all the choice posts in French science were controlled ultimately by the government, it was impossible to succeed in a scientific career without the approval of at least one faction of the Academie. Ambitious young researchers presented memoirs to the Academie at the earliest opportunity, for

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this was the way to become known to those of influence. In fact more papers were presented by nonmembers than by members. When a nonmember read a paper, the Academie chose a commission consisting of two or three members to present a report at a later meeting. In the cases of the physiologists Flourens and Magendie, and the zoologist Audouin, reports by Cuvier on their work were instrumental in establishing them in their careers.97 Regarded by the government and public alike as the final arbiter on scientific questions, the Academie contributed to the centralization of French science and, later in the century, to its general conservatism. As leader of the naturalists in a centralized and government-sponsored system of science, Cuvier was in a unique position to dispense favors to disciples in return for their allegiance. Although one cannot find a clearly delimited "school of Cuvier," Cuvier did collect about him a number of collaborators and proteges who were in accord with the essentials of his doctrine. First, they agreed that the major goal of zoology was to arrange animals according to the natural system, and they assumed in their work the four embranchements, the use of comparative internal anatomy (rather than external characters), and final causes. Second, they followed him in rejecting all those theories he had labeled as speculative, in particular the great chain of being, spontaneous generation, phrenology, evolution, and Geoffroy's "unity of organic composition." And finally, in varying degrees, they echoed his methodological precepts on restricting science to "positive facts." Some of his followers rejected any theorizing whatever. For all of his major publications, Cuvier employed assistants and collaborators to do the more mundane tasks: to prepare specimens, sketch plates, check references, and write up details in accordance with his master plan of organization. It has been said of Cuvier's proteges that most were second-rate men whose work Cuvier exploited.98 Blainville bitterly accused his former master of allowing his followers no independence. The naturalist-voyager JeanRene-Constant Quoy observed that "M. Cuvier had the grand art of knowing how to employ men and plucking from them all the fruit possible." Quoy refused to become a disciple and accept Cuvier's offer of patronage, because "the star of Jupiter would not have eclipsed her satellites as much as I would have been."99 Cuvier's chief disciples, all of whom figure to some extent in the controversy with Geoffroy, were his aide at the Museum, Charles Leopold Laurillard; his collaborator on the first two volumes of the Lecons d'anatomie comparee, Constant Dumeril; his relative and collaborator on the last three volumes of Lecons, Georges Louis Duvernoy; his younger brother, Frederic Cuvier; and his collaborator on the Histoire naturelle des poissons, Achille Valenciennes. All spent their careers in the shadow of the great naturalist. In return for their loyalty and subordination, Cuvier found them positions and helped them to enter the Academie des Sciences. Laurillard, more a technician than a trained naturalist, became keeper of the galleries of comparative anatomy. Dumeril became professor of comparative anatomy at the Ecole de Medecine (1802), suppleant to Lacepede (1804), and finally titulary professor of reptiles and

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fishes (1825). Duvernoy was chosen professor of natural history at Strasbourg (1827) and eventually inherited his mentor's chairs at the College de France (1837) and at the Museum (1850). Frederic Cuvier, with no training as a naturalist before he joined his brother in Paris in 1797, became keeper of the menagerie (1804), a functionary in the administration of public education (1810), and finally professor of comparative physiology at the Museum (1837). Valenciennes, despite his mediocre talents, became professor of annelids, mollusks, and zoophytes at the Museum (1832) and maitre de conferences at the Ecole Normale (1831). All except Laurillard were elected members of the Academie des Sciences.100 Not all of Cuvier's immediate disciples were second-rate naturalists, but the exceptions tended to prove the rule. Pierre Flourens, who inherited Cuvier's post of permanent secretary of the Academie, won recognition as a brilliant experimental physiologist whose ablation experiments distinguished the functions of the cerebrum, cerebellum, and medulla. Beginning his career as a disciple of Geoffroy, Flourens gradually moved into Cuvier's orbit in the 1820s. Although he modified his research to retain Cuvier's patronage, his creativity was less hampered than it might have been were he a naturalist rather than a physiologist. Nonetheless, his later work did not live up to the promise of his initial successes.101 Henri Marie Ducrotay de Blainville, on the other hand, was able to become an innovative naturalist only because he broke away from Cuvier's domination.102 Blainville affords an especially good example of the way in which Cuvier's patronage could start an aspiring naturalist on a successful career. Nearly thirty when he decided on a scientific career, Blainville entered medical school, obtaining his degree in 1808, and simultaneously attended courses at the Museum. In 1809 Cuvier and Geoffroy admitted him into their galleries, where he embarked upon independent research on reptiles, fishes, and comparative myology (muscle anatomy), the areas which he regarded as the least studied of vertebrate anatomy. His opportunity for advancement came in 1810 when Cuvier offered to make him a collaborator on a comprehensive work on comparative anatomy (a work that never materialized). It was agreed between them that Blainville would communicate to Cuvier all his research on the myology of mammals and birds in return for an annual indemnity, the arrangement to endure until such time as Cuvier could find an independent post for his disciple. It was not long before Cuvier was able to honor his commitment. At his request the Athenee chose Blainville as his successor. While he was away on administrative assignments, he allowed Blainville to teach his courses at the College de France and the Museum. Cuvier's most important service for Blainville was assisting him to obtain the adjunct professorship at the Faculte des Sciences in 1812. All these benefits of patronage came to an abrupt end in 1816 when Blainville and Cuvier broke relations. Given their personalities, a rupture was almost inevitable. Cuvier, authoritarian and vain, preferred disciples who would flatter him and do his bidding. Blainville, difficult and oversensitive, was too proud to be content in Cuvier's shadow, and chafed under the

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conviction that Cuvier was exploiting him. He angered his benefactor by deliberately changing the outline of Cuvier's course on "sensibility" at the Museum and teaching his own doctrines in opposition to Cuvier's. The final break came over a trifling incident of wounded pride at one of Cuvier's Saturday night salons. Henceforth, Blainville was denied access to the collections of comparative anatomy at the Museum and forced to make his way against the opposition of his former benefactor. Fortunately for Blainville, he already had an independent position and reputation by 1816. More determined than ever, he made it his mission in life to succeed despite Cuvier. With the patronage of the astronomer Francois Arago, he entered the Academie in 1825 after a heated election in which the rival candidate was Frederic Cuvier. In 1830 he won one of the two chairs into which Lamarck's chair was divided, but his crowning triumph came when he succeeded to Cuvier's chair of comparative anatomy in 1832. Although Cuvier commanded a great deal of patronage, Blainville's success showed that his control over natural history was far from absolute. Increasingly in the 1820s new factions led by Arago and Brongniart supplanted the "old guard" led by Laplace in the physical sciences and Cuvier in the biological sciences. That Cuvier less often had his way in the 1820s was not without significance for the Cuvier-Geoffroy debate. Georges Cuvier went unrivaled during his lifetime as the greatest naturalist France had ever produced. His memory and the breadth of his knowledge were nothing short of phenomenal. The order and clarity of his lectures won praise from all who heard them.103 Among his colleagues, however, he was said to have been a man more esteemed than loved. A severe and overly serious man, irritable and impatient, he was constantly occupied with his work. Having little time for pleasant chit-chat, he revealed his social instincts only in his Saturday night salons, attended by the celebrated men of the age. Yet Quoy observed that Cuvier's colleagues, while they showed him great deference, rarely attended, "which indicated little sympathy."104 Despite his lack of personal warmth, Cuvier was able, through both his enormous scientific talent and his ability to control patronage, to transform natural history into a prestigious and professional discipline. By combining the comparative anatomy of Daubenton and Vicq d'Azyr with the taxonomy of the Linnaean naturalists and the natural system of Jussieu, he was able to lay the foundations for an ongoing research program that was both rigorous and analytic. In his commitment to teleological functionalism, Cuvier lent his support (in a peculiarly French fashion) to the traditional natural theology that reigned in Britain and America. By his methodological prescriptions he deliberately sought to divorce natural history from the speculative natural philosophy of the philosophes. Whereas the philosophes had enjoyed the freedom of theorizing on the nature and origin of life, the history of the globe, and the universal properties of matter, Cuvier, motivated by religious and social as well as scientific concerns, limited natural history to descriptive comparative anatomy and paleontology and to the perfecting of the natural system of classification. In so doing, he became a legislator of science in a negative as well as a positive sense. Authors of controversial theories found it difficult to obtain a hearing

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among serious scientists in nineteenth-century France. In England, a Robert Chambers could pen a speculative theory of evolution and have it reviewed (albeit viciously) in the major journals by the best-known scientists of the day. In France, Lamarck and other such authors faced a conspiracy of silence. It is against this background of biological and methodological orthodoxy that one must study the genesis and reception of philosophical anatomy in France.

CHAPTER 4

Geoffroy and the Emergence of Philosophical Anatomy

In Philosophie anatomique (1818) Etienne Geoffroy Saint-Hilaire proclaimed the dawn of a new era when works of classification and description would be superseded by the demonstration of a new principle, that all vertebrates were formed on a single plan. The same fund of "organic materials," he boldly declared, could be traced throughout the vertebrates from the mammals down to the fishes. Geoffroy thus became the foremost proponent of the morphological or transcendental approach to comparative anatomy, which came to be known in France and England as "philosophical anatomy." For Geoffroy, the final cause of an animal, the role that it was destined to play in nature, did not foreordain its structure. On the contrary, the anatomical organization of an animal determined its mode of life—whether it crawled on land, flew through the air, or swam in water. Geoffroy, more than any other comparative anatomist, introduced and popularized the concept of homology, the central tenet of philosophical anatomy. Homologies, now defined in terms of evolution, formerly were interpreted in a transcendental sense. Whereas homologous parts are now considered to have descended from a common ancestor, in the pre-Darwinian era they were usually looked upon as evidence of an ideal pattern imposed on nature, or a blueprint in the mind of the Creator. For Geoffroy and other philosophical anatomists, homologous parts were those parts in different animals which were "essentially" the same, even though the parts might have different shapes and be employed for different purposes. The idea of homology arises naturally out of the attempt to compare different species of animals, yet for a long time naturalists did not consciously seek for homologies. Any anatomist concerned with naming the parts of animals would sooner or later be faced with the problem of when to use the same name for structures found in different animals, and when to invent a new name. For most comparative anatomists, this was not a particularly worrisome problem. When parts had approximately the same shape and performed a similar function, they were given the same name. However, if the parts appeared to have a different form and function than in man, a different terminology was intro69

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duced. Cuvier, for example, in Lecons d'anatomie comparee named and described the furcula (wishbone) in birds and the operculum (gill-cover) in fishes, implicitly assuming that these bones occurred only in birds and fishes because they were specialized pieces required for specialized functions, namely flying and respiration under water, respectively.1 Equivalent names, moreover, were only loosely applied. Anatomists were not concerned with breaking down structures such as the sternum or the parts of the skull into a number of primitive bones which had sutured together. As long as the form and function agreed, organs were given the same name regardless of the fine details of structure. Thus the explicit search for homologies involved asking a set of new questions about animal organization. Although in general comparative anatomists were not concerned with establishing homologies before the early nineteenth century, the recognition of homologies can be traced back to antiquity. Aristotle had observed the correspondence among the fin of the fish, the wing of the bird, and the arm of man and the similarity in structure of scales and feathers, but he did not specify wherein the similarities lay. One of the most striking early depictions of homologies appeared in the sixteenth-century ornithological treatises of Pierre Belon. Belon exhibited the skeletons of a man and a bird juxtaposed, the bones labeled with corresponding letters. But these drawings had little to do with the rest of his works, and the ideas implicit in them were not followed up.2 In 1784, unknown to Geoffroy, the German naturalist-poet Johann Wolfgang von Goethe, having assumed a unity of structure among the vertebrates, discovered the intermaxillary bone in man. In 1790 he published Metamorphosis of Plants, which identified serial homologies—homologies of parts within a single individual—of plants.3 Among the works readily accessible to Geoffroy was Vicq d'Azyr's memoir of 1774. This memoir not only compared limbs in man and the quadrupeds, but also compared the arm to the leg in man and the forelimbs to the hindlimbs in quadrupeds with respect to bones, muscles, and nerves. Thus Vicq d'Azyr proposed the earliest examples of serial homologies among the vertebrates. Considered a curious and ingenious piece of research by contemporaries, Vicq d'Azyr's memoir likewise bore no immediate fruit, for it inspired no further researches.4 Geoffroy was the first naturalist to promote an awareness of the pervasiveness of homologies, to develop a self-conscious method for determining them, and to found a continuing program of research. He was one of the first to analyze osseous structures in different animals into individual bones, even parts of bones, and to seek bone by bone correspondences among mammals, birds, reptiles, and fishes. Geoffroy consistently used the term "analogue" for what is now called "homologue." In effect, he appropriated a common word and gave it a specific meaning, much to the confusion of some of his colleagues. German authors, some time before 1820, had begun to employ the term "homology" for what is now called serial homology, that is, for correspondences among different parts of the same individual, a type of comparison in which Geoffroy had relatively little interest. The confusion of terminology was alleviated in 1843 when

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the British comparative anatomist Richard Owen established the modern distinction between homology and analogy. Owen formally defined "homologue" as "the same organ in different animals under every variety of form and function" and "analogue" as "a part or organ in one animal which has the same function as another part or organ in a different animal." As he explained, the legs of a crab or an insect were "analogous" to the limb of a quadruped because they all served for locomotion, but they were not "homologous" to them. In what follows, we will use the modern terms, homologue and homology, in writing about Geoffroy, but retain Geoffroy's own terms, "analogue" and "analogy," in translated quotations.5 Geoffroy's "principle of the unity of organic composition for all the vertebrates" did not appear in full flower until 1818, but his morphological orientation, as we have seen, was evident from the very beginning of his career. Philosophie anatomique was the culmination of a series of researches beginning with his 1795 memoir on the makis. Only after a long gestation was his earliest vague notion of unity of plan in the animal kingdom, which probably derived from Buffon, transformed into a workable program of research, which he embodied in a series of memoirs in 1807.6 The metamorphosis of Geoffroy's thought took place largely as a result of the three years he spent in Egypt as part of Napoleon's party of savants. There he shifted his attention from zoology to comparative anatomy and made his first essays at reducing anomalous organs to more familiar organs in other animals. Removed from the concerns of his colleagues, Geoffroy also found in Egypt the freedom to indulge in Enlightenment-style philosophizing. In his later memoirs from Egypt, none of which were published, Geoffroy rejected reasoning from final causes and displayed a deep commitment to seeing nature, organic as well as inorganic, in terms of a dynamic interaction of universal laws. The view of nature that he sketched out in the final months before returning to Paris was diametrically opposed to that espoused by Cuvier. It was in writing up his Egyptian materials for publication that Geoffroy first faced the problem of naming bones in fishes that did not have obvious equivalents in other vertebrates. Unlike Cuvier, he took the bold step of deciding that instead of resorting to specialized names for bones in fishes, he would seek their equivalents among the bones of mammals. Geoffroy's three papers on the homologies of the bones of fishes in 1807, laying the groundwork of philosophical anatomy, were so favorably received that they helped him to obtain at last a seat in the Academie. Philosophical anatomy brought to Geoffroy the professional recognition that had so far eluded him. In the period covered by this chapter—from 1798 through the reception of Philosophie anatomique in 1818-1819—Geoffroy rose in his profession to become one of the leading zoologists in Europe. His "theory of analogues" was widely acclaimed as an original, ingenious, and valuable contribution to comparative anatomy and zoology. The unexpected nature of his homologies, together with his far-reaching claims for the novelty and applicability of his doctrine, generated excitement among his students at the Museum and the Faculte des Sciences, and led several other naturalists—among them Jules-

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Cesar Savigny, Blainville, Dumeril and Henri Dutrochet—to extend philosophical anatomy beyond the skeleton of vertebrates. In the first decade of the nineteenth century, contemporaries had little inkling of an impending confrontation. Only with the publication of Philosophie anatomique did a hint of dissension become evident, as Cuvier and others who believed in the primacy of function began to voice their concerns that morphological considerations, if overextended, would lead to disturbing consequences. Intellectual Adventure in Egypt

In the beginning of 1798, Claude Louis Berthollet, a prominent chemist and personal friend of the young general Napoleon Bonaparte, came to the Museum to offer both Cuvier and Geoffroy a singular opportunity. "Come," he is supposed to have said, "Monge and I will be your companions and Bonaparte your general."7 The proposition Berthollet laid before the two young naturalists was that of accompanying Bonaparte as a member of a party of scientists on a military expedition to an undisclosed destination. Cuvier quickly declined, realizing that it was to his professional advantage to remain in Paris and consolidate his positions at the Museum and the Academie des Sciences. In his manuscript autobiography, he explained his reasoning: My calculation was soon made. I was at the center of science in the midst of the finest collection, and I was sure to do my best work there, more coherent, more systematic, and containing more important discoveries than in the most fruitful voyage. I suggested Savigny in my place; he accepted and I have always been happy with my decision.8

Having no idea of the destination or the duration of the voyage, Geoffroy enthusiastically agreed to take part in the adventure. It was in keeping with his naive romantic character that he should have accepted, as it was equally in keeping with Cuvier's more practical and politically astute character that he should have refused. While Geoffroy was away from 1798 to 1802, Cuvier was elected professor of comparative anatomy at the death of Mertrud, was chosen to fill Daubenton's post at the College de France, published the first two volumes of Lecons d'anatomie comparee, and rose to a high rank in the administration of public education. By 1803 he had been elected permanent secretary of the First Class of the Institut, the most prestigious and powerful of scientific posts, whereas in that same year Geoffroy was again unsuccessful in an election to become a member of the Institut. In 1803 Geoffroy could boast of no more than his original chair at the Museum, nor had he published a single major work. The expedition to Egypt was nonetheless of seminal importance in Geoffroy's career. Until 1798 Geoffroy had dealt exclusively with the classification of mammals and birds. His research in comparative anatomy, stimulated by Cuvier, was limited to unraveling narrow problems of taxonomy. It was in Egypt that he began to study comparative anatomy for its own sake. Intrigued

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by peculiarities of anatomical structure, he began the attempt to reduce anomalies to more common and well-known forms. Egypt also gave Geoffroy the opportunity to give full vent to his penchant for generalization. Away from the restraining hand of his colleagues, enchanted by an unexplored land with an ancient history, overwhelmed by constant changes of scenery, by hunger, disease, and war, Geoffroy forsook more than ever close detailed studies for speculation on the broadest of topics. He tackled a host of perennially debated physical and physiological problems, which were for the most part not amenable to observation and experiment and could only be investigated by rational argumentation. Had he been in Paris, Daubenton or Cuvier might have convinced him that such topics were outside the scope of zoology and comparative anatomy and that, moreover, his didactic approach would be unproductive of "positive" results or of honor. But in Egypt, Geoffroy was not to be deterred by any notion of scientific specialization. Before he returned to Paris, he had constructed a grandiose theory of fluids and forces that was supposed to explain all the phenomena in the universe. Only when he was about to rejoin his colleagues in France did he fear the repercussions to his professional reputation, and request Cuvier's intervention to save him from ridicule by the leaders of the physical sciences. Bonaparte took with him on his military expedition to Egypt a Commission on Science and the Arts, a microcosm of French culture, to reflect the glory of France and her leading general.9 Among the members of the illustrious company were Berthollet, the mathematicians Gaspard Monge and Joseph Fourier, the geologist Deodat de Dolomieu, the physicist Etienne Louis Malus, and the botanist Alire Raffeneau-Delile. Savigny, designated as a substitute by Cuvier, worked along with Geoffroy on zoology, and Henri-Joseph Redoute, brother of the celebrated floral artist Pierre-Joseph Redoute, sketched the specimens they found. On 19 May the expedition set sail, its destination still a secret to all but a handful of the participants. "Goodbye, my friend," Geoffroy wrote to Cuvier from Toulon, "love me always. Do not cease to consider me as a brother."10 The first two years of the expedition to Egypt were a happy and productive period for Geoffroy, and gave him a welcome feeling of importance.11 The savants were to form part of the Institut d'Egypte, an imitation of the Institut in Paris, which was officially established on 22 August 1798 and housed in a spacious palace on the outskirts of Cairo. Geoffroy was a member of the committee which on 21 August drew up the details of organization, and he was designated along with Berthollet, Dolomieu, Savigny, and others a member of the section of physics (the sciences were divided into physics and mathematics).12 One of the most zealous academicians, Geoffroy presented frequent papers, sat on commissions, and published his work in the newly founded "journal of literature and political economy," La decade egyptienne. He had the opportunity to go on local excursions with Bonaparte to the Pyramids and to the quarries in the neighborhood of Cairo, as well as on three major journeys to the eastern part of the Nile Delta (winter 1798-1799), up the Nile as far as the first cataract (August-September 1799), and to Suez and the Red Sea (win-

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ter 1799-1800). From the waters of the Nile, Lake Manzala, the Red Sea, and the Mediterranean he collected new and unusual species of fishes, and from ancient tombs along the Nile he gathered a rich collection of human and animal mummies. No one could henceforth reproach him, he wrote to Cuvier, for not being a true naturalist because he had not ventured forth from the confines of the Museum.13 A respected member of an Institut at last and a privileged companion of General Bonaparte, Geoffroy came to see himself as a hero of science about to achieve immortal glory. Although Geoffroy contributed to systematic zoology sufficiently to merit the compliment of his colleagues at the Museum that "no voyager since the celebrated Dombey has given our collections so considerable an increase,"14 his real interests lay beyond mere collecting. In any case, Egypt was hardly conducive to systematic specialized research. Almost all of the members of the Institut took up research problems well outside the areas of their expertise. In such a setting one could not help but be preoccupied with the grand sweep of civilization. Thus Geoffroy presented papers on the legends of Herodotus, on the religious customs of the Egyptians, on embalming, and on establishing the site of ancient Memphis.15 There would have been little audience for dry taxonomic or anatomical memoirs, as there were no other zoologists except Savigny. At first Geoffroy seemed concerned with the effect of this lack of professionalism on his career. On the occasion of the publication of his first memoir on the wing of the ostrich in La decade egyptienne, Geoffroy apologized to Cuvier in a letter dated 20 October 1798: You note that it is written for the army, [and] that I could not dwell on details of anatomy. In order to make the few facts that I published stand out, I have been forced to present a general tableau. Thus, forestall the objections that could be made against my publication, and please let the savants know that it was not for them that I have written.16

Further into the expedition, as will be seen, Geoffroy lost all cognizance of the requirements of his colleagues at home. Several of Geoffroy's memoirs written in Egypt presented special cases of his later doctrine of "unity of organic composition." In his memoir on the wing of the ostrich, read to the Institut d'Egypte on 7 September 1798, he reasoned from structure to function to show that the ostrich's bone structure made it impossible for the bird to fly. He pointed out that although the ostrich, being flightless, had no need of a furcula (wishbone), vestiges of the bone were nevertheless present. "These rudiments of the furcula," Geoffroy explained, "have not been suppressed, because nature never advances by rapid leaps, and she always leaves the vestiges of an organ even when it [the organ] is entirely superfluous, if that organ has played an important role in other species of the same family." On the same principle, Geoffroy suggested, vestiges of a wing could be found in the flightless cassowary, and the rudiments of the nictitating membrane, found in many birds and quadrupeds, could be seen in man. At this time, Geoffroy still regarded the furcula as "a bone that has no analogue at all

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in the quadrupeds." It was assumed to be found only in birds.17 In another early memoir presented to the Institut d'Egypte on 1 November 1798 on the frontal prolongations of ruminants, Geoffroy attempted to show that the antlers of the deer, the horns of the giraffe, and the horns of cows "are absolutely of the same nature."18 Other papers dealt with the anatomy of the flamingo and of the crocodile of the Nile, but to Geoffroy the fishes were the source of his greatest anatomical triumphs. Although he retained some interest in the classification of fishes, his chief preoccupation was to make sense out of the strange anatomical structures he found. He read papers on the appendices of sharks and rays, on the anatomy of Polypterus, a fish so extraordinary that Cuvier claimed it alone was worth the entire expedition,19 and on the inflation mechanism in Tetraodon, the puffer. To Cuvier and Lacepede he wrote extensive notes on the Siluridae and Mormyridae. In his letters to Cuvier, he suggested several intriguing anatomical comparisons. For example, in a fish he called Silurus anguillaris, he claimed to have found two ramified organs similar to lungs in man and three hearts, which he compared to the hearts in the cuttlefish, a mollusk. So satisfied was he with his work that he boasted in jest to Cuvier that he would "demand no less than the throne of anatomy."20 Geoffroy later claimed that while working up his notes on fishes for publication, he first attempted, without success, to find homologies between the skeleton of fishes and those of other vertebrates. In 1798 and 1799 Geoffroy concentrated his energies on comparative anatomy and produced some fine work which he published when he returned. In the final years of the expedition, however, Geoffroy turned to a different and broader set of problems, less productive of scientific results, but important for what they reveal of Geoffroy's fundamental ideas about God, nature, and physical law. When the Institut d'Egypte reopened in September 1800 after a long break, Geoffroy presented an extraordinary series of memoirs which took him far beyond the bounds of his previous research in Egypt. On the basis of little or no new research, he tackled a number of long-debated physiological problems, issues of manifest importance, but ones for which taxonomy and comparative anatomy offered little in the way of solution. At best Geoffroy suggested the possibility of experiments which he was neither equipped nor inclined to carry out. The first paper of this series, "Exploration of a Plan of Experiments to Arrive at the Proof of the Coexistence of the Sexes in the Germs of All Animals," read on 6 November, dealt with a significant problem, perhaps suggested by Buffon, namely the role of the environment in modifying animal structure, and in particular in modifying the sex of the developing embryo.21 Whether or not epigenesis was the true process of generation, Geoffroy believed that he could demonstrate that the sex of the embryo, at least, was not predetermined. At the moment of conception, he conjectured, both sexes were present simultaneously. In other words, every animal was initially a hermaphrodite, and only afterwards did the organs of one sex develop while those of the other sex

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remained rudimentary. In fact, Geoffroy claimed that the existence of rudimentary organs had convinced him that the germs of all organs that can be observed, for example, in the different families of animals with pulmonary respiration, exist together in all species, and that the cause of the infinite diversity of particular forms and of the existence of so many half-effaced or totally obliterated organs is due to a greater proportional development of some [organs], a development which always operates at the expense of those found nearby.22

This was an early statement of what became known as the principle of balance of organs. An organ was enlarged only at the expense of other organs that were rendered rudimentary. Geoffroy presumed that the determination of sex depended on such material factors as the quantity and quality of the nourishing fluids and the size, shape, and motion of the reproductive organs. In his next paper, "History of the Formation of the Egg," read on 21 November, he suggested that the sex of a bird might be related to the shape of the egg, the more elongated eggs containing males and the shorter eggs containing females. The shape of the egg was in turn determined by the ease of passage of the egg through the bird's oviduct. If this presumption could be proved, he asserted, then "the sex is determined in a purely mechanical manner according to the various states of the generative organs."23 To test his hypothesis, he proposed a number of experiments. Silkworm larvae might be raised under various regimes, some in confined spaces, some with minimal nourishment, and some with abundant nourishment, to determine the correlation, if any, between sex and environmental factors. Domestic animals might be dissected at various intervals after conception to see how sexual characteristics developed.24 But Geoffroy placed his greatest hopes for success on an ambitious program of experiments on bird eggs. He planned to incubate separately the long eggs and the short eggs of chickens and pigeons to determine whether the shape of the egg correlated with sex. To remove all doubts on the matter, he actually hoped to convert long eggs into short eggs and vice-versa by removing the ends of the shell and grafting on new ends. To carry out these and related experiments Geoffroy asked for an incubator, 500 to 600 chicken eggs, an enclosure for 500 to 600 chickens, 112 pairs of pigeons, two dovecotes, male and female ostriches and ostrich eggs, measuring instruments, and two Egyptian assistants.25 On 6 December a committee consisting of the physician Rene-Nicholas Desgenettes, General J.-L.-E. Reynier, and Jacques-Nicholas Conte presented a favorable, if somewhat skeptical, report on Geoffroy's lavish plans, recommending that a request be made in the name of the Institut d'Egypte to the government to provide the necessary materials. Even if the proposed experiments failed to lend support to Geoffroy's theories, the reporters concluded, the experiments promised to be useful to science.26 It is not likely that Geoffroy ever received the birds, eggs, or incubator. However, he did commence research on the effects of the environment on the determination of sex from another direction. In December 1800 he planted spinach

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and hemp seeds according to an elaborate set of diagrams, giving some plants too much water, some the optimal amount, and some too little water. Some plants were crowded together while others were given ample room to grow. By these means, Geoffroy hoped to determine the cause of the "abortion" of one of the sexes in dioecious plants, that is, plants in which the pistils are on one individual and the stamens on another. Although Geoffroy's results were bound to be inconclusive, it is probable that the members of the Institut left Cairo before the plants could be analyzed.27 Not content with the extensive plans he had already laid out, Geoffroy turned to other, and even more diverse, topics of research which he treated in a deductive manner. In a paper read on 6 December entitled "Considerations on the Muscular Fiber," he joined the ranks of physiologists of centuries past who offered speculative theories of muscular contraction.28 Yet another paper, read on 5 and 20 January 1801 and dealing with respiration, offered an explanation based on the caloric theory of heat of why some animals (e.g., birds) exhibited more vitality than other animals (e.g., fishes). In a Cartesian mechanistic fashion, he attributed the differences between hibernating and nonhibernating animals to the "greater narrowness of the aerial vesicles which terminate the branches of the bronchial system" in the former. Although he suggested that his hypothesis might be experimentally verified, he admitted that he lacked the proper injection equipment to do it himself.29 None of these papers ever saw print; his colleagues in Paris no doubt convinced Geoffroy that publication could only do his scientific reputation more harm than good. Yet Geoffroy's memoirs of November 1800 to January 1801, as far-reaching as they were, were only a prelude to the grand synthesis that he developed during his last three weeks in Egypt before embarking for France. By the time Geoffroy returned from Upper Egypt in the fall of 1799 with his valuable collection of human and animal mummies, he was weary of adventure and more than ready to go home. Bonaparte had already departed in August 1799, taking with him Monge and Berthollet. In November a coup d'etat established him as first consul of France. It appeared as though the rest of the savants would soon be leaving Egypt, for France had consented to the evacuation of Egypt after her armies were defeated by the British at El Arish in January 1800. But as events turned out, they had to endure another year and a half before being allowed to sail. From late 1799 on, Geoffroy complained of stomach trouble, frequent ophthalmia (he was blind for 29 days), lack of books, and loneliness. He feared his collections would all be lost if he did not get out of Egypt quickly. Nearly every letter to Cuvier begged a word from him and chastised him for scarcely writing.30 Pathetically, he called for Cuvier's intervention in bringing him back to Paris. "It is certain that without your help," he wrote on 9 November 1800, "I will remain here until I don't know when."31 The Institut d'Egypte held its last meeting on 21 March 1801, at which Geoffroy presented a memoir on the anatomy of the crocodile of the Nile. With the defeat of General Menou in the Battle of Canopus, the scientists were evacuated in April to Alexandria, under siege by the British. Their expected departure to France continued to be delayed.

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The most fascinating episode of Geoffroy's expedition to Egypt took place during the final weeks of the siege in August up to the time of his final embarkation for France in September. More than any other single incident in Geoffroy's career, it provides insight into the conflict between Geoffroy's synthetic and materialist vision of nature and his deep-seated need to win the approval of his colleagues. The capture of two electric fishes, the torpedo ray from the Mediterranean and the electric catfish (Malapterurus electricus) from the Nile, served as the point of departure for an extravagant and all-encompassing system of subtle fluids and forces. Never in his life, his son reported, did he work harder. Recalling the event in 1835, Geoffroy wrote that his complete absorption in his work made him totally oblivious to the commotion of the siege of Alexandria. "The bombing, the local fires, the surprises of the besiegers, and the plaintive cries of the victims"—all paled before his examination of "my problems of natural philosophy." The anatomies of his two fish led him to consider the nature of nervous action, and finally "all the phenomena of the material world:" It was necessary, in the short interval of three weeks, to pass through in my mind sixty-four times all my knowledge of science because of the sixty-four hypothetical formulas that I had to examine and compare.

"Knowledge is so sweet," Geoffroy fondly recalled, "when one has arrived at a series of deductions which appear to the mind with the character of a perfect lucidity."32 Geoffroy's ambition was no less than to explain the relationship among the major "imponderable fluids" of the universe: light, caloric, electricity, and the nervous fluid. His thought can be gleaned from several portions of his theory that survive in manuscript. Of particular interest are a memoir entitled "On Caloric Considered as the Vital Principle and on Its Laws of Circulation in Organized Beings" and a long essay pretentiously called "My Physics, or System of Ideas on the Nervous Fluid, and Secondarily on the Real Elements of Bodies."33 As inappropriate as such memoirs might appear for a naturalist, Geoffroy was not entirely isolated from the science of his contemporaries. In this lofty undertaking, he was influenced by the aspirations of the physical scientists of his day, and in particular by Laplace's program for physics. Trained as a mineralogist and in contact with physical scientists from his earliest exposure to science, Geoffroy could not help but take an interest in the outstanding physical problems of the time. In the 1830s, when Geoffroy returned to his speculations of 1801, he related on several occasions an episode that allegedly occurred on the eve of Bonaparte's departure from Egypt. In a pensive mood, Bonaparte was supposed to have declared that if he had his choice he would rather have been a Newton than an Alexander. Monge, quoting Lagrange, replied that no one could attain the glory of Newton, for there was only one world to discover. Not so, was Bonaparte's rejoinder, there was still "the world of details," the investigation of the laws governing the movement of the smallest particles of matter at very short distances.34 The elucidation of "the world of details" was in fact the ultimate goal of the

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ambitious and influential program set out by the two most prominent physical scientists of the day, Laplace and Berthollet.35 By utilizing experiment and sophisticated mathematics, they hoped to reduce the phenomena of the physical world to attractions and repulsions between matter and various imponderable fluids such as heat, light, electricity, and magnetism. Geoffroy absorbed the vision of the Laplacian program without any of the discipline required of those seeking to carry it out. He seemed largely oblivious at first to the fact that his speculative synthesis was more likely to antagonize than to impress his colleagues in Egypt.36 Geoffroy's theory was thoroughly materialistic. A benevolent Creator was no more a hypothesis of Geoffroy's universe than He was of Laplace's. Geoffroy decided that the vital fluid, previously misunderstood because of "a prejudice of our education that the vital principle ought to be of a supernatural order," was nothing more than caloric disengaged from matter in the lungs and circulated through the nervous system.37 At small distances there was no other law than Newtonian attraction, but attraction took place in a special and hitherto unrecognized manner. Geoffroy hypothesized that whenever two effluvia of caloric were directed toward each other, the "molecules" of caloric, assumed to be conical or pyramidal in form, would attract base to base according to the square of the distance between them, and by an "electric" process, lose their elasticity and become transformed into a molecule of carbon. This law of attraction at short distances, Geoffroy claimed, could be applied to a host of phenomena. In muscular action, for example, two streams of caloric, one emanating from the nerve and the other from the arteries, encountered each other in the muscle, causing the muscle to contract. In generation, the male produced one stream of caloric and the female the other. Enlarging his system to encompass other subtle fluids, Geoffroy further supposed caloric to be composed of seven generating elements (corresponding to the seven colors of the rainbow), distinguishable only by specific weight and affinity for oxygen. The seven elements together with oxygen constituted light and all other material bodies. All change occurred as the result of two opposing processes: the electric process by which generating elements lost their "expansibility" and the process of "combustion" by which they regained it. "It suffices to tell you," he wrote to Cuvier, that in accepting the general principles I have established regarding the differentiation of caloric elements according to their weight and ability to combine with oxygen, and general attraction, all the rest can be deduced in a simple manner. One can explain all galvanic, electric, and magnetic phenomena, the nervous fluid, germination, development, nutrition, generation . . . I say also that one can explain the intellectual function by physics [la physique], something that I have been told has already been done, and that 1 will dare to undertake [myself].38

At first Geoffroy hoped his system would make his scientific reputation. On first announcing his discovery to Cuvier on 26 September 1801, shortly before leaving Alexandria, he wrote: "As soon as I arrive at quarantine, I will send you a very extended work on physics, chemistry, and physiology: the discovery of the nervous fluid and the vital principle has led me to a very vast theory. I hope to re-enter France worthy of you and my illustrious colleagues. I have

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already obtained from you your love of work; I will renew myself in your company so as to gain all the other qualities still lacking in me."39 However, Geoffroy's hope for instant glory soon proved illusory. His system met with immediate resistance from his colleagues in Egypt, especially from the mathematician Fourier, permanent secretary of the Institut d'Egypte, who had little patience with Geoffroy's enthusiasms. Geoffroy naively tried to rationalize Fourier's hostility by suggesting to Cuvier that Fourier was jealous of his discovery. He explained to Cuvier on 19 December from quarantine at Marseilles shortly before his return to Paris, that ever since Berthollet left Egypt with Bonaparte, Fourier had been assuming a haughty manner around his colleagues, perhaps hoping to gain by such means a reputation to match that of Lagrange and Laplace in Paris. Geoffroy continued: When my discovery concerning the nervous fluid, caloric, light, electricity, etc. became known, there were some people who extolled it, whether by persuasion or by resentment against Fourier. The latter felt the blow brought against him; he wanted to prove that it [the discovery] was only nonsense [une niaiserie], and being unable to reply to me, he tried, seeing me so enthusiastic about my discovery, to make me pass for a fool. I was angered. I wrote. I assembled all my proofs and proposed no other revenge than to begin convincing him, and I have very certainly succeeded.40

Geoffroy was, however, not entirely sure all was well. He feared that Fourier's assent was merely politic. If, in fact, Fourier was still jealous of his discovery, he might misinform Monge and Berthollet concerning the state of Geoffroy's health and work, especially since the system had attacked one of Berthollet's chemical laws. For these reasons Geoffroy begged Cuvier to sound out Fourier and act quickly to protect his scientific reputation: Keep, my dear friend, all this babbling to yourself. Do not expose me to the ridicule of the great mountain that would give birth only to a mouse! I regret having made this recitation to you without all the facts that have authorized me in these singular opinions, yet I must ward off the unfavorable impressions that might be spread concerning me in Paris and furnish for your friendship some arms for my defense.41

Released from immersion in his new theory only by the orders to embark for France, in the final days in Egypt Geoffroy performed a heroic act for which he was long after remembered. The capitulation agreement contained a clause whereby the British would receive all the notes and collections made in Egypt by the savants. It was only after Geoffroy threatened to burn the collections rather than cede them to the British that the British command relented.42 The Rosetta Stone discovered by the French was confiscated and now rests in the British Museum, but Geoffroy's collections arrived safely in Paris. Following a long quarantine at Marseilles, Geoffroy at last rejoined his colleagues at the Museum in late January 1802. Separated from the guidance of Cuvier and Daubenton, Geoffroy in Egypt had pursued a path of his own. Although systematic zoology had been the expected and appropriate area of concentration for a naturalist exploring an unknown country, Geoffroy instead turned to morphology. In the papers on

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comparative anatomy that Geoffroy presented in Egypt, there was little of the Cuvierian concern for functional adaptation. Instead, Geoffroy focused on structure, and in particular on the assimilation of unusual forms of animals to more common forms. He had as yet only an intuitive notion of an underlying unity of plan, no real explanation for similarities of structure, and no method for making rigorous comparisons. By the end of 1799, as he became even more isolated from the concerns of his fellow professors at home, he followed his penchant for exploring subjects chosen not because they were amenable to solution given the means at hand, but because they were intrinsically significant. His theory of physics, he admitted, would require a great many observations and experiments, but he was justified in presenting it, he felt, if he could show that "the era of the solution of the most transcendent questions of physiology is not as far off as has been until now believed."43 His proposed experiments on birds, eggs, and plants in 1800 attested to a belief in an environment capable of producing significant structural modification of animals (although not necessarily transmutation of species). Implicit in his experiments was the idea that all animals were built on a similar plan in which both sexes were present, but that the environment modified the sexual organs, developing some and aborting others. In Egypt, therefore, we find the germ of much of Geoffroy's later morphological theorizing. In Geoffroy's system of elementary materials, forces, and processes, the sciences of physics, chemistry, and physiology were intertwined. Like the Enlightenment philosophes before him, he recognized no boundaries between the sciences and no limitations imposed by specialized training. Geoffroy rejected any vital force acting in opposition to the destructive action of physical and chemical forces in the animal body. Nor did he allow for the activity of a wise and benevolent Creator who preoccupied Himself with the minutiae of fashioning means to ends. The Creator of Geoffroy's imagined universe was at best a First Cause who defined the elementary forms of matter and devised the universal laws to set things going. Unable to convince his colleagues of the value of his speculative theorizing, Geoffroy was overcome by fear and caution and suppressed his work. None of the memoirs on generation or on his system of the universe ever saw print. The specific principles of his system of the universe were quietly dropped. But although he returned to writing circumscribed memoirs of comparative anatomy and zoology, as was expected of a savant in his position, he did not destroy his manuscripts and he did not abandon the overall view of nature and science that they contained. It was a view of nature and science which, if pursued, would bring him into inevitable conflict with Cuvier. Return to Paris and Circumspection

The next few years saw Geoffroy settling down as a professor in the Museum, publishing a large number of memoirs on zoology and comparative anatomy, and generally avoiding controversy. Through his publications and especially

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through the report of his colleagues at the Museum on his collections, he was able to make for himself a modest reputation based on his travels. But he was unable to achieve the level of fame and recognition that he coveted. In particular, a seat in the Institut de l'Egypte was unable to gain for him a place in the Institut in Paris. Soon after Geoffroy's return home, Lacepede, Lamarck, and Cuvier presented a highly laudatory report to the professors of the Museum on Geoffroy's collections, which raised issues that were to be of major consequence in the history of evolutionary biology. The remains of mummified animals brought back by Geoffroy from Egypt were to become one of the strongest arguments against evolution in the period before Darwin. "For a long time it was desired to know if species have changed form with the passage of time," his colleagues wrote. "Never has one been in a better position to decide with respect to a large number of remarkable species and several thousand years."44 It is impossible to recapture the original excitement of unwrapping animals—cattle, crocodiles, ichneumons, monkeys, and ibises—which had been embalmed three thousand years before and discovering them to be in no way different from present animals of the same species. With the hindsight of the evolutionary time scale, three millenia is a very short time, but in the first decade of the nineteenth century it was reasonable to conclude that if species had remained stable for so long, no significant change had occurred over any period of time. This conclusion was not, however, explicitly drawn in the report, possibly because Lamarck had a hand in its drafting. It was simply observed that "these animals are perfectly similar to those of today,"45 and that it would be interesting to obtain information on animals of an even older origin hidden in more inaccessible tombs in Egypt. But the implication was clear. If the official report was noncommittal, Lamarck and Cuvier soon took their individual stands on the evidence brought back by Geoffroy. In Philosophie zoologique (1809), Lamarck argued that the animals of Egypt had not changed because the climate of Egypt had remained stable for over three thousand years: "The birds that live there now, still found in the same environment as they were then, could not have been forced into changing their habits."46 Cuvier drew the expected antievolutionary conclusion from the same evidence in his preliminary discourse to Recherches sur les ossemens fossiles (1812). Some naturalists, he wrote, relied on "thousands of centuries which they accumulate with the stroke of a pen." But the effects of a long period of time could only be judged by multiplying the effects that a shorter time produced. Thus, he maintained that the specimens brought back from Egypt provided no foundation whatever for the belief that Paleotherium, Anoplotherium, Megalonyx, Mastodon, Pterodactylus, etc. (all new genera of fossils that he had established) were the ancestors of current animals.47 The same antievolutionary argument drawn from Geoffroy's animal mummies was to figure in Charles Lyell's Principles of Geology in 1831.48 For his part, Geoffroy pronounced no opinion on the matter until 1825, when his first memoir on paleontological evolution appeared. The controversy with Cuvier, when it came, was to deal not with evolution per se, but with philosophical anatomy, a doctrine that might be combined with evolution, but that might also be held independently of it.

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Back in Paris, Geoffroy returned to taxonomy, publishing competent memoirs on several new families of marsupials and bats and describing a number of new species from around the world, but failing in his one attempt at publishing a book dealing with taxonomy at a broader level. Lamarck produced a multivolumed classification of all invertebrates, and Cuvier in Le regne animal published a guide to classification of the entire animal kingdom, but Geoffroy's contributions to taxonomy did not deal with taxa above the level of the order (e.g., primates). As curator of mammals and birds at the Museum, Geoffroy might have been expected to publish a catalogue of these classes together with a systematic classification. In late 1802 Geoffroy did indeed begin a Catalogue of Mammals in the Museum d'Histoire Naturelle, which was to offer a new classification (or rather a revision of the earlier classification developed with Cuvier), concise descriptions, and summaries of synonymy (i.e., a list of all the names by which an organism has been called). It was to have been his first book-length publication. But he fell ill, his assistant made some errors, and in early 1803 he suddenly abandoned the entire project. Printed copies of the work without a title page were handed out to his colleagues, but the book never reached the booksellers.49 Isidore Geoffroy regarded his father's decision not to complete the work as a turning point in his career, signifying his divergence from Cuvier, his belief that classification was arbitrary, and his conviction that "true science should be sought on a broader and higher plane [plus loin et plus haut]." 50 In addition to taxonomic memoirs, Geoffroy published several anatomical papers based on materials gathered in Egypt, some of them establishing new homologies. Polypterus bichir, Silurus anguillaris, and the crocodile of the Nile were each the subject of memoirs published in the Bulletin of the Societe Philomatique or in the newly founded Annales of the Museum. The paper on Polypterus (a genus named by Geoffroy), published in the first volume of the Annales in 1802, foreshadowed some of Geoffroy's later theorizing on fishes. He noted homologies between limbs in mammals and the unusual pectoral fins in this fish, remarked that the omoplate, clavicle, humerus, and sternum were separate bones in the young but then sutured together, and found examples of rudimentary organs as well as other organs enlarged to compensate for the rudimentary organs.51 From the perspective of his later morphological work, however, his most interesting memoir of 1802-1803 dealt with the electric organs of the torpedo and other electric fishes. In this paper, also published in the first volume of the Annales, Geoffroy posed the question of why only the torpedo, among all rays, had the power of electrocuting other fishes. Geoffroy was certain that a homologue of the highly complex electric apparatus of the torpedo ray had to exist in other rays, for, he explained, "It is not necessary to have compared many animals to learn that there are never any new organs among them, especially in species which resemble each other as much as the rays. It was more natural to believe that the tubes enclosing a gelatinous substance in the torpedo existed masked in the other rays." By dissecting other rays, he was indeed able to identify "an analogous organization, with differences that should correspond to the manners of existence and action of each species."52

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Despite a large output of publications, Geoffroy's career was only moderately successful, as demonstrated to him by his repeated failure to gain a seat in the Institut. Ever since 1796 Geoffroy had been reading papers at the Institut and awaiting positive reports from the commissions of members chosen to review the papers presented by nonmembers.53 There were, however, few opportunities to become a member of the Institut, for suitable vacancies normally depended on the death of one of the six members of the section of anatomy and zoology. In May 1798, when Geoffroy was on his way to Egypt, he was nominated for a vacancy among the nonresidents where competition was less stiff, but he was declared ineligible and his name was struck from the list of candidates. Although on a voyage, he was still considered a resident of Paris because he retained his place as a professor of the Museum.54 In 1800 Daubenton's death created the first opening in the section of anatomy and zoology since the section was set up in 1795. Despite a favorable report by Cuvier and Lacepede on several of his unpublished memoirs on fishes of the Nile, Geoffroy, away in Egypt, placed only sixth among eleven candidates. In the final voting, the Linnaean naturalist Olivier was elected.55 When the next vacancy occurred in 1803 upon Cuvier's being made permanent secretary, Geoffroy waged an active campaign but lost once again. The two men who carried the most weight in deciding the outcome, Geoffroy realized, were Laplace and the Emperor himself. Geoffroy reported to Cuvier, who was away inspecting schools, on 8 February 1803, that, with the help of Lacepede, Laplace and Napoleon were convinced that a zoologist should be elected rather than a physician-anatomist. Laplace had prepared a list of three candidates: Dumeril, Brongniart, and Geoffroy. Geoffroy begged Cuvier to intervene with the patriarch of the physical sciences. "Please my friend," he wrote, "write a word to him. Encourage him to hold to his list and say something helpful on my behalf."56 Despite Geoffroy's plea, additional nonzoologist candidates were in fact nominated. He lost to the physician Philippe Pinel, twenty-six votes to thirty. Pinel had only slight qualifications as an anatomist or a zoologist and would have made a more appropriate member of the section of medicine and surgery, but competition in that section was so fierce that his friends had to make use of whatever vacancy was available.57 Thus, by 1803, ten years after he became professor of zoology at the Museum, Geoffroy had achieved only a modest reputation as a taxonomist and comparative anatomist. He had not yet published a book, and was still not admitted to the tribunal of French science, the Academie des Sciences. Recognition, when it finally came to Geoffroy in 1807 and 1808, was in good part the result of Geoffroy's identification of a research program which he could call his own. That was the novel program, later known as philosophical anatomy, of searching for homologies among the skeletons of the four classes of vertebrates. Foundation of a Research Program: Memoirs of 1807

In 1807, in a series of memoirs presented to the Institut and published in the Annales of the Museum, Geoffroy opened a new direction for research in com-

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parative anatomy. Whereas traditional comparative anatomy dealt with the description and the function of organs, Geoffroy posed an entirely new set of questions to anatomists and zoologists. His research program was morphological, and centered on a new concept, that of homology. He proposed to demonstrate not only that the four classes of vertebrates shared a general plan of organization (as was generally assumed), but also that they were susceptible of very specific kinds of comparison. Through the determination of "analogous" (homologous) parts, the four classes of vertebrates could, in fact, be reduced part by part to a common plan of organization. Underlying the diversity of animal organizations, a hitherto unexpected identity would emerge. Specifically, Geoffroy's goal in these pioneering memoirs was to show that despite obvious differences in appearance and milieu, the "same" structures were present in the fishes as were found in reptiles, birds, and mammals. The skeletal system was the focus of his efforts, for it was not only the easiest system to analyze, but also (as it appeared to Geoffroy) the framework for all the other systems of organs. For the first time, osseous structures in fishes were systematically analyzed into individual bones and compared bone for bone to structures in other vertebrates. Instead of function determining structure, Geoffroy brought to light the disquieting possibility that homologous bones in different animals could be modified to perform entirely disparate functions. Bones could henceforth be perceived in abstraction as "materials of organization" that in different animals could be developed, atrophied, modified, sutured together, or even annihilated. The guide for determining homologies, Geoffroy had begun by 1807 to recognize, was neither shape nor function, but the connections of the parts. Geoffroy also quickly seized upon another methodological precept of morphological anatomy, namely that the fetus provided a more reliable guide to the determination of homologies than the adult. If homologies could not be made evident by the examination of adult organisms, unity of plan might yet be revealed by a study of the bones of the fetus. Although the "unity of composition," the "theory of analogues," and the "principle of connections" were not mentioned by name, Geoffroy's three memoirs on the skeleton of fishes and two memoirs on the skull of vertebrates contained most of the theory and method of the later Philosophie anatomique. The immediate impetus for Geoffroy's program of research was no doubt the publication of Cuvier's five-volume Lecons d'anatomie comparee (18001805), the first work offering a systematic and detailed comparison of the four classes of vertebrates. To someone like Geoffroy with a presentiment of the unity of plan underlying the structure of vertebrates, Cuvier's Lecons were disappointing and inconsistent. Not at all concerned with naming individual bones or with identifying structural affinities among the classes of vertebrates, Cuvier's aim was rather to exhibit the different structural means by which animals performed the same function. Thus his volumes were organized by functional system, such as respiration, circulation, digestion, and movement. In his treatment of the skeleton of fishes, Cuvier relegated the pectoral fin to a separate section from the anterior extremities of other vertebrates, and made no effort to describe the bones in detail or to compare them to bones in other vertebrates. Most of the bones of the pectoral fin were lumped together as "os

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en ceinture." However, Cuvier did name a clavicle and a scapula (shoulderblade) in fishes, his determinations differing from both those of his predecessor, the Linnaean naturalist Antoine Gouan, and those of Lacepede. When bones in one class of vertebrates appeared similar and performed a similar function, Cuvier usually carried over the name from one class to the other. But when the bones in one class of vertebrates appeared sufficiently different in form and function from those in another class, Cuvier, without giving justification, used a different name. All this must have appeared to Geoffroy quite arbitrary. In any case, when Geoffroy began his investigation of homologies among the vertebrates, he deliberately set out to show that the parts which Cuvier assumed were present in only one class were actually present in all four classes. Thus, in reaction to Cuvier, he looked for a furcula (wishbone in birds) in fishes and for the bones of the operculum (gill-cover of fishes) in mammals.58 When, shortly before his first memoir on fishes, Geoffroy set out to describe the skeletons of his Egyptian fishes for the monumental Description de l'Egypte to be published by the Commission of Sciences and Arts, he began to transform his as yet vague Buffonian insights on the unity of plan among vertebrates into a new understanding of animal organization.59 Recounting his career in 1830, he recalled that he was faced with determining a name for a large bone which played an important role in the inflation mechanism of Tetraodon (the puffer). Instead of inventing a name for the piece, Geoffroy's predilection for unity of plan in nature led him to seek its homologue. He concluded that the bone in question was equivalent to the coracoid process, a part of the scapula in man, and hence gave to it the name coracoid. With this success, Geoffroy passed on to the neighboring bones of the shoulder and arm. "And it is from this epoch," he claimed, "that my first researches on the philosophical resemblances of organs date."60 The bones of the arm and shoulder in fishes formed the subject of Geoffroy's "First Memoir on the Fishes," written in late 1806 and read to the Institut on 2 March 1807. Aristotle had long ago suggested that the pectoral fin of fishes was a piscine equivalent of an arm, but Aristotle had not specified the similarities in further detail. No one before Geoffroy had thought it possible to formulate an exact equivalence of bones.61 The comparison was far from obvious, because, as Geoffroy explained, the bones in fishes were differently situated than in the other vertebrates. In fishes, the bones of the arm and shoulder girdle were to be found behind the thoracic cavity, but at the same time articulated to the skull. Geoffroy managed to determine a clavicle, scapula, humerus, radius, ulna, carpals, and metacarpals. "I have had in fact the satisfaction of discovering that even the organs which had most consistently resisted all comparison found their analogues in the other vertebrates," he boasted.62 Among the "bones of the shoulder" in fishes, Geoffroy identified not only a clavicle and scapula but also, surprisingly, a furcula, a bone hitherto assumed to be restricted to birds. Moreover, he argued the radical thesis that the bone changed its function as it passed from one class to another. In birds the furcula played a significant role in flight. In fishes it lost this function, but it was retained because of "the tendency of nature to cause the same organs contin-

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ually to appear in [all] animals formed on the same type."63 Thus, Geoffroy implied, the furcula was not a bone specifically designed by the Creator to aid birds in flight, but rather an abstract element of organization which could serve multiple functions as it was placed in different circumstances: Without a direct object in swimming animals, without a utility determined in advance, and thrown, so to say, by chance into the field of organization, the furcula enters into connection with the organs near it; and according to the manner in which this association is formed, it takes on uses which are in some sense prescribed by them.64

Geoffroy observed that some fishes had no furculas, that in other cases the furcula acted as an additional rib, and that in anglers it aided in opening the gills. Thus the furcula, when not needed for flying, either disappeared or took on a wholly unrelated function. In his second memoir on the fishes, read to the Institut on 23 March 1807, Geoffroy pursued the subject further, considering the various "metamorphoses" of form and use of the furcula in the families of fishes.65 In his third memoir on fishes, read on 27 April and 11 May 1807, Geoffroy identified homologues of the sternum in the fishes. The eighteenth century anatomist Joseph-Guichard Duverney, Vicq d'Azyr, and Cuvier, among others, had already attempted to identify a piscine sternum, but, according to Geoffroy, there had been no agreement thus far, because most of the bones situated between the abdominal viscera and the jawbone in fishes had not yet been reduced to their true "analogues." The bones in this region, he claimed, had hitherto been examined and described in isolation, and given names such as operculum and branchiostegal rays which implied that they were part of "an entirely novel organization, uniquely applicable to the class of fishes." To find the homologues, Geoffroy argued that it was first necessary to take into account the modifications in the structure of fishes due to differences in respiratory milieu. Because the fishes lived in a watery environment, the form of their parts had been modified and their relative position changed, hence making the parts difficult to recognize. Although the anterior extremity in fishes "was transposed and carried back from the front to the back of the thoracic cavity [cavite pectorale]," the sternum was "transported" along with the gills to a point anterior to the "arms" and under either the cervical vertebrae or the head. Geoffroy located a sternum and sternal annexes in a bony apparatus under the gills and maintained that the branchiostegal rays, spines which supported the membrane covering the gills, were the piscine equivalent of sternal ribs. He showed how by an appropriate transformation the sternum of a bird might become the sternum of a fish.66 Geoffroy sought confirmation of his homologies by identifying anomalous species to serve as transitional forms and by counting centers of ossification in the fetus. For example, to support his homology of the single furcula in birds with the two supposed homologous bones in fishes, he exhibited several anomalous fishes in which the two bones were prolonged, inclined toward each other, and sutured together as in birds. Likewise, he found cases of birds in

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which the wishbone was separated into two pieces as in fishes. The homology was further corroborated by an appeal to the avian fetus. In the fetus of birds, he could point to two primitive bones that during the process of development sutured to form the wishbone. Thus, while the "furcula" consisted of one bone in most birds and two bones in most fishes, the homology could be demonstrated by a series of intermediary forms. In a similar manner, Geoffroy found traces of five separate bones in the sternum of avian fetuses to correspond to the five separate bones of his proposed sternum in fishes.67 Geoffroy's language, and that of other philosophical anatomists, was as vague and confusing to contemporaries as it often seems to historians. It is uncertain what was to be understood by words such as "modified" and "transposed." At times, indeed, Geoffroy sounded nearly Lamarckian. However, Geoffroy intended homologies to be interpreted in ideal terms rather than in real and physical terms. Geoffroy appears to have envisioned an abstract common plan of vertebrate structure whose elementary parts could be minimally or maximally developed, and modified as needed to serve different functions in different environments. This common plan had a prior ontological status to the modifications of the plan for particular purposes. One set of bones still eluded Geoffroy in his project of reducing all anomalous bones in fishes to their homologues in the higher animals. These were the bones of the operculum or gill-cover, the bones which, to Geoffroy, most clearly marked the difference between fishes and vertebrates with pulmonary respiration. If the Creator were to create any bones especially for fishes, they would be the opercular bones. Thus to demonstrate a unity of plan among the vertebrates, it was essential for Geoffroy to identify the homologues of these bones. He suspected that the operculum corresponded to bones which formed part of the skull in other vertebrates, in particular to the parietals. In order to pursue this idea, Geoffroy undertook a general study of the bones of the skull in the four classes of vertebrates. On 17 August 1807 he read to the Institut a memoir on the skull of vertebrates, with special reference to birds, and in 1807 also he published a detailed analysis of the bones of the head of the crocodile, presumably as a representative of the reptiles.68 Although unproductive in identifying homologues of the operculum, the study of the skull led Geoffroy to a number of other unexpected results. He claimed that the skull of birds was composed of the same number of pieces as in mammals. In particular, he formed a homology between the quadrate bone in birds (an important mobile bone which joins the lower jaw to the skull), and the tympanal bone (a bone in the skull which partially encloses the tympanic membrane) and styloid process of the temporal in mammals. More significantly, he also claimed an equivalence in the number of bones in the skull of mammals and fishes. Colleagues, and Cuvier especially, were extraordinarily impressed with Geoffroy's ingenious method of demonstrating this equivalence by counting centers of ossification in the fetuses of mammals. Fishes had many more bones in their skull than adult mammals, but Geoffroy was able to find the additional bones still separated in mammalian fetuses. His justification for the procedure foreshadowed his later controversial theory of "arrests

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of development." Fishes, he claimed, had arrived at the same stage of development as the mammalian fetus.69 It was in working with the bones of the skull that Geoffroy began to recognize that the connections of the bones to each other formed the most reliable guide to establishing homologies: Nature, I have said above, tends to cause the same organs to reappear in the same number and in the same relations, and it varies to an infinite degree only their form. According to this principle, in the determination of the bones of the head of fishes, I will never have to decide according to consideration of their form, but only according to that of their connections.70

Tracing of connections did indeed become the chief means by which nineteenth century morphologists identified homologies. As Geoffroy himself later argued, neither function nor form ought to be considered in determining homologies, but solely the relations of the parts to each other. Yet all this effort still did not solve the problem of the operculum of fishes. Geoffroy could only reiterate his predetermined conviction that the operculum ought to have homologues in the other vertebrates. The alternative, the commonly held belief that nature had so completely modified the organs of fishes to enable them to live in water "that there remains of the primitive plan only a few scattered and masked traces," could provide "nothing of satisfaction": It is known that nature works constantly with the same materials. She is ingenious to vary only the forms. As if, in fact, she were restricted to the [same] primitive ideas, one sees her tend always to cause the same elements to reappear, in the same number, in the same circumstances, and with the same connections.71

Unfortunately, even connections were of no help in the case of the operculum, for each of the bones of the skull of mammals was surrounded by several other bones, while each of the opercular bones had one of its edges free. Having failed at discovering the exact homologues of the bones of the operculum, Geoffroy contented himself with publishing only his general idea that the bones were the equivalent of parietal bones in other vertebrates. In fishes the frontal bone articulated directly with the occipital, and the parietals, on this supposition "of no use to the cerebral cavity," were "transformed into the opercular bones" and took on "functions related to the mechanism of respiration."72 Although Geoffroy pursued the determination of the operculum several more years, by 1812, according to his recollection, he finally despaired of a satisfactory solution and put the matter aside. His sudden successful resolution of the problem in 1817 led him to write the celebrated Philosophie anatomique. Historians have offered several conflicting sources for Geoffroy's concept of homology. Isidore Geoffroy suggested that philosophical anatomy was his father's replacement for Bonnet's chain of being, which he had only reluctantly rejected. Still seeking a single law to account for variability in nature, Geoffroy filled the void left by the demise of the chain with his theory of unity of composition.73 Isidore's suggestion, if it has merit at all, can only be true in the most general sense. An even less likely source of Geoffroy's thought, suggested by the German

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historian J.H.F. Kohlbrugge and others, was German Naturphilosophie, transmitted to Geoffroy through Cuvier.74 Although Cuvier may have had some familiarity with the work of Blumenbach and Kielmeyer, he was unlikely to have had much contact with the romantic tradition of Naturphilosophie associated with men like Schelling and Goethe. Goethe's morphological writings, although some of them dated from the 1780s, were little known in Germany, let alone France, before the 1820s, when Naturphilosophie reached its full flower. Philosophical anatomy in France and German Naturphilosophie were largely independent traditions, which later merged to some extent, but only after 1818. Geoffroy's concept of homology is more likely to have derived from French antecedents.75 More recent historians, most notably Franck Bourdier and F. Dagognet, have stressed the parallel with Hauy's crystallography. Hauy's theory, so the argument goes, was essentially a morphological theory which reduced the chaotic variety of crystals to a few basic types. Geoffroy sought a similar order in zoology. The comparison is intriguing, and in a general sense, is probably justified, but there is no evidence for other than an indirect debt. Geoffroy, like his mentor Hauy and also like Lavoisier, was seeking a theory whereby a variety of beings could be reduced to combinations of elements, in Hauy's case elementary crystals and in Lavoisier's, chemical elements.76 The most direct and immediate stimulus to philosophical anatomy was near to hand, namely the writings of Buffon, Daubenton, and Vicq d'Azyr. From them Geoffroy derived a general notion of a plan of structure in nature, which he then transformed in his 1807 memoirs into a fruitful methodology and program of research. The Reception and Spread of Philosophical Anatomy

Geoffroy's papers on philosophical anatomy met with a warm reception, even from Cuvier, who reported upon them to the Academie. Cuvier's approval— although moderated in tone—ensured that when the next seat became available Geoffroy would become a member of the Institut. Other accomplishments soon followed. By far the most important for Geoffroy's future career was his appointment in 1808 as professor of zoology at the Imperial University. This second post, held concurrently with the Museum professorship, gave Geoffroy added income and, moreover, the welcome opportunity of teaching broadly conceived courses to a group of enthusiastic young students. During the next decade, while Geoffroy himself published mainly taxonomic memoirs, others, clearly inspired by Geoffroy's success, extended philosophical anatomy beyond the skeletons of vertebrates to encompass other animals and other systems. Even before the publication of Philosophie anatomique, morphological studies had become a familiar feature of French zoology. As Geoffroy was not yet a member of the Institut, the papers he read in 1807 were each submitted to an examination by a commission of three members who reported upon them at later meetings. Several members of the Institut—

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the elderly anatomist and surgeon Jacques Rene Tenon, the botanist and invertebrate zoologist Louis Augustin Guillaume Bosc (recently elected to the section of rural economy), Lacepede, and even Lamarck—participated in the three commissions on Geoffroy's memoirs, but in each case it was Cuvier who wrote and delivered the reports.77 This in itself was significant, for Cuvier's praise counted most strongly. Cuvier, in his reports, appeared to be almost entirely in sympathy with Geoffroy's methodology and objectives. Not only did he judge Geoffroy's approach eminently philosophical, but he declared that his "analogies" appeared "plausible" and had not been seized before him in the same manner. Cuvier made special note of "the importance for philosophical natural history of these striking relations [rapports] of structure which extend sometimes over one class and sometimes even over all four classes together of the vertebrates." All of Geoffroy's appeals to anomalous cases to justify homologies were described with approval, but Cuvier greeted with most enthusiasm Geoffroy's method of counting centers of ossification in the fetus. "These unions and separations [of pieces]," Cuvier readily admitted, "are therefore only the effects of age." In the "primitive plan of nature or in the original directing of the nutrition of parts," he explained, there existed the same number of points of ossification.78 Research such as Geoffroy's on structural constancy independent of function, he went so far as to suggest, might even one day shed light on that most obscure of subjects, the origin of organized beings: It is perhaps in the constancy of these analogies that one ought to seek the first flashes [of insight]. This constancy is, in fact, independent of all use, and consequently, it results solely from the forces which have determined the formation of the organic body. One is thus led to suppose that this determination is due, at least in part, to some common principle. But we are not going to engage the Class any further in this order of reflections.79

It is possible, however, to detect in Cuvier's reports a note of hesitancy (as seen in the quotation above), reflecting, perhaps, as much Cuvier's wish to present Geoffroy to his colleagues in a favorable light as his personal misgivings. This hesitancy was manifested first by a reluctance to claim any originality for Geoffroy's method, and second by an emphasis on Geoffroy's empirical results at the expense of his theory. Indeed, Cuvier recognized no essential novelty in Geoffroy's approach. Evidence for a "common plan" among the vertebrates, Cuvier claimed in his first report, had been sought as far back as Aristotle. "For a long time," he observed, "anatomists have observed that the vertebrates seem constructed on a common plan, and that most of their differences are due only to some variation in the number or respective proportions of the pieces that compose each of their organs."80 Geoffroy himself had not insisted in his memoirs on credit for the discovery of a new doctrine or a new method in anatomy. A reader of his papers might legitimately conclude that by assuming a priori that structures in fishes were homologous to structures in other vertebrates, Geoffroy had merely discovered a few more affinities than his predecessors. Cuvier's reports tended to praise the newly observed "facts," while deliber-

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ately sidestepping the implicitly raised issue of whether one could reduce all bones in fishes to bones in mammals. Although Cuvier described all of Geoffroy's arguments regarding the furcula sympathetically, he nonetheless seemed concerned lest his colleagues might find the homologies a bit speculative, for he concluded his second report with a caveat: "In any case, the interest of M. Geoffroy's memoir is not at all limited to these affinities. He gives a positive description of the osseous apparatus of respiration in various fishes and thus furnishes to comparative anatomy new data independent of all opinion."81 In his final report on 7 September, while acknowledging that Geoffroy's memoir "could lead to yet more elevated views in the general theory of organization," he concluded with the reminder that Geoffroy "has moreover the uncontested merit of presenting numerous facts to which anatomists until now have given little attention."82 Soon after Cuvier pronounced these words, at the same meeting of the Institut, candidates were named for a vacant seat in the section of anatomy and zoology, Geoffroy among them.83 The death of Broussonet gave Geoffroy his first chance since 1803 to become a member of the Institut, and this time he, with Cuvier's support, had prepared the ground well. Geoffroy's main rival, Dumeril, had also been busy presenting memoirs at the Institut, reported upon by Tenon and Pinel. Yet Geoffroy emerged the victor in the election held on 14 September, obtaining an absolute majority on the first ballot.84 Cuvier was supposed to have congratulated him by saying, "I am all the more happy because I reproached myself for occupying a place that was due to you." Geoffroy modestly demurred.85 Soon after his victory at the Institut, Geoffroy's career was given a further boost by two new opportunities: an imperial commission to visit the natural history collections of Portugal, then under French control, and a professorship at the newly created Faculte des Sciences of the Imperial University. Geoffroy spent the greater part of 1808 in Portugal and Spain on a mission as dangerous as it was prestigious. For three days he was held in a Spanish jail during an insurrection against French control of the country. Although Geoffroy had the power to demand specimens from the Portuguese collections, he won the admiration of the Portuguese and of his colleagues at home by trading Museum duplicates for voluntary gifts.86 When Geoffroy returned to Paris he was offered the professorship of zoology at the Faculte des Sciences in accordance with the decree of 1808 organizing the University, which stipulated that the professor be selected from the professors of zoology at the Museum. According to Isidore Geoffroy, Geoffroy was at first reluctant to accept because he felt the position ought to be given to Lamarck, his senior at the Museum. Lamarck, however, was said to have refused Geoffroy's offer of the position, because at sixty-five he felt himself too old to undertake the additional studies necessary to teach a general course of zoology and comparative anatomy.87 For Geoffroy, a member of the Faculte from 1809 until his death in 1844, the general nature of the course to be taught was a decided advantage. The courses at the Museum, attended by medical students and specialized scholars, were narrowly defined by the subject of the chair, in Geoffroy's case, mammals

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and birds. At the Faculte, where a large part of the auditors were students at the Ecole Normale, he was free to teach whatever he chose in the much broader field of zoology and comparative anatomy. He was given a platform from which he could try out new ideas before a relatively uncritical group of enthusiastic young men. His controversial homologies between insects and vertebrates, for example, were advanced for the first time in a course on the structure of the vertebra at the Faculte des Sciences in 1820. J.-B. Dumas, dean of the Faculte, recalled at Geoffroy's funeral in 1844 the essential role that Geoffroy's teaching at the Faculte played in fostering philosophical anatomy. With the support of his students, Dumas declared, Geoffroy's thinking "was launched more freely into the area of abstraction. . . . Until then anatomical philosophy such as he conceived it did not exist. It is with us, for us, I will even say by us that he has developed this doctrine."88 The intellectual stimulation Geoffroy derived from his teaching was reflected in his exuberant conclusion to the opening discourse of Philosophie anatomique: Oh, that I might learn that they [my researches] have been useful to the youth in our schools! What class in all of our fair France is more worthy of interest! Such devotion, such effort, such ardor for study! Amiable youth, entirely occupied by the noble work of the mind, you seem absorbed in a single thought—in the thought that led Virgil to say: Felix qui potuit rerum cognoscere causas!89

Even before Geoffroy synthesized his researches on philosophical anatomy by the publication of Philosophie anatomique, other French naturalists began to extend his concepts and methodology. Although Geoffroy was rarely cited, all the writers of memoirs of philosophical anatomy were well aware of his work. Of all this research the most provocative was the application of philosophical anatomy by Savigny and Blainville to the exoskeletons of articulates, that is, the insects, spiders, and Crustacea. Others found examples of homologies in the digestive system of insects, and in the nervous, muscular, olfactory, reproductive, and integumentary systems of vertebrates, while Augustin-Pyramus de Candolle inaugurated a parallel school of "philosophical botany." Many of these workers, like Geoffroy, compared parts among different animals, while others compared parts within the same animal (a comparison later to be known as serial homology). Between 1807 and 1818, philosophical anatomy with its search for homologies and symmetries threatened to displace the more traditional functional anatomy and to become the most exciting and challenging branch of comparative anatomy. Everyone, even Cuvier and his disciples, dabbled in it. One of Geoffroy's earliest followers was his companion on the expedition to Egypt, Jules-Cesar Savigny. Savigny claimed to have been led into philosophical anatomy as a by-product of his effort to classify insects gathered in Egypt according to the system then in use, that of Johann Christian Fabricius, which employed characters taken from mouthparts. The main work on which Savigny's reputation as a theorist rests is a small volume, Memoires sur les animaux sans vertebres (1816), containing two papers on the skeleton of the articulates which he read to the Academie in 1814 and 1815. Inspired by Geoffroy,

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although he did not mention him by name, Savigny sought in these papers homologies between the mouthparts in the various orders of articulates.90 In his first memoir, Savigny showed that the flexible coiled tubule of butterflies used for sucking nectar from flowers was composed of the same pieces as the horny mouthparts of the caterpillar employed for the very different function of chewing leaves. Cuvier had not found a trace of the mouthparts of the caterpillar in butterflies, but the difference between butterfly and caterpillar, Savigny claimed, was more apparent than real. The tubule of the butterfly was in fact simply a modification or transformation of the maxillae of the caterpillar. Following Geoffroy's procedure for formulating homologies, Savigny named individual pieces of the mouthparts, and demonstrated in detail by means of illustrations that the same parts—the labrum, labium, mandibles, and maxillae—could be found in all the major orders of insects.91 His second memoir extended these results to arachnids, myriapods, and Crustacea. Here matters were more complicated, because, as Savigny observed, the mouths of many articulates were provided with auxiliary pieces which had no "analogues" in the heads of Hexapoda (insects, proper). However, Savigny maintained that the pieces could be found instead in the thorax and legs of Hexapoda. In Scolopendra (a genus of myriapods) the auxiliary lips were merely a modification of the first pair of legs of insects, while in the crabs, all six legs of insects had been transformed into jaws! "This power that nature possesses of transforming organs of locomotion into organs of mastication," now fully accepted, was indeed to contemporaries an astonishing idea.92 Despite their novelty, Savigny's conclusions were in general well received and helped him to be elected to the Academie des Sciences in 1821. Suffering from ill health most of his long life, Savigny took little part in scientific affairs after 1817 and never held a teaching post. His two memoirs of philosophical anatomy, however, opened an exciting new field for study and led several other naturalists, including Audouin, Latreille, and Milne Edwards, as well as Geoffroy himself, to continue the search for homologies in the exoskeleton of articulates. Another early advocate of philosophical anatomy was Blainville. Contesting Savigny's priority, Blainville claimed that he had taught since 1814 that in all orders of insects the mouth was composed of the same parts modified according to function.93 In several short memoirs on insects written from 1817 to 1820 he asserted that all insects were formed on a common plan. In effect, Blainville posited an archetype for insects—an ideal plan which was obscured in particular insects by the fusion or loss of parts. The ideal insect, Blainville claimed, was constructed of fourteen identical segments, the first of which was modified to form the head. Associated with each segment was a pair of appendages which could be modified according to need to form mouthparts, respiratory organs, or organs of locomotion.94 Blainville also sought homologies among the vertebrates. In response to Geoffroy's 1807 papers, he looked for homologues of the opercular bones in the higher vertebrates, and in a memoir read in 1812 but not published until 1817, he believed he had found them in a part of the lower jaw. In other mem-

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oirs, he found the equivalent of nails or claws of mammals in the wings of birds, discovered olfactory nerves in the dolphin, which was assumed to lack them because it had no sense of smell, and demonstrated that throughout the animal kingdom, male and female organs of generation were essentially identical.95 A good friend of Geoffroy and yet another early proponent of philosophical anatomy, the physiologist Henri Dutrochet also found homologies in both insects and vertebrates. In a memoir of 1818 Dutrochet traced the alimentary canal from esophagus to anus in the various orders of insects and showed that it was always composed of the same parts. In particular, he sought to demonstrate that "the alimentary canal of perfect insects, however different it is from that of their larvae, is, however, only the same canal modified in diverse ways and adapted to the nature of the new food of which the insect is to make use. These observations accord with those of M. Savigny, who had discovered that the organs of the mouth of the butterfly are nothing but the organs of the mouth of the caterpillar rendered almost unrecognizable by their changes of form and dimension."96 In another paper in 1819 Dutrochet demonstrated the essential identity of the quills of the porcupine, the hair of mammals, and the feathers of birds.97 Primarily concerned with physiological problems outside the scope of taxonomy and comparative anatomy, Dutrochet wrote only a few memoirs from the standpoint of philosophical anatomy, but became an ardent supporter of Geoffroy in his battle with Cuvier. Independently wealthy, Dutrochet carried on most of his research away from Paris on his family estate. In that respect he was an anomaly among French scientists who achieved celebrity, most of whom held paid teaching positions in the capital. Geoffroy, with whom Dutrochet maintained close ties, acted as his agent in dealings with the Academie des Sciences and warmly supported his election to the Academie in the section of rural economy in 1831.98 Even Cuvier's acknowledged disciples were willing in the period before 1820 to venture into research in philosophical anatomy. But, as the case of Dumeril illustrates, philosophical anatomy when pushed too far could leave one vulnerable to severe criticism. Dumeril, editor of part of Cuvier's Lecons d'anatomie comparee, suppleant to Lacepede, and worker in Cuvier's laboratory, was one of the earliest naturalists to advocate the vertebral theory of the skull, a theory more commonly associated with German Naturphilosophie. In a paper read at the Institut in 1808, Dumeril sought to demonstrate that all the vertebrae of the vertebral column and their associated nerves and muscles were "analogous" in structure. This part of the memoir might have raised little objection. But Dumeril went further, suggesting that the skull was a single inflated vertebra, united to other vertebrae by the same muscles as united the vertebrae of the vertebral column to each other. According to Geoffroy, he was roundly criticized by his colleagues, who poked fun at "the thinking vertebra of M. Dumeril." From the manner in which Dumeril altered his paper before publishing it in the Magasin encydopedique, Geoffroy concluded "that one could see that he would not persevere in the same route."99

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Parallel to philosophical anatomy, there arose in France a school of "philosophical botany," which adapted many of the same explanatory devices to the anatomy of plants as Geoffroy had employed in the anatomy of animals. Its founder, Augustin-Pyramus de Candolle, had resided in Paris and was in close contact with the professors at the Museum and Institut before leaving in 1808 to take a post as professor of botany at Montpellier. Candolle's classic work Theorie elementaire de botanique (1813) demonstrated how in a natural system of classification anomalous plants could be reduced to the common plan of their order or family by morphological considerations. Thus, plants which at first glance did not meet the criteria of a particular taxonomic group (defined, for example, by a particular number of stamens or petals) could be shown to possess the required characters in an obscured form. Candolle and his many followers—among them Michel-Felix Dunal, Auguste de Saint-Hilaire, LouisMarie Aubert Dupetit-Thouars, P. J. F. Turpin, and Alfred Moquin-Tandon— called upon such explanatory devices as metamorphoses of parts (e.g., leaves to petals), abortions, adherences, balance of organs, and monstrous duplications of parts to reduce anomalous plants to a unity of plan. From the beginning subordinate to the program of establishing a natural system of classification, "philosophical botany" never became as controversial as "philosophical anatomy." But because of the similarity of concepts, several of the leaders of this school, particularly Moquin-Tandon, were supportive of Geoffroy when he came into conflict with Cuvier.100 Even Cuvier tried his hand at philosophical anatomy in a paper in 1812 which reviewed the problem of determining the bones of the skull in vertebrates. In this, his only excursion into philosophical anatomy, Cuvier once again praised Geoffroy for his work on the skull "which offers very ingenious researches and very happy results." By taking the fetus of mammals as an object of comparison, Geoffroy "has thus managed to reduce to a common law conformations which at first glance could be judged extremely diverse." While enthusiastically adopting Geoffroy's views on the "metamorphoses" of the temporal and maxillaries, he still disagreed with Geoffroy on the frontal, ethmoid, and sphenoid. But Cuvier was not wholly committed to Geoffroy's goal of finding a common plan, nor had he abandoned his belief that special bones could be created for one class of animals. Geoffroy later took exception to Cuvier's claim in this 1812 paper that the frontal of mammals included additional bones not found in other vertebrates.101 From 1807 to 1817, Geoffroy himself wrote almost nothing on philosophical anatomy. Most of his research in this period was taxonomic; in 1812 he offered a new classification of primates, and from 1810 to 1813 he published several papers on new families of bats. These years were also occupied with raising a family. In 1804 he had married Pauline Briere de Mondetour, who bore him a son, Isidore, and twin daughters, Stephanie and Anais. The next few years after Geoffroy began teaching at the Faculte des Sciences were, according to Isidore Geoffroy, the most peaceful of his life, and were disrupted only by the political events leading to the fall of Napoleon and Geoffroy's one entry into politics as a representative of Etampes during the Hundred Days, Napoleon's brief return

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to power in 1815.102 It was not until 1817 that Geoffroy turned once more to the anatomy of fishes. Geoffroy's Master Work: Philosophie Anatomique

Geoffroy's return to philosophical anatomy was brought about by the old problem of the homologies of the opercular bones. In 1807 he had discovered the homologues in fishes of the bones of the shoulder, the arm, and the sternum, but he was still confounded by the most challenging bones of all, the bones of the gill-cover or operculum. Cuvier had named four bones of the opercular apparatus in fishes—the operculum, preoperculum, suboperculum, and interoperculum—implicitly assuming that these bones were only to be found in fishes. If fishes were ever to be reduced to the same plan as other vertebrates, it was incumbent on Geoffroy to find the homologues of these bones. In 1807 he believed he could find them among the bones of the skull in mammals, reptiles, and birds. However, he worked in vain from 1809 to 1812 trying to establish the correspondences in detail. Cuvier's single foray into philosophical anatomy in 1812 put a damper on Geoffroy's effort. The important result of Cuvier's study of the skull, as far as Geoffroy was concerned, was his discovery that the skull of fishes contained all the bones of the mammalian skull with the exception of the squamous portion of the temporal bone. In other words, all the bones of the mammalian skull were accounted for in fishes; there were no bones left over that might serve as homologues to the bones of the operculum.103 Cuvier's study led Blainville, then Cuvier's protege, to try his own hand at finding homologues of the operculum. Agreeing with Geoffroy that such homologues had to exist, he believed he had found them in a part not included by Cuvier in the skull, namely the lower jaw. Since the lower jaw of fishes appeared to lack some of the pieces found in the lower jaw of the crocodile, Blainville suggested in a memoir read to the Societe Philomatique in 1812 that these pieces had been transformed into the operculum.104 While Geoffroy remembered being satisfied with Blainville's solution, he must not have been fully convinced, because he did not publish a memoir he wrote on the operculum at the time. There the matter was left hanging until 1817, when Cuvier communicated to Geoffroy an anatomical preparation of a pike, showing that the lower jaw of this fish was not missing any pieces which might be transformed into the operculum. Blainville's homologies, then, had to be wrong. With renewed excitement, Geoffroy returned to philosophical anatomy, this time determined to succeed in the solution of his problem. In a flash, as he recalled, the answer came to him: the bones of the gill-cover were homologous to the only other bones left out of Cuvier's enumeration of the bones of the skull, namely the chain of tiny bones in the middle ear of mammals: the malleus, the stapes, and the incus! "From this moment on," Geoffroy reminisced in 1830, "I regained courage and recommenced my studies never to abandon them again."105

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Events now moved quickly. Geoffroy read a memoir on his discovery to the Academie des Sciences on 23 and 30 June 1817 and had it published in extract form in the Bulletin of the Societe Philomatique. To Geoffroy's perturbation, Blainville was induced by Geoffroy's article to publish his own article of 1812 in the same volume. Geoffroy feared, quite legitimately given Cuvier's proclivity for noting that speculation inevitably led to contradictory results, that the two articles presenting alternative solutions to the same problem might place the study of homologies in a bad light.106 Retiring to his retreat at Chailly with his aide-naturaliste Pierre-Antoine Delalande, Geoffroy quickly composed his first book and master work, Anatomical Philosophy: Of the Respiratory Organs with Respect to the Determination and the Identity of their Osseous Pieces, published in 1818. In 1807 Geoffroy had not presented his work as a novel venture, but by 1818 he was acutely aware of the originality of his doctrine. The very title, Anatomical Philosophy, in imitation of Lamarck's Zoological Philosophy, but also of several eighteenth-century volumes such as Fourcroy's Chemical Philosophy (1792), proclaimed the broad vision of the work. Even some of his admirers would have preferred the more reserved "Philosophical Anatomy," the designation by which research on homologies thereafter came to be known in France and Britain.107 Indeed, much of the introductory portion of Philosophie anatomique was a personal and polemical defense of Geoffroy's right to publish controversial ideas. In the manner of the social reformers of the day, he pretentiously traced in the preliminary discourse the previous epochs leading up to the publication of this work, which marked the inevitable transition to a new and greater epoch. No longer was the reader permitted to accept the new homologies without confronting the theory inspiring them, for at the outset of his book, Geoffroy posed the fundamental question: "Can the organization of vertebrates be reduced to a uniform type?" To his belief that "we will always find in each family all the organic materials found in another," he gave the designation "theory of analogues."108 The key to determining homologies was provided by Geoffroy's new method embodied in the "principle of connections." Naturalists had always instinctively discovered homologies, Geoffroy observed, but such comparisons had generally been limited to cases where the form (shape) or the function of the parts was unaltered. When the form and function of the parts changed, naturalists, no longer able to discern homologies, abandoned the search and contented themselves with classifying and describing. "The thread of Ariadne escaped from their hands."109 Despite the recent advances in comparative anatomy, the search for homologies remained limited, because the method of comparative anatomy, Geoffroy claimed, required the existence of intermediary forms so that a structure could be traced from one extreme point to the other. If intermediary forms were lacking, as they were in the comparison of the pectoral fin of a fish and the arm of a man, nothing further could be accomplished. A new method was needed, Geoffroy declared, and he believed he had discovered it in the principle of connections. Armed with this principle, the researcher could "dispense with passing

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degree by degree through all the transformations of organs" and instead formulate homologies directly between animals of different classes.110 "Strictly, it will suffice for you to consider man, a ruminant, a bird, and a bony fish. Dare to compare them directly, and you will reach in one stroke all that anatomy can furnish you of the most general and philosophical [nature]."111 Geoffroy maintained that while the form and function of osseous pieces were subject to wide variation, the connections of the pieces to each other remained constant. The materials of organization, Geoffroy held, existed in some sense prior to function, for the same pieces could be called into the service of different functions in different animals. Thus if one wished to proceed beyond obvious homologies, those based on identity of form or function, connections offered the most reliable guide. For the first of many times to come, Geoffroy proclaimed, "An organ is sooner altered, atrophied, or annihilated than transposed."112 It was by the principle of connections that Geoffroy claimed to have arrived at the astonishing and unexpected homologues in mammals of the bones of the operculum. But before one could apply the principle, one first had to admit a transformation or "metastasis."113 As Geoffroy had indicated in his 1807 papers, the organs of fishes were crowded together under the cranium. The abdominal cavity, which began at the middle of the vertebral column in mammals and toward the posterior end of the vertebral column in birds, was translated to the anterior end of the vertebral column in fishes. Once this shift was admitted, then the principle of connections could disclose the homologues of every structure in fishes. By tracing connections, Geoffroy determined to his satisfaction that the tympanic cavity of mammals, birds, and reptiles corresponded to the gill cavity, and the Eustachian tube to the respiratory canal connecting the gill chamber with the mouth in fishes. In mammals the tympanic cavity contained four bony pieces, namely the incus, malleus, stapes, and a fourth piece that Geoffroy called the lenticular. These, Geoffroy proudly concluded, were precisely the four pieces necessary for the determination of the homologues of the bones of the gill-cover. The operculum, he decided, corre-, sponded to the stapes, the interoperculum to the malleus, the suboperculum— which was actually formed of two bones—to the lenticular and the incus, and the preoperculum to the tympanic, a bone in mammals which partially encloses the eardrum.114 The transformation in function implied by this homology appeared incredible to contemporaries. Bones which served for hearing in mammals became bones assisting respiration in fishes. Moreover, Geoffroy invited further incredulity by insisting that the primary function of the bones of the middle ear was not audition but rather respiration. The primary function of anatomical pieces, as Geoffroy defined it, was their function when they were at their maximum of development. Since the bones were clearly more developed in fishes than in mammals, then, Geoffroy contended, they were essentially bones of the respiratory system. In mammals, he explained, the opercular bones were no longer needed for respiration, and were therefore rendered rudimentary and placed in the service of a secondary function, that of hearing.115 The bones of the operculum formed the subject of the first of five memoirs

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on the bones of fishes making up the Philosophie anatomique. The remaining four memoirs went over the ground covered in Geoffroy's 1807 memoirs, but in much greater generality. They treated the sternum; the hyoid; the bones associated with the larnyx, trachea, and lungs; and the bones of the shoulder and arm in the four classes of vertebrates. Projected volumes, never written, were to continue the demonstration of unity of composition in the pieces serving as support to the organs of sense and movement, and in the muscles.116 Geoffroy's procedure for determining homologies can best be illustrated by his treatment of the sternum. He sought the vertebrate in which the sternum reached a maximum of development to serve as a basis of comparison for the sterna of all other vertebrates. That animal Geoffroy considered to be the tortoise with its large and complex plastron. In the "sternum" of the tortoise Geoffroy counted nine pieces, some paired and some unpaired, which he assumed to be the maximum number. He then supplied them with names: entosternal, episternals, hyosternals, hyposternals, and xiphisternals. By tracing connections of bones he could then determine which of the pieces found in the tortoise had disappeared in fishes and which of them still remained. In the fishes he found no entosternal or xiphisternals; the remaining pieces were separated and closely associated with bones of the hyoid. Geoffrey claimed that the hyoid achieved its maximum development in fishes in which it consisted of nine pieces also named by him: a basihyal, entohyal, and urohyal, and two each of glossohyals, apohyals, and ceratohyals. As in 1807, the branchiostegal rays were identified with sternal ribs. Although Geoffroy's determination of the sternum in fishes was controversial from the beginning, the terminology and the effort to apply it to all the vertebrates was an important legacy to comparative anatomy.117 Philosophie anatomique was laced with a variety of gratuitous physiological explanations to account for metastases of structures, for seeming intercalations of bones distinct in other species, and for the atrophy or disappearance of bones. Geoffroy even offered a dubious physiological basis for the principle of connections. On the assumption that the blood vessels distributed molecules of blood to form each bone in sequence, he argued that since the smaller vascular branches were attached to the principal branch in a fixed order, the bones formed by each branch would also retain a fixed order.118 Such hypotheses left him vulnerable to justifiable criticism. If Philosophie anatomique was not controversial enough already, Geoffroy injected into his memoir on the larnyx, trachea, and lungs a part of his subtle fluid speculations from his Egyptian period. Although he recognized that the topic was inappropriate in a work on comparative anatomy—"I know that I am not in a position to be an authority on such questions, and that I have much more to fear than to hope in this enterprise"—he nevertheless ventured into a digression on the nature of sound, taken from his grandiose system of 1801. In opposition to the universally held belief that sound was due to a movement of the air, Geoffroy, following Lamarck's Hydrogeologie, insisted that sound was a separate elastic fluid. Whatever they might think of the main

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argument of Philosophie anatomique, most reviewers agreed that the theory of the nature of sound was "entirely out of place" and totally lacking in proof.119 Despite some evident hesitation, Philosophie anatomique met with an overall enthusiastic response. Reviewed by Cuvier in his annual report to the Academie and by several naturalists and physicians in scientific and medical journals, it was praised for its originality and breadth of view. The work clearly had something new to say, and to some it bid fair to open a new era of research. To other reviewers, however, Geoffroy's approach to anatomy, while undeniably promising, threatened to overturn common sense. What was the meaning of homologies, especially of those comparisons in which supposedly homologous organs took on widely differing functions? In what sense could the "same" organs be said to be transformed from one animal to another? Although not yet as opposed to Geoffroy's undertakings as he was to become in the 1820s, Cuvier showed obvious signs of skepticism in his annual reports to the Academie for 1817 and 1818. He began applying to Geoffroy the strategies he had found so effective in dealing with earlier questionable doctrines. Employing his oft-used tactic of separating solid fact from suspect theory, Cuvier announced that he would consider Geoffroy's book under three separate heads: (1) the enumeration and description of all the pieces composing each of the organs contributing to respiration in the fishes; (2) correspondences alleged by Geoffroy; and (3) Geoffroy's theoretical considerations. Predictably, he had little but praise for the factual part of the book: "This part of the work, consisting of certain facts, clearly exposed and in large part new, will always remain a precious acquisition for science."120 As might be expected, he was less happy with the theoretical aspects of the work. Geoffroy's homologies disturbed Cuvier insofar as they neglected the functional approach to anatomy. Surprisingly, Cuvier found Geoffroy's comparison of the bones of the operculum to the bones of the mammalian ear less objectionable than many of the other homologies. "His opinion on this matter is very bold," he commented, "and yet it will be perhaps the most difficult in all his theory to attack, at least if one employs only the method of comparison."121 It seemed, nonetheless, incredible to Cuvier that organs of hearing ought to be "considered as merely a sort of superfluity in a rudimentary state. . . in animals with lungs, and as fully developed and indicative of a strictly necessary organization in fishes."122 Even more disturbing were the complicated transformations of organs presupposed by Geoffroy's homologies. Cuvier treated with evident disbelief Geoffroy's determination of bones of the sternum in fishes and his contention that the branchiostegal rays supporting the branchiostegal membrane in fishes corresponded to the sternal ribs in birds. Such a comparison appeared to him meaningless, for the transformation of bones from birds to fishes, if taken in physical rather than ideal terms, was topologically impossible to conceive. To transform in the mind the sternum and hyoid in birds to Geoffroy's supposed homologues in fishes, one had to imagine "a sort of fusion and interlacing of the sternum and the hyoid, and to suppose that the sternal annexes have become intercalated between the styloid bones and the rest of the styloid

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cornua of the hyoid." This, Cuvier ventured, "will be, without doubt, we repeat, one of the strongest objections that can be raised against him."123 Although in 1817 and 1818 Cuvier refrained from pronouncing final judgment, his descriptions of Geoffroy's results carried an underlying note of sarcasm and ridicule that was to become more pronounced in later years: It is almost impossible for us to enter in detail into all the transpositions, and all the movements of the pieces of the organic machine that these analogies presuppose, and still less to analyze all the reasons that the author assigns for these movements.124

By the 1820s, as we shall see, Cuvier pushed his objections to philosophical anatomy further, and became convinced that homologies could only be established where there was constancy of function in addition to constancy in the number and arrangement of parts. Other reviews were more favorable than Cuvier's, but they too voiced reserve on Geoffroy's subordination of physiological to morphological considerations. Etienne Pariset, future permanent secretary of the Academie de Medecine, was skeptical of the generality of Geoffroy's claims: But, as little as I am versed in these matters, I have much fear that in research of this nature, at each step the rule will be encountered by the exception, and that the supposition of an exact parity among such different organizations will in the end be relegated to chimerical ideas.125 To say [of birds] that their wings represent our arms and our hands is a pure play of the mind. The materials are similar, but the disposition, the form, and the use differ completely.126

Pariset, however, concluded in Cuvierian fashion that the problem of discovering homologies between vertebrates was worthy of the efforts of naturalists. If it failed to lead to a complete solution, at least it would provide naturalists with the opportunity of studying animal organization more closely. Another reviewer, A.-J.-L. Jourdan, a future member of the Academie de Medecine and a later proponent of philosophical anatomy, resisted on functional grounds Geoffroy's conclusion that the larynx was not essentially an organ of voice. Geoffroy had argued this seemingly paradoxical statement on the grounds that the bones of the larnyx were found in fishes although fishes were voiceless. Jourdan, too, at this time, preferred functional considerations to purely morphological ones. "It seems to us," he avowed, "that whenever one wishes to state incontrovertible principles in physiology, it is necessary to establish them on the essence of function and not on the mechanical means that nature has employed in order to arrive at the goal."127 Nonetheless, he believed that Philosophie anatomique was "a book that will mark an era in the history of anatomy," a work in which readers would find "in addition to a doctrine, disputable, perhaps, in certain respects, but nevertheless quite ingenious, a multitude of insights and observations which could modify or rectify the ideas that they have drawn from treatises of ordinary anatomy."128 It might come as a surprise that the two most laudatory reviews of Philosophie anatomique should have been written by men who later became Cuvier's

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closest disciples—Frederic Cuvier and Pierre Flourens. Frederic Cuvier, then keeper of the menagerie of the Museum under the direction of Geoffroy, was unhesitatingly sympathetic to Geoffroy's bold handling of anatomy. On Geoffroy's determination of the homologies of the bones of the operculum, he wrote: Free of the presupposition [that the ossicles of the ear are essentially associated with hearing], M. Geoffroy, having attentively examined these bony parts, has demonstrated their analogy with those of the operculum. And he has done so with the aid of such curious comparisons and such plausible reasoning that one cannot help admitting his determination, as extraordinary as it may at first appear.129

Savants would find in the work "wealth and a new direction," and if the work had been written more in their idiom, he concluded with a flourish, "moralists and philosophers would have found in it, in their turn, new grounds for admiring the infinite power of nature . . . to form, in some sense, the all out of emptiness, to create everything from nothing."130 Flourens, just beginning his career, was exhilarated by the prospects opened by Geoffroy's work. Trained in medicine at Montpellier, Flourens had arrived in Paris to study the natural sciences with letters of recommendation from Candolle and obtained admission to collections at the Museum in 1814. Geoffroy, a collaborator on the Revue encyclopedique (successor to the Magasin encyclopedique), may have helped Flourens to obtain the post of reviewer of scientific books for the journal. In any case, Geoffroy could not have hoped for a warmer review. Flourens's essay not only appeared in the journal, but he had it published separately in a brochure which he distributed to all the members of the Academie des Sciences.131 Later an arch-defender of "positive facts" in science, Flourens in 1819 specifically praised Geoffroy's book for its appeal to ideas. The time had come, he enthusiastically proclaimed, to warn savants "that naked observations inevitably become sterile, and that reasoning alone can fertilize them." Philosophie anatomique "fulfills a genuine need of the times," because all previous work had been marred by insufficient use of reasoning and exclusive attention to differences of animal organization: The publication of this work will thus fix the date of a new direction for anatomical studies by bringing them into permanent contact with an eminently philosophical idea, namely, the unity of organic composition, or, if you will, the theory of analogues.132

In Flourens's estimation, Geoffroy had succeeded in demonstrating that all the materials of which fishes are composed are "entirely the same" as the materials found in mammals, reptiles, and birds. The class of fishes had no need for "the intervention of new organs, or of organs created only for them: all is therefore uniform in the vertebrates."133 In part, Flourens's infatuation with Geoffroy's doctrine stemmed from what he perceived as its materialistic nature. The antivitalist aspect of Geoffroy's theory, which Cuvier and others found objectionable, was precisely what gave the work favor in the young Flourens's eyes. Indeed, philosophical anatomy

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provided for Flourens a means of dispensing with antiquated vital forces. "Ever since Galileo reduced the horror of the vacuum to be no more than the effect of the weight of the air, modern physicists [physiciens] have discarded the yoke of occult forces," Flourens wrote. So too would the progress of physiology replace vital forces—which were to Flourens nothing more than "occult" laws—by physical laws. "I am persuaded," Flourens avowed, "that even the provisional acceptance of the term vital force is an evil."134 For a few years Flourens became Geoffroy's devoted disciple, but not long afterwards he was to make a major about-face, eventually to become the defender of Cartesian dualism against Gall. In 1819, however, he, like many others, was enthralled by the promise of philosophical anatomy. Geoffroy's long-awaited master work, while undeniably controversial, was enthusiastically welcomed by scientific contemporaries. Whatever their doubts over the meaning of homologies, few seriously questioned the overall originality and significance of the book. In many respects, Geoffroy was at the height of his career in 1818. Professor at the Museum and at the Faculte des Sciences, and a member of the Institut, he had achieved a financially secure and comfortable base, and had become a leading member of the scientific community, the most respected French naturalist after Cuvier. With his memoirs of 1807, his name had become associated with a novel doctrine which gave a new direction to comparative anatomy. Through his publications on philosophical anatomy, he had focused the attention of naturalists on a new set of problems in comparative anatomy and had suggested a fertile method for their solution. Inspired by Geoffroy's apparent success in reducing vertebrates to a common type, other French naturalists such as Savigny, Blainville, Dutrochet, and Dumeril were led to write memoirs of philosophical anatomy, while bright young men such as Flourens surrounded him as disciples. Almost all the wellknown naturalists in France were attracted in some degree by the lure of pure morphology. Even Cuvier shared in the excitement generated by Geoffroy's discoveries. While emphasizing the priority of functional considerations and of facts in natural history, Cuvier was still willing to grant that there existed resemblances of structure independent of all use. In the period before 1820, he not only condoned, but even encouraged, philosophical anatomy, as long as theorizing was kept within reasonable bounds. Philosophical anatomy was at high tide in 1818, and had as yet given rise to but a hint of discord. This more or less harmonious state of affairs was to end abruptly in the 1820s.

Etienne Geoffroy Saint-Hilaire (1772-1844). (Courtesy of the Bibliotheque Centrale du Museum National d'Histoire Naturelle.)

Georges Cuvier (1769-1832). (Courtesy of the National Library of Medicine.)

Natural History Galleries of the Museum d'Histoire Naturelle, 1822. (From J. P. F. Deleuze, Histoire et description du Museum Royal d'Histoire Naturelle, 2 vols., Paris, 1823. Courtesy of Princeton University Library.)

Homologies of "opercular bones" in the vertebrates, from Etienne Geoffroy Saint-Hilaire, Philosophie anatomique, Paris, 1818. 1-2, man; 3-4, ram; 5-7, owl; 8, pike; 9-11, crocodile; 12, grouper; 13, gar (lepisosteus). o, stapes or operculum; m, malleus or interoperculum; 1, lenticular or suboperculum; e, incus; p, tympanic or preoperculum; st, stylohyal; c, eardrum; t, temporal; j, jugal; x, coronal; u, dentary. (Courtesy of the University of Ottawa.)

Homologies of the bones of the arm and shoulder girdle in fishes, from Etienne Geoffroy SaintHilaire, "Premier memoir sur les poissons . . . ," Annales du Museum d'Histoire Naturelle 9 (1807), pp. 357-372. (Courtesy of the American Philosophical Society Library.}

Homologies of sterna of mammals, birds, and reptiles (including amphibians) from Etienne Geoffroy Saint-Hilaire, Philosophie anatomique, Paris, 1818. 14, tapir; 15, chicken; 16, starling; 17, woodpecker; 18, seal; 19, ornithorhynchus (platypus); 20, Tupinambis (a South American lizard); 21, tortoise; 22, frog; 23, green lizard; 1, episternal; o, entosternal; m, hyosternal, n, hyposternal; p, xiphisternal; ss, sternal ribs; f, furcular; c, coracoid; a, acromion. (Courtesy of the University of Ottawa.)

Geoffroy's determination of bones of the sternum and hyoid in fishes, from Etienne Geoffroy Saint-Hilaire, Philosophie anatomique, Paris, 1818. 24, 25, 28, grouper; 26, 27, 29, ling; 30, pike; 31, St. Peter's fish or John Dory. Bones of the hyoid: a, apohyal; c, ceratohyal; g, glossohyal; b, basihyal; e, entohyal; u, urohyal; st, stylhyal. Bones of the sternum: 1, episternal; m, hyosternal; n, hyposternal; ss, sternal ribs. (Courtesy of the American Philosophical Society Library.)

Geoffroy in the 1820s.

Segment of an arthropod compared to a vertebra, from Etienne Geoffroy Saint-Hilaire, "Considerations generates sur la vertebre," Memoires du Museum d'Histoire Naturelle 9 (1822), pp. 89-119. i, "vertebral nucleus"; a', a", e', e", superior pieces of the vertebra; o', o", u', u", inferior pieces of the vertebra; p, cutaneous appendages. The superior pieces are separated and the inferior shown in place. (Courtesy of the American Philosophical Society Library.)

Geoffroy's ideal plan of the skull of vertebrates consisting of seven vertebrae, from Etienne Geoffroy Saint-Hilaire, "Composition de la tete osseuse de Thomme et des animaux," Annales des sciences naturelles 3 (1824), pp. 173-192, 245-299. (Courtesy of the Canada Institute for Scientific and Technological Information, Ottawa.)

Geoffroy's papers on evolution for the most part dealt with technical arguments concerning the pieces of the cranium of living and fossil crocodiles. Shown here is the foldout plate on cranial pieces of crocodiles from Geoffroy's series of five memoirs, "Divers memoires sur de grands sauriens... ,"Memoires de I'Academie des Sciences 12 (1833), pp. 3-138. (Courtesy of the Canada Institute for Scientific and Technological Information, Ottawa.)

Pierre-Andre Latreille (1762-1833).

E. R. A. Serres (1786-1868).

(Courtesy of the National Library of Medicine.)

(Courtesy of the New York Academy of Medicine.)

Jean-Victor Audouin (1797-1841).

Antoine-Louis Duges (1797-1838).

(Courtesy of the Bibliotheque Centrale du Museum National d'Histoire Naturelle.)

(Courtesy of the Osier Library, McGill University.)

Drawings of (A) a quadruped bent over backwards and (B) a cuttlefish, intended to refute the claims of Meyranx and Laurencet, from the plate accompanying Georges Cuvier, "Considerations sur les Mollusques, et en particulier sur les Cephalopodes," Annales des sciences naturelles 19 (1830), pp. 241-259. a, brain; b, ear; c, jaw or superior mandible; d, inferior mandible; e, e', e", esophagus, stomach, and intestinal canal; f liver; g, respiratory organ; h, "aortic heart"; i, "veinous heart"; k, principal vein; /, principal artery; m, organ of generation; n, kidney (mammal); o, bladder (mammal); p, siphon (cephalopod); r, anus; s, genital orifice; t, spinal cord (mammal). (Courtesy of the Canada Institute for Scientific and Technological Information, Ottawa.)

"The great philosopher fell into a profound sadness." Caricature of Geoffroy and Cuvier by J. J. Grandville in Honore de Balzac, "Guide-ane a l'usage des animaux qui veulent parvenir aux honneurs." (Courtesy of the University of Ottawa.)

"The savants sent an Academician armed with his works." An ass in the costume of the Institut with the Institut in the background, by J. J. Grandville in Honore de Balzac, "Guide-ane a 1'usage des animaux qui veulent parvenir aux honneurs." (Courtesy of the University of Ottawa.)

"My master obstinately called the illustrious Marmus." Caricature of a successful savant by J. J. Grandville in Honore de Balzac, "Guide-ane a 1'usage des animaux qui veulent parvenir aux honneurs." (Courtesy of the University of Ottawa.)

Isidore Geoffroy Saint-Hilaire (1805-1861). (Courtesy of the Bibliotheque Centrale du Museum National d'Histoire Naturelle.)

Pierre Flourens (1794-1867). (Courtesy of the National Library of Medicine.)

Henri Milne Edwards (1800-1885) (Courtesy of the National Library of Medicine.)

Henri Marie Ducrotay de Blainville (1777-1850). (Courtesy of the American Philosophical Society Library.)

Owen's Archetype, from Richard Owen, On the Archetype and Homologies of the Vertebrate Skeleton, London, 1848. (Courtesy of the National Library of Medicine.)

CHAPTER 5

The Battle Lines Are Drawn: 1820-1829

When, in 1828, Cuvier published the first volume of his long awaited Histoire naturelle des poissons, he began with a lengthy introduction which allowed him to give vent to his frustration over the philosophical currents that were swirling around him. The very first sentence delivered a swift blow to all theorizing on the links between fishes and the other classes of vertebrates: "L'histoire naturelle est une science des faits." Grouping together a number of speculative theories—the chain of being, unity of composition, and evolution—Cuvier dismissed them all at once: Let us especially conclude that they [fishes] are neither links in this imaginary chain of successive forms. . . nor in that other chain, no less imaginary, of simultaneous and nuanced forms, which has reality only in the imaginations of some naturalists [who are] more poets than observers.1

Reaffirming his commitment to a strict functionalism, Cuvier insisted that fishes had their assigned place in the creation and were neither ennobled mollusks nor fetuses of mammals. If they possessed any organs in common with other vertebrates, it was solely because these organs were created to perform similar functions. To Geoffroy, Cuvier's final phrase, "more poets than observers," came as a personal slur. In his course at the Museum in 1828, soon published, he vigorously defended himself against the "grave opposition" coming from "on high, from the greatest authority in the natural sciences."2 Much had changed in the ten years since Geoffroy published Philosophie anatomique. Before 1820 there had been little overt animosity between Cuvier and Geoffroy. As skeptical as he may have been of philosophical anatomy, Cuvier had discouraged only such excesses as Dumeril's unfortunate vertebral theory of the skull. With his tacit approval, philosophical anatomy was finding a respected place for itself alongside more traditional descriptive and functionalist anatomy. The first break came in 1820. Geoffroy's astounding suggestion that the common plan of the vertebrates was also applicable to insects marked the turning point in the relations between the two rivals. Thereafter, a series of events—scientific, institutional, and political—steadily intensified the con105

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troversy over philosophical anatomy. While Geoffroy became ever bolder in his pronouncements, Cuvier grew increasingly defensive. By the end of the decade, the two men found themselves divided on every level: anatomical doctrine, methodology, metaphysics, patronage, and politics. The most obvious cause of the deterioration of relations was Geoffroy's extension of philosophical anatomy into new territory. Whereas previously his homologies had been confined to the vertebrates, in 1820 Geoffroy began to suggest homologies between the exoskeleton of insects and Crustacea and the skeleton of vertebrates. These comparisons, which completely ignored the constraints imposed by Cuvier's functional teleology, threatened to eliminate the gaps between Cuvier's four embranchements. Soon after, with the aid of his disciple E.R.A. Serres's work in embryology, Geoffroy extended unity of composition to yet another arena, that of human and animal malformations, in his formulation of the new and often controversial science of teratology. As he elaborated his theories, Geoffroy made ever more explicit the antiteleological context in which he conceived them. His attacks on preformation and vitalism tainted him with pantheism or, worse, materialism. Finally, he drew what would seem to the modern mind an obvious conclusion from unity of composition, namely a theory of evolution. Cuvier's four distinct embranchements and his functionalist teleology were beset from all sides in the 1820s. Audouin adapted Lorenz Oken's vertebral theory of the skull to insects. Building upon the work of Savigny, Audouin and Latreille established homologies of appendages throughout the articulate embranchement. Latreille sought connecting links between fishes and invertebrates, while Blainville continued to publish and profess his unorthodox views on the animal series. Having for the most part given his teaching posts over to suppleants, Cuvier was no longer able to exert a direct influence on the younger generation who were gaining their initial exposure to zoology and comparative anatomy from Geoffroy, Blainville, and Latreille at the Museum and Faculte des Sciences. Across the Rhine, the leading anatomists publicized a German style of transcendental anatomy, which, although it may have borrowed something from the French—German anatomists such as Oken, Johann Friedrich Meckel, and Johann Baptist von Spix had studied in Paris in the first decade of the nineteenth century—was also firmly rooted in an independent German philosophical tradition, based upon the writings of Kant. While some German anatomists—the so-called Gottingen school, in which Meckel can be placed—took a relatively empirical approach, Oken and others of the school of romantic Naturphilosophie endowed transcendental anatomy with a mystical form and cosmic dimensions that were entirely absent in the French counterpart.3 The French, however, did not distinguish among the schools of German science, and friends and foes alike linked Geoffroy with an undifferentiated "German philosophy of nature." Whereas Geoffroy had compared the parts among different classes of animals, the characteristic homology of the German anatomists was the serial homology—the repetition of parts within a single animal. These correspon-

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dences they called "homologies," in distinction to Geoffroy's "analogies."4 The most provocative and debated of "homologies" among the vertebrates was that between the vertebrae and portions of the skull. Oken proposed in 1807 that the vertebrate skeleton was in essence composed of a series of vertebrae, the first four of which had been modified to form the skull.5 Repugnant to Cuvier, the vertebral theory of the skull was widely adopted in Germany and elaborated upon by a considerable number of anatomists. Moreover, Oken and Spix believed they could discern an even deeper level of homology in the vertebrate skeleton, a repetition of the organs of the trunk in the parts of the head. According to Oken's system, the mouth corresponded to the stomach, the nose to the lungs, and the jaws to the arms and legs. Whenever Cuvier wished to discredit all of German theorizing, he paraded this fantastic notion before his audience. Although it shared some ideas with Naturphilosophie, Geoffroy's doctrine was based on a different foundation. German biology was teleological, fundamentally indebted to Kantian philosophy, and gave primacy not to matter but to forces and their transformations. Because German biologists believed nature possessed an internal dynamism, their science was considered by the French to be pantheistic. Geoffroy's doctrine, in contrast, was nonteleological, and as in the eighteenth-century Newtonian tradition, forces were assumed to be superimposed upon matter.6 Geoffroy shared none of the mysticism found in the writings of romantic Naturphilosophen such as Oken. Both he and Serres readily granted with Cuvier that Oken's comparison of the head and trunk was an abuse of a priori reasoning, yet they believed the Germans had a right to be given a fair hearing. While they disassociated themselves from the worst excesses of the Germans, they therefore defended them as best as they could against Cuvier's attacks. Geoffroy adopted from the German school the vertebral theory of the skull, as well as the idea, first suggested by Kielmeyer as early as 1793, of a parallelism between the stages of development and the series of living organisms. In particular, Geoffroy, and especially Serres, borrowed and extended the doctrine that the developing fetus of the mammal repeated in succession the forms of animals lower in the scale. In the 1820s, the heyday of Naturphilosophie, the formerly separate French and German traditions began to merge. From 1817 on, Geoffroy's works were translated into German and published in the major journals of Naturphilosophie, Oken's Isis and Meckel's Deutsches Archive fur die Physiologie.7 And in the 1820s, especially after the founding of the Annales des sciences naturelles in 1824, the writings of the German anatomists became available in French translation. New trends threatened Cuvier not only on an intellectual plane but also on a professional and institutional level. In the Academie des Sciences, Cuvier's position was being undermined by a "new guard" of scientists led by the liberal, Francois Arago, which sought to wrest power to control scientific patronage by taking over key positions in the scientific community. Supported by Arago, Blainville, a sworn enemy of Cuvier, was able to enter the Academie in 1825 despite Cuvier's determined opposition. Faced by this new challenge to his authority, Cuvier redoubled his efforts to obtain the allegiance of young

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scientists and to keep his followers in line. While in the 1810s it was still possible for Cuvier's disciples to venture into controversial areas, by the late 1820s the limits were clearly set. Any author of a paper that attempted to bridge the four embranchements or establish homologies within an embranchement without consideration of "the conditions of existence" risked losing Cuvier's support. Geoffroy, who controlled much less patronage than Cuvier, saw his students slip away from him to become proteges of his rival. Cuvier's tactics in this period were cleverly satirized in the 1840s by Honore de Balzac in a littleknown short story (see Chapter 7) in which the students of "The Philosopher" (Geoffroy), lured on by promises of positions, sell out to the "Baron Cerceau" (Cuvier), and adapt their once controversial theories to fit his requirements. The political environment in France only compounded the alarm with which Cuvier and his followers viewed opposing scientific doctrines. The year 1820 saw liberal revolts against several established monarchs in Europe.8 In France the Due de Berry was assassinated, leading to the fall of Louis XVIII's moderate ministry and its replacement by a ministry of ultraroyalists. Whatever ministry was in control, it was constantly buffeted from the right and the left. In a country still divided by wounds inflicted during the Revolution and Empire, no ministry was stable and no single policy satisfactory to a majority of Frenchmen. One of the most contentious issues of this period was the place of the clergy in public education. In general, the government supported an increasing role for the Church. In 1821, a bishop, Denis Antoine Luc Fraysinnous, was made head of the University system, and after 1824 he became minister of ecclesiastical affairs as well. When the comte d'Artois, the ultraroyalist brother of Louis XVIII, ascended the throne in 1824 as Charles X, the reactionary policy of the government became even more pronounced. The political situation in France was potentially explosive, and ultimately did blow up in the Revolution of 1830, which removed Charles X from his throne only a few months after the Cuvier-Geoffroy debate erupted in the Academie. Concerned as always with maintaining order, Cuvier acquiesced in increasing Church control, despite the fact that he was a Protestant. His association with the repressive policies of the ultraroyalists lost him friends among scientists and left him open to the charge of political expediency.9 The romantic movement, especially prominent in Germany, only added to Cuvier's fear of social uprising. Romanticism, which in the early years of the Restoration had been associated with Catholic and Royalist thought, came in the 1820s to be linked with anticlericalism and the liberal opposition to the government. Cuvier's anxiety concerning the future of France carried over into the scientific arena. Insofar as possible, he utilized his patronage to discourage theorizing that might be perceived to be heretical. Among the theories that he came to regard openly as not only false but dangerous because they placed limits on the Creator's power were a number that Geoffroy professed, among them unity of composition and evolution. As the decade wore on, Cuvier became increasingly insistent on banning speculation from natural history. Aspiring naturalists, especially those desirous of pursuing philosophical anatomy, found it pru-

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dent to avoid the taint of irreligion by limiting their claims and differentiating their work from Geoffroy's. By the end of the decade the tension between the two naturalists was apparent to all, and weighed heavily not just upon the principals but upon the entire scientific community. In all their lectures, articles, books and reports at the Academie, Cuvier and Geoffroy continued to snipe at each other, but neither responded to the other head-on. Geoffroy accused Cuvier of denouncing him behind his back, rather than presenting a reasoned critique of his work. For ten years, beginning in 1820, he tried to lure his scientific opponent into a public confrontation, but Cuvier preferred the strategy of indirect attack. In this uneasy environment, nearly all felt the need for caution in their theoretical statements lest they inadvertently be drawn into the feud to the detriment of their careers. Not just for Cuvier and Geoffroy, but for the entire community of comparative anatomists and zoologists, the battle lines were drawn. Geoffroy, Entomologist: The First Break with Cuvier

Geoffroy's work of the 1810s culminated in his first and most significant book, Philosophie anatomique, which proclaimed unity of composition among the vertebrates. In the following decade, Geoffroy continued to elaborate on this theme, extending the "theory of analogues" to soft parts of the body as well as to other parts of the skeleton. A series of memoirs dealt with homologies of the urinary tracts, digestive tracts, and reproductive systems in vertebrates. A major paper in 1824, published in the newly founded Annales des sciences naturelles, identified all the components of seven cranial vertebrae in the skulls of vertebrates.10 Still another group of memoirs defended his homology of the bones of the operculum of fishes with the bones of the middle ear of mammals against an attack by the German anatomist E.H. Weber, who discovered in some bony fishes a chain of bones extending from the ear to the swim bladder (Weberian ossicles), which he claimed were the true homologues of the ossicles of the ear.11 But Geoffroy's most interesting and significant work in the 1820s took him far beyond the confines of homologies of the vertebrates. Of all his research in this decade, none drew more negative criticism than his excursion into the philosophical anatomy of insects and Crustacea. Until 1820 Geoffroy had restricted his homologies to a single one of Cuvier's four embranchements, that of the vertebrates. With some reservations, Cuvier had generally given warm support to his colleague's research. Already, as we have seen, friends and followers of Geoffroy had begun to establish homologies within another embranchement, that of articulates (now known as the phylum Arthropoda). Going a major step beyond the work of Savigny and Blainville, Geoffroy attempted to establish homologies between the articulate and vertebrate embranchements. This effort, largely a failure, damaged Geoffroy's reputation and brought on Cuvier's first explicit condemnation of his work. When Geoffroy presented his three memoirs on insects to the Academie des Sciences in January and February 1820, he was not the only naturalist to be

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occupied with the comparative anatomy of articulates. Within the space of a little more than two months, the Academie was inundated by a series of novel and unexpected researches on the philosophical anatomy of insects and crustacea. The stream of papers began on 27 December 1819, when Cuvier's collaborator on entomology for Le regne animal, Latreille, presented a memoir in which he sought the homologue of the wing of the insect in Crustacea. The following week, on 3 January, Geoffroy read his first contribution, "On Insects Reduced to the Embranchement of Vertebrates." The very next week, on 10 January, Latreille followed suit with his own version of the link between vertebrates and invertebrates. Geoffroy read his second and third memoirs on 17 January and 21 February. And on 24 January Audouin presented his first paper to the Academie—an adaptation of the vertebral theory of the skull to insects—which was reported upon by Geoffroy on 7 February. A veritable feast of speculation on homologies among articulates and between articulates and vertebrates was provided.12 Of all these papers, Geoffroy's were by far the most controversial, for he argued the seemingly preposterous thesis that the exoskeleton of insects was a true skeleton and could be compared piece for piece to the bones of vertebrates. Insects and vertebrates, Geoffroy pointed out, had numerous parts with names in common: they had a mouth, head, eyes, thighs, hips [hanches], tibias, a thorax, and an abdomen. Only by comparison under a general point of view could one determine which of these usages were just. If Latreille had found so many affinities among the insects, was it not reasonable that one could also observe affinities between insects and vertebrates? "I have never ceased, in fact, to follow the great idea of nature, the unity of organic composition.'" As usual, Geoffroy left the detailed demonstration of unity of composition to later memoirs that were promised but never published. His intent in his memoirs of 1820 was merely to sketch how the comparison might be done.13 The first problem Geoffroy faced was to explain why the insect "skeleton" contained all the internal organs of the insect instead of just the spinal marrow, as in vertebrates. His ingenious justification of this radical difference, based on Serres's still unpublished work on osteogenesis (the development of bones), was at best highly speculative. As in his previous research on vertebrate homologies, Geoffroy assumed the parts were formed by exudates deposited by branches of vessels. Serres had shown that the vertebra in its first stage of development was a single tube which formed from four centers of ossification. In the vertebrates, according to Geoffroy, the nervous system distributed materials around its axis within the vertebral canal, while other organs were formed by the circulatory system outside the vertebral column. The distinctive characteristic of insects, he hypothesized, was that they had a defective circulatory system. Lacking a heart, insects were left with only a single system for the distribution of materials of organization, namely the nervous system. The "skeletal" system of insects therefore remained at the stage of development of the embryonic system in higher vertebrates, that is, the "vertebrae" retained the form of a tube. In insects all the parts were formed by the nervous system and

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distributed within the "vertebral canal." Insects were in essence vertebrates, Geoffroy concluded, and the exoskeleton corresponded to the vertebral column, not to the epidermis as was commonly supposed. Indeed, he generalized, "every animal lives within or without its vertebral column."14 One objection readily raised against all attempts to link insects and vertebrates was that the nervous system in insects was found on the ventral side, while in the vertebrates it was located on the dorsal side. Geoffroy's solution to this problem was repeated by many later philosophical anatomists, and is now believed to have some validity. The definitions of dorsal and ventral, Geoffroy claimed, were purely arbitrary, because they were based solely on the orientation of the animal to the sun. If it was assumed that the insect walked with its "ventral side" rather than its dorsal side toward the sun, then all of the organs of the insect would have the same arrangement as the organs of vertebrates.15 Having explained the peculiar mode of development of insects, Geoffroy went on to sketch the particular homologies that followed. The four points of ossification of the vertebrae in vertebrates corresponded to the four pieces of the segment of the exoskeleton in articulates. He compared the first segment or head of the insect not to the entire head of vertebrates, but to the bones of the face, the bones protecting the cerebrum and the hyoid. The second insect segment corresponded to the bones protecting the cerebellum, the palatine bones, and the larnyx, while the third segment was homologous to the interparietals, the parietals, and the "opercular bones" (i.e., the bones of the middle ear). Geoffroy thus compared the first three segments of insects to three vertebrae which he believed to form the vertebrate skull. The last three segments of the insect, according to Geoffroy, corresponded to the vertebrate thorax, abdomen, and coccyx. Determining the homologues of the second pair of wings presented some difficulty, but Geoffroy eventually declared them to be the equivalent of the swim bladder of fishes, or the air sacs of birds. He even went so far as to consider the appendages of articulates to be homologous to vertebrate ribs.16 In a memoir published in 1822, he displayed for his readers two diagrams, one of a typical vertebra and the other of a segment of a crustacean, with the supposedly homologous parts labeled.17 Such comparisons seemed no less fanciful to his contemporaries than they appear to us today. The members of the Academie gave Geoffroy's memoirs a generally poor reception, although there were some who, like the physicist Andre-Marie Ampere, were, sympathetic.18 According to Geoffroy, the "geometers" at one point instigated an argument over whether insects could be said to possess vertebrae as Geoffroy had claimed. The question was submitted to the entomologists, who, in no uncertain terms, decided against Geoffroy. "I was condemned," bewailed Geoffroy, and he added self-righteously, "Eppur si muove, I could have replied along with the celebrated philosopher of Pisa."19 Cuvier explicitly rejected his colleague's conclusions at a meeting of the professors of the Museum and in private discussions, but not in print. Geoffroy, who from the beginning tried to draw the controversy out into the open,

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reported to his readers on the incident at the Museum in his second memoir on insects: 'Your memoir on the skeleton of insects lacks logic from start to finish,' M. Cuvier said, expressing himself on my first work with extreme ardor before the professors of the Museum. 'You compare things which are in no way susceptible of being [compared]. Nothing in common, absolutely nothing, between the insects and the vertebrates, [or] at the most, a single point, animality.'20

In public, in his annual analysis of the work of the Academie, Cuvier chose not to condemn Geoffroy outright, but instead adopted the more indirect strategy of pointing out (with noticeable sarcasm) that while attempts to bridge the gap between vertebrates and articulates might not achieve their desired ends, they at least had the merit of producing new facts.21 Geoffroy complained in his memoirs that his work was being dismissed without a proper hearing. Instead of confronting him openly, Cuvier was criticizing his work in private. "Disapproving allegations made in general terms and spread in places behind my back are not arguments," Geoffroy wrote, and he challenged Cuvier to "explain himself." "Let him attack my doctrine, let him attack it as heatedly as his conviction prescribes. But at least let it be publicly."22 Philosophical Anatomy and Patronage Politics: Latreille, Audouin, and Flourens

Cuvier's indirect but clear disapproval had only a limited effect on Geoffroy, but weighed more heavily on the other authors of memoirs on articulates, Latreille and Audouin, as well as on Geoffroy's young disciple, Flourens. Stung by the rebuke from most of his colleagues, Geoffroy after 1823 quietly let drop homologies between articulates and vertebrates. But Geoffroy, secure in his career, was not to be deterred for long by Cuvier's displeasure. Latreille, Audouin, and Flourens, however, were still making their way. In such circumstances, Cuvier could be a powerful ally and a dangerous enemy. Though they remained sympathetic to philosophical anatomy, they all subsequently modified their work, in particular by renouncing all efforts to link embranchements, so as not to further offend the acknowledged leader of French naturalists. Latreille, trained as a cleric during the ancien regime, spent the greater part of his career in subordinate positions at the Museum.23 Hired in 1798 to arrange the collections of entomology, he became an aide-naturaliste in 1805 and suppleant of Lamarck in 1820. Since 1814, he had been a member of the section of anatomy and zoology of the Academie. In 1820, Latreille was 58 years old, but he still had not achieved his ambition of a professorship. Latreille had joined in the general excitement over philosophical anatomy in the second decade of the century. Even before Savigny's memoirs on insects, he had intimated the essential identity of the maxillae of caterpillars and the tubule of butterflies.24 Following Savigny's lead, he attempted in 1819 to demonstrate unity of plan throughout the articulate embranchement. As part of this

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program of research, he read to the Academie on 27 December 1819 a paper published under the title "On the Formation of the Wings of Insects, and on the External Organization of These Animals Compared in Various Points with That of the Arachnids and the Crustacea."25 Like Geoffroy, Latreille believed that structural differences were only deceptive appearances. The wing of the insect appeared to Latreille to be an anomalous structure in articulates, since neither the arachnids nor the Crustacea were winged. He concluded from his investigation that wings were simply a modification of the "tracheal feet" of Crustacea, in the same way that the wings of birds and the fins of fishes were regarded as modifications of the arm in man. Latreille reduced several additional "anomalies" to their homologues in other classses. Spiders, for example, appeared to lack antennae. Guided by the principle of connections and observations on the transformations of antennae in other classes, Latreille arrived at the conclusion that the antennae of insects were modified in spiders to form the mandibles. As a former cleric and assistant of Cuvier, Latreille was aware of the implications of morphology for traditional functional teleology, but argued that philosophical anatomy only increased admiration for the Creator: "Insofar as the number of these supposed anomalies is diminished," he wrote, "our respect for nature grows. The harmony of her plan is revealed to our eyes to possess greater unity and simplicity, and hence greater wisdom."26 In "Passage from the Invertebrates to the Vertebrates," read before the Academie on 10 January 1820, Latreille offered his own solution to the linkage between embranchements. Attacking the problem from a slightly different perspective than Geoffroy, he sought continuities between the highest Crustacea and the lowest fishes, the cyclostomes, in the chain of being. He began his argument by tracing the position of the gills up the series of Crustacea, and from the batrachians (frogs and toads) down through the series of fishes. In both cases he found that the gills moved from the anterior to the posterior part of the body. From this initial point of comparison, Latreille deduced a number of specific homologies. The branchiostegal rays of fishes, he decided, corresponded to the last four "feet-jaws" and the ten feet of decapod Crustacea. The carapace of Crustacea corresponded to the fish hyoid, and the appendages of the crustacean tail to the ventral and anal fins of fishes.27 Cuvier's critique of Latreille's memoirs in his report on the work of the Academie was subtle but effective. "M. Latreille," wrote Cuvier, "whose immense works on the positive part of Entomology have made him so celebrated, has felt obliged to devote himself also to some theoretical researches on the means of bringing together insects and vertebrates."28 The clear implication was that Latreille would be best advised not to venture into the uncharted territory of establishing homologies between embranchements. Cuvier pointed out that even the task of finding homologies among the articulates was fraught with hazard. Savigny, Audouin, and Latreille, he observed, had discerned three totally different and contradictory sets of correspondences between the mouthparts of Crustacea and the legs of insects.29 The hallmark of speculative theories for Cuvier was that they failed to arrive at conclusive or positive results. At first, Latreille did not heed the warning and persisted in his attempt to

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link vertebrates and invertebrates. In 1823 he presented a new memoir on the subject to the recently re-established Societe d'Histoire Naturelle. In this, his final excursion into philosophical anatomy, he sought continuities between the highest class of mollusks, the cephalopods, and the cartilaginous fishes. He compared the hard piece in the interior of the cephalopod body to "a sort of general vertebra."30 It was not long, however, before Latreille perceived where his best interests lay. Employed by the Museum and a member of the Academie, he nevertheless lacked the security and prestige of a professorship, and aspired to become Lamarck's successor. By the latter part of the decade, he had come to realize that research on homologies between embranchements placed Cuvier's support in jeopardy, and he restricted his work to classification and description of insects. With Cuvier's aid, he did indeed inherit one of the chairs into which Lamarck's chair was divided in 1830, although by then he was too old to handle the duties. "They give me bread when I no longer have any teeth," the aged entomologist was said to have lamented.31 He expressed his indebtedness and loyalty to Cuvier in a letter dated 30 August 1831: If I finally enjoy some happiness, have you not been the dispenser of it? Since 1795 when I had the precious advantage of becoming personally acquainted with you, you have not ceased to gratify me with all sorts of kindnesses, and I can well say that you are, of all our colleagues, almost the only one on whose friendship and support I can count.32

Audouin, the third naturalist to occupy the Academie with memoirs on insects in 1819-1820, was just starting out on his career. Protege and later sonin-law of the powerful Alexandre Brongniart, Audouin's career was from beginning to end supported by the Brongniart faction in the Academie.33 In 1820 he was still a medical student. From his personal journal for the years 1817-1818 emerges an engaging picture of the life of an aspiring naturalist during the Restoration. In addition to his medical courses at the Faculte de Medecine, the young Audouin filled his days with attending meetings of the Societe Philomatique and the Institut and lectures all around the capital, at the Museum, the College de France, and the Faculte des Sciences. Over the course of the two years he attended lectures on mineralogy by Brongniart, physics by Jean-Baptiste Biot, chemistry by Louis Jacques Thenard, and zoology by Cuvier, Geoffroy, Dumeril, Latreille, and Blainville. Audouin was already occupied with the study of entomology when the concepts he encountered in Geoffroy's and Blainville's courses redirected his work. In his journal entry for 17 July 1818, Audouin made note of working with his friend Lachat on the organization of insects: "We had some ideas concerning unity of composition, and everything led us to believe that the differences noted among the various orders of insects were due only to modifications of parts."34 This research on the philosophical anatomy of insects formed the subject of his first memoir and the occasion of his debut before the Academie. Audouin's memoir, read on 24 January 1820, was a decidedly bold presentation for a beginning naturalist. As it was never published, its contents can

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only be gleaned from the two commission reports on it.35 Audouin apparently began by acknowledging his debt to Geoffroy and Savigny. The paper was to be the first part of a larger study in which Audouin planned to reduce the entire exoskeleton of the various orders of arthropods to a common plan. In this first presentation, however, he proposed to limit himself to the structure of the head in the insects, arachnids, and Crustacea. What Audouin proposed was the equivalent of Oken's vertebral theory of the skull for insects. Oken had advanced in 1807 the theory that the human skull was composed of four modified vertebrae. Without referring to the German anatomist, Audouin adapted the theory to articulates by showing that the head of these animals could be decomposed into the equivalent of three modified body segments, each identified by its associated appendages. The appendages of the first segment of the head were the upper lip (labrum) and mandibles, those of the second segment, which was confounded with the third, were the antennae and lower lip (labium), and those of the third segment were the eyes and maxillae. Extending these results to the other articulates, Audouin proceeded to establish homologies among the appendages of the head in insects, arachnids, and decapod Crustacea. In drawing comparisons, he, like Geoffroy, insisted that consideration of connections ought to prevail over consideration of form or function. In fact, he reproached Savigny for having made false determinations for the pieces of the head of Crustacea, because he had not rigorously followed the principle of connections. One would expect that Geoffroy, charged to report on the memoir along with Latreille and Bosc, would have greeted such a work with great enthusiasm, but such was not the case. Geoffroy was offended by the attack on his friend Savigny and devoted a good part of his report of 7 February to reconciling the latter's determinations with the principle of connections. However, Geoffroy's report failed to be approved by the Academie for quite a different reason, namely Geoffroy's unwarranted attempt to assimilate Audouin's memoir to his own dubious theory of the homologies between insect segments and vertebrae. It became habitual for Geoffroy in the 1820s and 1830s to appropriate the work of other scholars in this way. For example, when Geoffroy mentioned the word "segments" in his report, he added "or, as we prefer to say it, vertebrae," and he referred in passing to the "vertebral column" of insects. An argument arose in the Academie over whether a report which was to receive the Academie's blessing could legitimately employ the word "vertebrae" to designate a segment of the body of insects. Here, as in other examples to come, Geoffroy's propensity to read into memoirs more than was there could get a young author into more trouble than he bargained for. In the end, Geoffroy withdrew his report, which he published in a medical journal, and the Academie named a new commission, for which Geoffroy declared himself ineligible.36 Representing Cuvier's sentiments, this second report was even more damaging than Geoffroy's. Drafted by Lacepede, Cuvier, Bosc, and Dumeril, it was read by Cuvier's disciple Dumeril on 20 March. Dumeril denied outright the premise of Audouin's paper—that any articulate possessed a head formed of

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three body segments. The conclusion of the report, separating fact from theory and praising Audouin for his careful research on insect anatomy, bears the unmistakable mark of Cuvier: However, your Commissioners have recognized in the memoir of this young naturalist the acquisition of exact knowledge on [the subject of] organization, as well as a real talent for the art of observing and exposing facts. And, although they do not adopt his ideas, they regard his work as interesting and worthy of the encouragements of the Academie.37

As in the case of Latreille, Cuvier hoped by such hints to redirect Audouin's efforts away from philosophical anatomy to a more traditional and empirical study of insect anatomy. After this inauspicious debut, Audouin learned his lesson. His second memoir on insect anatomy, read on 15 May 1820, though it still owed something to Geoffroy, fared much better, in large part because Audouin moderated his theoretical claims. In this paper, entitled "Anatomical Researches on the Thorax of Insects," Audouin was careful to emphasize the extensive research on which his conclusions rested. He claimed to have examined at least 300 different species and over 1,200 anatomical pieces. Following Geoffroy's method for determining homologies, Audouin first separated and named the pieces of the thorax, dividing it into three segments: prothorax, mesothorax, and metathorax. Each segment was further divided into four major pieces: the sternum, two lateral pieces called flanks, and the tergum. Choosing to treat the Hexapoda or insects proper first, because in this class the pieces had achieved a maximum of development and were most distinct, Audouin then passed to a detailed comparative study of the modifications of these elements of organization in the various orders. Audouin's research was an evident outgrowth of the work of Geoffroy. Like Geoffroy, he maintained that too much attention had hitherto been paid to form (shape), leading naturalists to the false conclusion that every animal "had alone received as its portion certain organs completely denied to others."38 If such organs as the stinger of the bee, the fluid-launching apparatus of the ground beetle, and the singing organ of the cicada were considered completely different organs, then, according to Audouin, no philosophy would be possible. However, he asserted, once it is shown "that all the differences offered by the insects, all the anomalous organs that they present, are due merely to a lesser or greater development of certain parts existing generally in all of them," then "a philosophy of science has been created."39 His results for the thorax of insects, he indicated, could be extended throughout the articulate embranchement: In fact, all of the differences which are noted in the series of articulates depend only on the similar or dissimilar development of the segments, on the union or division of the pieces composing them, on the maximum development of some [pieces], and on the rudimentary state of others.40

Without referring to it by name, Audouin applied Geoffroy's principle of the balance of organs when he asserted that the maximum development of the

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metathorax always corresponded to the rudimentary state of the mesothorax, and vice versa. Like Geoffroy, Audouin used embryology to support his comparisons. Thus he noted that in larvae each segment was almost uniformly developed, whereas in adult insects certain segments attained an extraordinary development. Insects, he concluded, resembled each other more in the larval than in the adult state.41 While clearly indebted to Geoffroy, Audouin was nevertheless careful to avoid the excesses that cast doubt on Geoffroy's own work. This time he thanked both Cuvier and Geoffroy for their help, limited his comparisons to a single embranchement, and avoided discussion of unity of composition or any other controversial issue. Instead of formulating ad hoc physiological explanations for structural modifications, he was content to correlate structural modifications in the articulates with functional adaptations. For example, he observed that the mesothorax was poorly developed in Orthoptera (grasshoppers) and Coleoptera (beetles), where it bears elytra of little use for flying, while the sternum was most fully developed in those insects that made the greatest use of their legs. Where Geoffroy would have offered a causal explanation for these modifications, Audouin simply noted changes in functional requirements. Instead of introducing a metastasis or comparing two extreme forms directly, Audouin took the cautious route of proceeding by means of a series of intermediate forms in formulating his homologies. And he deliberately chose to sidestep the thorny subject of Geoffroy's homologies between insects and vertebrates, claiming that Geoffroy's generalization was yet to be demonstrated: Without denying the exactness of his observation, we will wait, before adopting his theory, until the numerous facts that it embraces have each been put to a severe examination. . . . If this manner of envisioning the skeleton of articulates had been useful to the understanding of our subject, we would have entered upon it without hesitation. But, not seeing the absolute necessity, we will abstain from saying more about it.42

Anxious to gain Cuvier's approval, the young man wrote to him on 17 July 1820, after the commission had been named at the Academie, to request an interview to discuss his work. He emphasized in his letter the five years of research that had gone into his presentation before the Academie, and avowed that it was through Cuvier's "precepts" and "examples" that he had learned "that it was not so much necessary to occupy myself in accumulating facts without method, as to coordinate them and deduce rigorous consequences from their examination."43 Because Audouin had made a deliberate effort not to offend Cuvier, Cuvier was able to support his research. But although Cuvier encouraged Audouin in his report to the Academie, presented on 19 February 1821 in the name of a commission composed of Lacepede, Dumeril, and himself, he also availed himself of the opportunity to point out the limits of acceptable research. By a conditional offer of patronage, Cuvier hoped to guide the young naturalist away from hazardous theorizing. The report totally avoided mention of Geof-

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froy or of the doctrine of unity of composition. Audouin's work, as Cuvier saw it, was simply a study of comparative anatomy, a science as old as Aristotle: Whoever has taken the trouble to compare a certain number of natural beings of the same kingdom regne or of the same class should perceive that in the midst of [all] these innumerable variations that they present in size, in form, and in color, there are certain similarities [rapports] in the structure, the position, and the respective functions of the parts, and that with a bit of attention, one can follow these similarities through the differences which on a superficial glance sometimes mask them.44

By such statements, Cuvier was able to disassociate Audouin's work on homologies from Geoffroy and from doctrinaire statements on unity of composition, whether between or within embranchements. If the message were not yet clear, Cuvier added a warning against carrying the study of homologies too far. He noted that the "philosophical spirit" had progressed to the point of leading some uncautious persons to seek links among all beings by means of "successive analogies" and "to deduce a priori the general and particular composition of universal laws of ontology and the most abstruse metaphysics." However, he hastened to add, Audouin's present work was not of this dubious character: "The work of M. Audouin belongs to the class of researches which are exempt from dispute, because they are based solely on positive observation."45 Cuvier's patronage was of immense importance in Audouin's career. When Audouin published his second memoir in 1824, he gratefully acknowledged Cuvier's support: Finally, if a sentiment of modesty ought to prevail over a vivid sentiment of gratitude, and if it were possible to satisfy one [sentiment] and not wound the other, we would pass over in silence the distinguished report that M. the baron Cuvier gave at the Academie des Sciences in the meeting of 19 February 1821. The sanction of such a master who, after having approved my debut, encourages me daily in my work, is a flattering pledge to which I attach the greatest value.46

In 1825, with the aid of Cuvier, Audouin was hired to teach at the Museum in the place of Latreille, who was then suppleant for Lamarck. Blainville had keenly desired the post. Efforts were made for Audouin to receive one of the two chairs into which Lamarck's chair was divided after his death. When it was realized by Audouin's supporters, including Cuvier, that Blainville was not to be denied a place in the Museum, Audouin was persuaded to withdraw in favor of the aged Latreille.47 He continued to act as Latreille's assistant and inherited the chair when Latreille died three years later. Through the 1820s Audouin continued to publish memoirs of philosophical anatomy, but within the bounds imposed by Cuvier. By a similar offer of patronage, Cuvier won the support of Flourens, formerly a devoted disciple of Geoffroy.48 Philosophie anatomique had appealed to Flourens because of its boldness and the materialistic tendency of its ideas. Likewise, Flourens, following Geoffroy's lead, was once sympathetically disposed to the German phrenologist Franz Joseph Gall, who was residing in Paris and whose private course Flourens had attended in 1815. (Geoffroy was nearly alone in supporting Gall's candidacy for admission to the Academie in

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1821.)49 Flourens assisted Geoffroy in his research for a second volume of Philosophie anatomique, published in 1822, and in turn Geoffroy lavished praise upon him and sought to establish a section of physiology in the Academie in which Flourens might hope to obtain a seat.50 However, once Flourens began to present his own research in experimental neurophysiology to the Academie, career considerations intervened, and he gradually switched his allegiance. In the years 1822-1828, Cuvier read to the Academie four encouraging reports on Flourens's works, the last two presented in the months preceding Flourens's December 1828 election to the Academie in the section of rural economy. Cuvier's first report, published in several contemporary journals, was largely responsible for publicizing Flourens's work. To suit Cuvier, Flourens modified his terminology in a postivist direction and was amply rewarded with Cuvier's patronage.51 In both 1824 and 1825 he won the Montyon prize for experimental physiology. In 1828 he acted as Cuvier's suppleant at the College de France, and after Cuvier's death he eventually inherited this chair as well as the post of permanent secretary of the Academie. In 1832 he also became professor of human anatomy at the Museum. Under Cuvier's patronage, Flourens's thinking underwent an almost complete about-face. He became a spokesman for final causes, the "positive facts," the four embranchements, and Cartesian dualism. As a popular essayist, he championed these orthodoxies first against Gall and then against Darwin. Academie Politics: The Election of Blainville

The election of Blainville to the section of anatomy and zoology of the Academie in 1825 showed that Cuvier was not always so blessed with fortune in his effort to manage scientific patronage. Despite Cuvier's repeated attempts to thwart the career of his one-time disciple, Blainville was able to prevail in an exceptionally close contest in which he defeated Cuvier's brother Frederic. Partly from conviction and partly from sheer perversity, Blainville had fashioned a doctrine directly at variance with Cuvier's. In the place of the four embranchements, he sought to establish an "animal series," an updated and modified chain of being, which entailed links between the major groups of animals. Blainville's system of classification of the animal kingdom, first published in 1816 and revised in 1822, discarded Cuvier's terminology, employed external characters rather than internal characters to form the major divisions, and deliberately established links between the four embranchements. In particular, he joined the two "types" or subdivisions, articulates and mollusks, by another "type" containing the barnacles and chitons, which he pointedly called "Malentomozoa" or "articulo-mollusks."52 His 1822 classification featured twenty-six classes in descending order from mammals to "Dendrolithaires" the connecting link with the plants. According to his peculiar and much-criticized jargon, vertebrates became "Osteozoa," mammals were "Pilifera" (having hair), birds were "Pennifera" (feathered), reptiles were "Squamifera" (having scales), amphibians (which he erected into a separate class joining fishes

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and reptiles) were "Nudipellifera" (naked skin), and fishes were "Pinnifera" (having fins).53 Blainville was, moreover, at this time favorably inclined toward spontaneous generation and critical of traditional vitalism, and believed that science could only progress by alternating the a posteriori and a priori methods.54 For both personal and intellectual reasons, then, Cuvier opposed any further advances in Blainville's career. Although by no means a follower of Geoffroy, Blainville remained sympathetic to philosophical anatomy, which he claimed to be far superior to and more profound than descriptive comparative anatomy. In discussing the various kinds of anatomy in 1822, Blainville stressed the importance of "philosophical or transcendental anatomy," because it rose from facts to abstractions, and rendered an account of organs throughout the series. "It follows an organ through all the variations it experiences, and recognizes it by general traits such as connections and uses." Its purpose, Blainville added, was not simply to perfect the natural system, but also to advance "physiology or the explanation of the phenomena of life by the application of general laws of nature."55 Despite his falling out with Cuvier in 1816, Blainville was able to obtain a seat at the Academie by winning the support of a rival patron, the astronomer Francois Arago. Throughout its history, the Academie had been under attack for controlling patronage through elitist "coteries." During the 1820s the Academie was the scene of an especially lively struggle for power. Until then, patronage at the Academie had been largely under the control of Laplace, Berthollet, and their proteges. Cuvier had long been associated with this older leadership. Besides being the leader of a faction at the Academie, Laplace was the initiator of a research program, that of explaining all elementary interactions in physics and chemistry by means of short-range forces acting at a distance between particles of matter or between imponderable fluids such as caloric or electricity. In the 1820s, however, the Laplacian program for physics was being attacked on several fronts. Augustin Jean Fresnel's wave theory (1815) and Ampere's electrodynamics (1820) were part of a larger movement among the younger generation of scientists to end the domination of physics by Laplace.56 Corresponding to the battle over concepts in physics was a parallel battle for control of scientific patronage. Arago, professor at the Ecole Polytechnique, director of the Paris Observatory, and co-editor of the influential Annales de chimie et physique, emerged as the leader of the anti-Laplacians. Warm, enthusiastic, volatile, and given to polemics, Arago stood in sharp contrast to the cold, austere Cuvier. So too did Arago's liberal politics differ from Cuvier's growing conservatism. Whereas Cuvier supported the increasingly reactionary ministries of the 1820s, Arago entered politics only after the Revolution of 1830 and reached the height of his political career as a member of the provisional government following the Revolution of 1848.57 During the 1820s the Arago faction took over many of the key posts in the scientific community. Most significantly, in 1823 Fourier, a member of the Arago group, defeated Biot, Laplace's protege, for the position of permanent secretary for the mathematical sciences. Biot, taking the vote as a personal

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rebuff, retired from active participation in scientific affairs for six years.58 In 1830 Arago himself became permanent secretary. The ascendancy of Arago represented a threat to Cuvier, not only in terms of naming candidates to vacant seats, but also in terms of Academie policy. The two scientific leaders disagreed profoundly on the much-debated question of how open the Academie ought to be. Whereas Cuvier and many of the older members wished the meetings to be closed to all nonmembers except for a few specially invited guests, Arago favored more access to the meetings. Since the founding of the Institut in 1795, more and more visitors had been allowed to attend the weekly Monday meetings. In the 1810s most of the nonmembers had been scientists or prospective scientists, some of them representing journals that published brief accounts of the Academie meetings. The presence of increasing numbers of nonscientists had occasioned sufficient alarm by 1825 that closed meetings of the Academie were called to discuss the matter, but no action was agreed upon.59 By the late 1820s the doors had become unofficially open to anyone who cared to enter. Journals published not just titles and brief extracts, but long accounts of memoirs and discussions, as well as the opinions of the science editors. Cuvier strongly disapproved of this type of publicity. The official records that he took of the meetings were scanty and remained unpublished until the twentieth century.60 Arago, however, desired to publish a Comptes rendus, containing an official extract of all papers presented as well as a summary of the discussion following the papers. Cuvier's disagreement with Arago on these issues came to a head in 1830, an episode which will be discussed in the next chapter. It was through the support of Arago that Blainville was able to defeat Frederic Cuvier. On three previous occasions Blainville had tried and failed to win election to the Academie. In 1825, for the seat left vacant by the death of Lacepede, the section of anatomy and zoology nominated Serres in first position and Blainville and Frederic Cuvier in second position. By the third ballot, the candidates were reduced to Blainville and Frederic Cuvier. In an unusually narrow vote, Blainville won, thirty-four votes to thirty.61 Without the backing of Arago, wrote his friend and biographer Pol Nicard, he would never have been able to overcome "the resistance that G. Cuvier was imposing."62 The election of Blainville illustrates that for all Cuvier's power, he was not always able to have his own way. His doctrine, like the Laplacian program, was under widespread assault, and his prestige at the Academie was being eroded. With Blainville and Geoffroy teaching their doctrines to the younger generation at the Faculte des Sciences, Cuvier, who was only sporadically teaching, found himself in a particularly vulnerable position. This in large part accounts for his increasing defensiveness in the 1820s and the care with which he courted the allegiance of bright and promising young naturalists. Serres, "Transcendental Anatomy," and the Theory of Arrests of Development

One naturalist whom Cuvier could not divert from his allegiance to Geoffroy was the embryologist Etiennne Reynaud Augustin Serres. Born in Clairac in

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the Southwest of France in 1786, Serres came to Paris to study medicine and had since 1814 been employed at the Hopital de la Pitie. In 1822 he was named chief physician at this hospital. First attracted to the study of comparative anatomy through attending the courses of Cuvier and Dumeril at the Museum and Faculte de Medecine, he only later made the acquaintance of Geoffroy. The latter, seeking skulls for his research on homologies in 1817, was introduced to Serres for this purpose by Magendie. Serres gladly supplied Geoffroy with the needed materials, and a lifelong friendship ensued.63 Apart from his son Isidore, Serres became Geoffroy's chief disciple. His loyalty to his mentor was unswerving throughout the 1820s, the debate of 1830, and even through the 1830s, when he tried in vain to moderate Geoffroy's by then outrageous pronouncements. Serres provided Geoffroy with skeletons, cadavers, and malformed fetuses from the Paris hospitals, supported and expanded upon his doctrines, dedicated books to him, and dined regularly at his home. In return Geoffroy did his utmost to assist Serres in the transition from medicine to science and to ensure that he would enter the Academie and the Museum. For his first memoir submitted to the Academie, on the "laws of osteogenesis," Serres was awarded in 1820 the Montyon prize for physiology, and for his extensive research on the comparative anatomy of the brain he won the prize for the physical sciences in 1821.64 Geoffroy was a member of the first prize committee and Cuvier of both. In 1819, 1824, and 1825 Geoffroy read three flattering reports on Serres's work before the Academie, which he published in scientific and medical journals.65 On his sixth attempt, in 1828, Serres, with the backing of Geoffroy, was finally elected to the Academie in the section of medicine and surgery, after having been nominated for every vacancy in this section and that of anatomy and zoology since 1821.66 Although Geoffroy tried on several occasions to secure him a professorship, Serres did not begin his career at the Museum until 1839 when he was chosen professor of human anatomy. The goal Serres set himself in his embryological research was to discover and apply general morphological laws of animal development.67 The ensemble of such laws Serres called "transcendental anatomy," a term which contemporaries identified as a synonym for philosophical anatomy.68 His research served to confirm Geoffroy's homologies and to provide a foundation for their joint study of teratology. Development, Serres was concerned to demonstrate, was not simply a process of filling in previously formed parts, as required by the eighteenth-century doctrine of preformation. Rather, he insisted, a true formation took place according to fixed and invariable laws. Of all such laws, the most fundamental was that of "eccentric development." All organs, Serres postulated, formed from the circumference to the center, rather than from the center to the circumference as was commonly held. Nerves, for example, did not arise from the spinal cord, as was thought, but instead formed first in the organs to which they were distributed, and only later inserted themselves in the cerebrospinal axis. Sockets in bones serving for articulations at joints were formed not by a single bone, but by two or more primitively distinct bones which coalesced. Serres's conclusion that such bones as the incus of the middle

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ear, the scapula, and the sternum were originally composed of several elements gave embryological confirmation to Geoffroy's analysis of the same bones in order to formulate homologies of the skeletal system. Related to "eccentric development" was Serres's contention that unique organs of the animal body located along the central axis were always primitively double. All canals and tubes, including the spinal cord, were formed by two distinct halves joining together. The two halves of such organs attracted each other ("law of affinity"), moved toward each other, and finally united ("law of conjugation"). Serres applied the laws of "transcendental anatomy" to solve the prize question posed by the Academie des Sciences in 1818 on the determination of the parts of the brain in the four classes of vertebrates (mammals, birds, reptiles, and fishes). If one were to compare adult forms directly, Serres argued, the task of making an accurate indentification would be hopeless. No homologies whatever could be found in such disparate structures as the brain of a monkey and that of a fish. However, if one examined the earliest stages of the development of the brain in the embryos of the four classes, the brains would appear so similar that the homologies of the cerebral lobes, optic lobes (which Serres determined to be the equivalent of the quadrigeminal bodies in mammals), cerebellum, and medulla could not be mistaken. This initial identity of parts was soon effaced by subsequent development. In each class of vertebrates, a part or combination of parts predominated, causing the remaining parts, by the law of balance of organs, to become almost rudimentary. Such transformations and metamorphoses were all mediated through changes in the number and caliber of the arteries supplying the parts. Like Geoffroy, Serres believed that the circulatory system was in large part responsible for regulating the formation of the other parts of the body. The higher the animal in the scale of beings, the more its brain deviated from its original embryonic form. Here again, as in the case of the laws of bone formation, Serres, by means of the study of embryology, arrived at the same conclusion as was predicted by the "theory of analogues." However varied the brains of adult vertebrates might appear, they were in essence the same and could be reduced to a "unity of composition": Thus in ascending the animal scale from fishes to monkeys you see the encephalon become gradually more complicated, as in descending from the adult mammals through the various periods of their embryonic formation, you perceive this organ successively decompose. You arrive by these 69two ways at the same result: the unity of their formation and their composition.

"The encephalon of vertebrates," Serres concluded, "is therefore constructed on a uniform type and with the same elements."70 Serres's guiding principle in making determinations of the parts of the brain was the "theory of arrests of development" or the recapitulation theory. Implicit in earlier formulations of German biological philosophy, such as that of Kielmeyer in 1793, the theory of arrests of development was clearly enunciated by Meckel in 1811. But Serres did so much to extend its application and promote it that historians have since called it the "Meckel-Serres Law," to dis-

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tinguish it from later evolutionary versions of recapitulation.71 "For certain of its parts," Serres postulated, "the encephalon of lower classes retains permanently the primitive forms of the embryos of higher classes."72 To arrive at an understanding of the "permanent forms" of vertebrates, therefore, it was necessary to follow the "fugitive forms" of their embryos in order to unravel the metamorphoses of parts and uncover the original unity. In his first memoirs, Serres applied arrests of development only to organs and not to entire embryos, but in his later writings he was less careful to limit his theory. He emphasized in his early research on the vertebrate brain that the embryo of a mammal was never actually a fish or a bird. Externally the embryo was always clearly identifiable as mammalian, although internally the nervous system might repeat the encephalic forms of the three lower classes. Already, by 1824, he began to apply the theory to invertebrates. Mollusks, he suggested, were, with respect to their nervous system, similar to the larvae or embryos of insects.73 By 1832 he claimed that the entire human embryo passed through fish-like, reptilian, and avian stages. "The embryogenesis of man," he declared, "thus reproduces in a transitory and short-lived manner the fixed and permanent organization of beings which occupy the various degrees of the animal scale."74 Serres thus clearly enunciated a form of what has become known as the recapitulation theory. Serres, however, interpreted the theory in a transcendental rather than an evolutionary context. His was a twofold parallelism between embryogenesis and the animal scale rather than the threefold parallelism, later popularized by the German Darwinian Ernst Haeckel, of embryonic development, the scale of nature, and evolutionary history. Beginning in 1821, Serres applied the theory of arrests of development to the study of malformations. Geoffroy's and Serres's researches on teratology, although published independently, were very much a joint effort, each contributing to the theoretical framework of the other. Serres's work culminated in a general theory of "organic formations and malformations," published in 1832 and dedicated to Geoffroy: "To the friendship, to the talent, to the virtues which honor the savant."75 The theory of arrests of development provided for Serres, as it did for Meckel, a ready explanation for many forms of monstrosity.76 When one or more organs of a higher animal were arrested in their development, the normally transitory organization of a lower form became fixed in time and rendered permanent. Thus, Serres argued, monstrous organs in higher animals would often reproduce forms found in lower animals. Monsters might also be formed at an earlier stage of development if the two halves of a unique organ were arrested in their movement toward one another, thus creating a fissure through which other organs could protrude. In all cases of monstrosity, Serres insisted, no part was aborted or duplicated unless its artery was also eliminated or doubled. Serres's results challenged Cuvier's doctrines on a number of fronts. He championed epigenesis against preformation, supported without reservation unity of composition, and tended toward the acceptance of a continuous chain of being. Moreover, he became the chief spokesman in France for a central tenet of Naturphilosophie, the transcendental theory of arrests of development.

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Differing with Cuvier on scientific methodology, he defended the right and even the necessity of the naturalist and anatomist to deal in hypotheses and abstractions. His work on the comparative anatomy of the brain, he admitted from the start, was based on a preconceived idea which he subsequently verified through observation. Even a single fact, Serres insisted, was in reality an abstraction, for no description of a thing was possible without a selective comparison to other things. Progress in the sciences, he maintained throughout all his writings, "depends essentially on the combination of facts and ideas."77 From Serres's own account, Cuvier made a concerted effort to win his allegiance, but Serres was not swayed. Introduced to Serres by Geoffroy, Cuvier at first gave him support and advice. He sat on the prize committees which crowned his memoirs at the Academie and presented a favorable report on his research on the anatomy of the brain. This review, which characteristically summarized Serres's results while ignoring his theoretical framework, Serres was pleased to reproduce in the introduction to his published work on the brain in 1824.78 At the same time, Cuvier endeavored to convince Serres to renounce philosophical anatomy. Alfred Moquin-Tandon, a provincial naturalist sympathetic to philosophical anatomy who visited Paris in 1834, left the following account in his journal of what Serres had told him: M. Cuvier gave plenty of advice to M. Serres but included in this advice was that of abandoning philosophical routes in order to follow the most absolute positivism, and to leave M. Geoffroy and attach himself to him. M. Serres was too convinced of the justice of the route he had embraced, and 79too grateful toward M. Geoffroy, to march under the banner of the great Cuvier.

Had Serres succumbed, Moquin-Tandon mused in his journal, he would most certainly have arrived at the Museum. Geoffroy had keenly supported his candidacy for professor of human anatomy left vacant by the death of Portal in 1832 shortly after the death of Cuvier, but the rival candidacy of Flourens prevailed. Moquin-Tandon cynically observed that "a man who is not much of an anatomist or savant" had managed to obtain "one of the finest chairs in Paris by the winds of favor and by the 'tail' of Cuvier's influence. The great man was sleeping in his tomb, but his shade was still giving places to his proteges."80 Eventually, however, Serres did succeed in becoming professor of human anatomy when in 1839 Flourens left that chair for the recently created chair of comparative physiology. Although already in his fifties when he began to teach, Serres held the post for nearly thirty years, during which he continued to profess the same doctrines he had formulated in the 1820s.81 The New Science of Teratology: The Philosophical Anatomy of Monsters

With the aid of Serres, Geoffroy set out to extend unity of composition to a new domain—teratology, or the study of malformations—a field of research that was both exciting and fruitful, but also wide open for speculation on laws of development. As early as his memoirs written in Egypt, Geoffroy referred to

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malformed animals as intermediary forms that could provide evidence for the transformations required by homologies.82 Now, in 1820, he began a series of works concentrating on malformations that culminated in the 1822 publication of a second volume of Philosophie anatomique on human "monstrosities." Although he began his research as a complement to his studies in comparative anatomy, he soon discovered that he had opened a rich mine, where, as his son wrote, he found "a whole new science to create."83 This field was given its current name, teratology, by Isidore Geoffroy, whose three-volume Traite de teratologie (1832-1836) was largely responsible for systematizing the field and giving it a level of respectability in France that his father's publications did not always attain.84 Teratology eventually became an integral part of morphological research and a widely used accessory means of determining homologies. Whereas human "monsters" (to employ the term then in use) had until recently been regarded as aberrations of nature, conforming to no fixed laws of organization, several of the German naturalists, especially Johann Friedrich Blumenbach, had argued that they were merely the result of modifications of the normal forces of nature.85 Geoffroy, one of the first to introduce the scientific study of malformations into France, set about to show that their formation obeyed the principles of unity of composition and of connections, and moreover that they provided an entirely new means of investigating the general principles of organization: I perceived, in fact, a completely different theater, that of organization in its irregular acts, of nature submitted to disturbances, embarrassed in its development [ses evolutions], surprised finally in its moments of hesitation or of powerlessness.86

"Monstrosity exists," proclaimed Geoffroy, "but not, however, exceptions to the ordinary laws [of nature]."87 One of Geoffroy's chief innovations was the attempt to classify monsters based on the supposed cause of malformation. Earlier efforts at classification had not gone beyond such broad categories as "monster by excess" and "monster by defect." For Geoffroy, as for Blumenbach and his students, monstrosities occurred as the result of perturbations in the normal order of development. Because the number of types of perturbations was limited, so must the resulting monstrosities be limited. Thus was created the possibility of a taxonomy of monsters. Indeed, Geoffroy believed that it would be possible to classify monsters using Linnaean binomial nomenclature according to the same natural method used in zoology. The classificatory scheme would by no means be limited to human malformations, but could serve as well for monsters of all the vertebrates. For example, the genus Rhinocephalus (head with a trunk or extraordinary nostrils) could be defined regardless of whether one were dealing with a child, a puppy, a foal, or a calf. His first memoir on teratology presented a classification of acephalic monsters, or monsters charactererized by an imperfect conformation of the head, in which he established thirteen genera. These included such malformations as Coccycephalus (head in the form of a

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coccyx or beak), Cryptocephalus (head invisible externally), Podencephalus (head with brain on a pedicle), Notencephalus (head with brain protruding on the back of the head), and Rhinocephalus.88 In the course of his subsequent work he created some thirty genera. By 1847, nearly fifty more were identified, mostly by Isidore Geoffrey, who arranged them all in twenty-three natural familes and five orders.89 As one might predict, however, the elder Geoffroy was less interested in the mere classification of monsters than in the cause of monstrosity. The study of monsters was especially important to Geoffroy because he believed it could throw light on the still-debated question of whether "germs" were preformed in the embryo or formed by epigenesis. In a footnote to one of his insect papers Geoffroy indicated that the "divergence of our ideas in the too famous question of the pre-existence of germs" was at the very heart of his disagreement with Cuvier.90 Cuvier, as we have seen, had tried to eliminate discussion of the issue altogether while at the same time leaning toward preformation, as had most religiously orthodox scientists of the eighteenth century. Geoffroy, following in the philosophe tradition of Maupertuis, Diderot, Buffon, and Lamarck, opposed preformation from the outset of his career. The issue was raised with particular force at this time because Meckel, whom Isidore Geoffroy regarded with Geoffroy as the cofounder of teratology, was perceived by Geoffroy as a preformationist. That is, Geoffroy interpreted Meckel as believing that the Creator had in the beginning formed monstrous germs along with the normal germs.91 Refuting the doctrine of preformation gave Geoffroy a golden opportunity to turn the tables on Cuvier. He could smugly argue that the doctrine was incomprehensible, hypothetical, metaphysical, and therefore poor science: Pre-existence of germs—these two words, difficult for me to understand even when they are detached from one another, appear to me completely unintelligible when united. They owe their origin to an idea of causality, to the metaphysical explanation of a fact which, it is well known, has neither been observed nor understood. To admit that a germ contains in miniature all the forms that will later be manifested in an organized being, and to develop a theory of pre-existence for so indefinable a thing, is to multiply at will the most gratuitous assumptions.92

Geoffroy declared preformation to be a detriment to the study of monsters, because it discouraged investigators from discovering the true causes of malformation. Monstrosity occurred, Geoffroy believed, when an accidental lesion modified the action of the "nisus formativus" (formative force), a term he borrowed from Blumenbach. For Blumenbach it had denoted an irreducible teleological force, but for Geoffroy it simply meant the tendency to normal development. He hypothesized that an accident induced either directly by a physical shock or indirectly by a severe mental shock could cause the fetus to adhere in one or more places to its placental envelope. "There is no monstrosity that does not depend on a pulling action [tirage]. . . bringing itself to bear on one of the superficial points of the body of the fetus," he wrote.93 The type of monster thus formed depended on where the attachment occurred. Geoffroy claimed to

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have discovered several examples of adherences while examining monstrous births. By assuming that the circulatory system formed the organs in the developing embryo, Geoffroy was able to provide a physiological basis for the formation of monsters. When, by an accident, the embryo adhered to the placenta, arteries would anastomose and circulation would become obstructed. Those organs nourished by the blocked vessels would be unable to develop normally, and would through arrest of development become monstrous. Geoffroy predicted that this explanation for monstrosity, once demonstrated, would settle for all time the problem of preformation of embryos.94 Central to the understanding of malformation was the theory of arrests of development. One of Geoffroy's more debatable propositions was his claim that an organ retarded in its development would necessarily resemble, to a greater or lesser degree, the form of an organ in an animal lower in the scale of being.95 Thus, in 1825 he named a human malformation Aspalasomas from "aspalax," meaning a mole, because the digestive, urinary, and reproductive canals were separated as in the mole.96 In practice, Geoffroy's comparison of human malformations to conditions of organs in lower animals was done in an ad hoc and thoroughly hypothetical manner. Although Geoffroy originally supposed that the formation of monsters by adherences of the embryo and placenta would account for all malformations, in 1827 he limited this explanation to unitary or single monsters, and proposed a new explanation for the other major category of monsters, the double monsters. In his article "Monster," written for a multivolume dictionary of natural history, Geoffroy adopted an explanation for this latter group (which included Siamese twins) that had previously served him for the explication of such types as the cyclops. In the cyclops, Geoffroy had reasoned, the removal of the olfactory organ had eliminated the barrier between the two eyes, which then united because they were similar structures.97 In any double monster, he generalized in 1827, the two beings were invariably united by homologous parts, down to the smallest blood vessels, nerves, and muscles. This phenomenon was only a special case of Serres's law of normal development, whereby the two primitively distinct halves of a unique and central organ united on a median line. In some sense, then, a double monster was composed of four halves rather than two. The law of attraction of similar parts Geoffroy christened in 1829 the "law of soi pour soi" or like for like, and hinted that it was of universal application to both organic and inorganic bodies.98 In the 1830s he explicitly assimilated the "law" to his theoretical system of 1801. Geoffroy's work in teratology was not limited to classification, description, and theorizing, but also involved experiment. He has been justly credited by many historians as the founder of experimental teratology for his repeated attempts (albeit unsuccessful) to produce malformations.99 In 1820-1822 and again in 1826-1827, Geoffroy resumed the experiments with eggs that he had begun in Egypt, with the goal of investigating the effect of the environment on animal organization, and in particular on the process of fetal respiration. In his first set of experiments, he sought to determine whether the fetus took in subtle

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fluids from the atmosphere or gave off fluids through evaporation. Having weighed six eggs at the beginning and toward the end of incubation and finding that the eggs lost 20% of their weight, he concluded that an interchange of fluids between the egg and the environment had indeed taken place. By designing experiments in which he could aid or obstruct the interchange of fluids, he hoped to modify the respiratory process. Thus, he pricked, scratched, and filed eggs, or covered them in whole or in part with linen, wax, gold-beater's skin, or varnish.100 In the manuscript notes of his experiments, he explained that if the air was "an agent of nourishment," covering the eggs in whole or in part by varnish would be "a means of causing perturbations in the development of the chick and procuring monsters at will."101 The results were, as one might expect them to be, inconclusive. Many of the fetuses died, and Geoffroy could not be sure whether the modifications in the nasal maxillaries and intermaxillaries that he observed in three of the surviving chickens were due to the varnish or to some extraneous cause. In a second early set of experiments, Geoffroy attempted to modify respiration in the fetus by tying the oviduct in hens. He wondered whether he might thus produce a uterine incubation as in mammals, but after twelve days, when the hens were killed, he found no trace of an embryo in the eggs. Although his experiments were thus far unsatisfactory, he remained convinced that he had discovered a new and fruitful method for the study of organization: A pathological condition is created at will, the intensity of which can be regulated by the duration of the experiment. And, master of all variables, one can dispose of the problem in such as way as to provide the greatest opportunity for a rigorous solution.102

Whereas in 1820-1822 Geoffroy had performed his experiments using natural incubation, in 1826 he was given the opportunity of carrying out experiments on a larger scale in the incubation ovens in the village of Auteuil near Paris. There he conducted several tests in the hope of creating monsters by mechanical accidents: he soaked eggs in water, incubated them in a vertical position with either the large or the small end up, partially covered them with plaster, or interrupted their incubation by taking them out of the oven for fifteen to forty-eight hours between the seventh and tenth day of incubation. Although he claimed to have created monsters, his results were, at best, highly equivocal.103 Yet the possibility of bringing about malformations in this manner continued to intrigue French naturalists throughout the century to the extent that the question was proposed for a prize at the Academie des Sciences in 1860.104 Cuvier, no doubt, appreciated the considerable anatomical skill and attention to detail that went into the classification and description of malformations, yet he surely must have been disturbed by several features of Geoffroy's teratological research. Geoffroy tended to leap from the description of a single malformed specimen to highly questionable generalizations concerning organization. His claim that the various unitary monsters arose by adherences of placental membranes was exceedingly hypothetical; even more so was his law

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of "soi pour soi" to account for double monsters. His recourse to the theory of arrests of development associated him with the suspect Naturphilosophie and led him to form the most far-fetched homologies between malformed structures in higher animals and normal structures in lower. Finally, his rejection of preformation and his appeal to the environment as a modifier of animal structure smacked of materialism and linked him to Lamarck. If that were not enough, Geoffroy further fanned the flames by the diffuse, egotistical, and polemical presentation of much of this material. He repeatedly rehearsed the results of his earlier researches and defended their novelty and import against all detractors. Scarcely an article appeared without a historical summary of the development of his theories written in order to justify himself in the face of some "unfair" criticism. Despite the very evident provocation, however, Cuvier refrained from a frontal attack on Geoffroy's teratological work. In his annual reports of the work of the Academie he limited himself to describing Geoffroy's monsters.105 A Teratological Theory of Evolution

In the 1880s many young French naturalists, angered at the Academie's reluctance to accept or even to discuss Darwinian evolution, traced French rejection of evolution back to Cuvier and his controversy with Geoffroy in 1830. Yet evolution was scarcely mentioned in the memoirs read by Cuvier and Geoffroy before the Academie between February and April 1830. The debate focused instead on unity of composition, a theory which can and should be separated from evolution, but the latter issue was not entirely absent. As early as 1825, in fact, Geoffroy indicated that, for him, unity of composition might imply common descent. For his part, Cuvier had also, in a sense, linked the two theories by condemning both on the same grounds in his 1825 theoretical essay, "Nature." There is no question that Geoffroy was influenced by Lamarck, but to what extent did he indeed become an evolutionist? Prior to 1825, Geoffroy had neither explicitly adopted special creationism nor mentioned evolution. From the outset of his career, he had obviously been exposed to the evolutionary speculations of Buffon, and it can also be assumed that, as Lamarck's friend, he must have discussed the issue with him. Although he may well have toyed with the idea earlier, there is no reason to assume that Geoffroy was a convinced transformist before 1825.106 Even then, he treated the subject in a hesitant manner, being more concerned with exploring the possibility of evolution than in demonstrating it as an irrefutable truth. Geoffroy's theory of evolution, unlike that of Lamarck, was neither comprehensive nor consistent. His desire in his first two papers on this subject—the only papers published before the debate— was simply to raise the issue of the possible modifying power of physical and chemical agents in the environment. He dealt with this possibility almost entirely with respect to a very specific case: whether Steneosaurus and Teleosaurus, fossils named by him, were the ancestors of the modern-day gavials.

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Thus Geoffrey's theory has been called a paleontological theory of evolution by the historian Franck Bourdier, but in its evolutionary mechanism, it is more fundamentally a teratological theory of evolution. Geoffroy's theory of evolution, if it can be called that, was constructed on the foundation of his earlier work in philosophical anatomy, and especially in teratology. Geoffroy regarded the chief evidence of evolution to be the natural system of classification and unity of composition among the vertebrates. If all vertebrates were built on a common plan, composed of the same organic elements, then he believed it was legitimate to inquire whether the "transformations" required by homologies among the vertebrates might also have a genealogical foundation. Teratology had provided him with a material means of bringing about the transformation of organic pieces. An accident in the course of development led to changes in the distribution of arteries and subsequently modifications of structure. The production of monsters, formed by a violent shock to the fetus, was an extreme case, and the monsters did not usually survive, let alone propagate. But if changes in the environment acted in a more modest way on the development of a normal animal, might not smaller and more viable structural modifications result? Geoffroy's first excursion into evolutionary theorizing, his memoir of 1825, was occasioned by the discovery of a new fossil crocodile, the "crocodile of Caen." Both Cuvier and Geoffroy considered themselves experts on crocodilian anatomy. Geoffroy had recently completed a series of papers on the nasal canal in vertebrates, particularly in the crocodile, when in 1823 Felix Lamouroux, professor of natural history at the Faculte des Sciences of Caen, sent to Cuvier part of a fossil "crocodile" skull and casts of other bone fragments. Cuvier announced the discovery to the Academie and hastily wrote up a description of the bones to be included in the second edition of Recherches sur les ossemens fossiles. In this account, published in 1824, Cuvier classified the bones as belonging to an extinct species of gavial.107 Geoffroy, recognizing, quite rightly, that Cuvier's attribution was inexact, applied his considerable anatomical skill to establish that the "crocodile of Caen" was of a different genus from all known crocodilians. The name Geoffroy bestowed upon the fossil was based upon a supposed structural affinity with the mammals. He claimed that by the structure of its palatine region, the modern crocodile appeared to be an amalgam of the saurian and the mammal. In the "crocodile of Caen," the form of the nasal passages, the bones of the palate, the nares, and the jugals all indicated to him an even closer conformity with the mammals. Thus Geoffroy named the fossil Teleosaurus, meaning "perfect saurian," the name it retains today, since it mistakenly appeared to him to announce the mammals from afar. In 1828 he suggested that the fossil animal formed a link in a progressive series from the swimming and flying fossil reptiles to the fossil mammals: Ichthyosaurus, Plesiosaurus, Pterodactylus, Mosasaurus, Teleosaurus, Megalonyx, Megatherium, Anoplotherium, Palaeotherium, and Mastodon.108 Instead of stopping at this point, Geoffroy took advantage of the occasion to make an inquiry "concerning the degree of probability that the teleosauruses

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and the steneosauruses, animals of antediluvian ages, are the source of the crocodiles dispersed today in the warm climates of the two continents."109 This "probability" was argued on the basis of highly specialized anatomical research. In fact, Geoffroy went out of his way to display his technical expertise. He believed that the "posterior palatines" (now called pterygoids), broad bones of the palate surrounding the nares in modern crocodiles, had reached their maximum of development in the crocodile. These pieces, whose homology with the pterygoid processes of the sphenoid in man Geoffroy pointed out, supposedly dominated and affected the forms of all the surrounding pieces. In great detail Geoffroy sketched out the modifications of the bones of the palatine region in passing from Teleosaurus to the modern crocodiles. These modifications, Geoffroy noted with satisfaction, were already slightly apparent in a mummified crocodile skull he had brought back from Egypt.110 The question of the descent was thus reduced to the question of how the bones of the skull of fossil crocodiles might have been transformed into those of modern crocodiles. The mechanism Geoffroy proposed was based upon the power of the environment to act directly upon the structure of the developing fetus. The principal differences between fossil and modern crocodile skulls, according to Geoffroy, centered on a modification of parts concerned with respiration, digestion, and the nervous system. All of these essential functions were claimed to be highly susceptible to the "pathological influence" of changes in the environment. Geoffroy pointed out that physicists and geologists did not doubt that great changes had taken place in the physical condition of the globe. If the physical and chemical properties of the environment remained unchanged, Geoffroy believed that animal forms would remain stable. If, on the contrary, the environment was modified "without breaking the vital action" (and thereby causing death or extinction), it necessarily caused the organisms submitted to its influence to vary. That which took a considerable time in nature, Geoffroy added, could be seen before our eyes in accidentally or voluntarily provoked monsters. In 1828 he claimed to have deliberately undertaken his teratological experiments at the incubation oven at Auteuil in order to obtain support for epigenesis and for evolution.111 In his paper of 1825, for the first time, Geoffroy drew attention in print to Lamarck: "These changes [in the environment] are of a nature to have acted on the organs of which I have just spoken, and to have done so precisely according to the two laws posed by M. de Lamarck in his Philosophie zoologique." Geoffroy was referring to the two "laws" in Lamarck's chapter on the influence of the environment whereby Lamarck had maintained that use strengthened an organ and that such modification was inheritable. In a footnote, Geoffroy recommended Lamarck's philosophical reflections preceding his laws "to the meditation of young men."112 However, he explained that although correct in general principles, Lamarck's proposed explanation for diversity (presumably the action of environment on the unconscious will of adult animals) was wrong. Variations were instead due to environmental changes acting upon the fetus and causing changes in the distribution of arteries in a manner analogous to that which occurs in human malformations.

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Geoffroy did not return to the subject of the modifying power of the environment until three years later when he and Serres were called upon to present a report to the Academie on a memoir of Francois-Desire Roulin, a physician and naturalist. Once again, Geoffroy exploited the opportunity afforded by the report to advance his own doctrines, which were far bolder than those of the author of the memoir. Roulin had described how domestic animals transported from the Old World to the New World and then returned to the wild had regained some of their presumably original traits: the variegated coat colors in horses and pigs returned to a uniform color, the courage of the ass reappeared, and the petulance of the goat and the rapidity of its gait increased. Instead of simply recounting Roulin's conclusions, Geoffroy took them for a point of departure to attack the doctrine of preformation and to speculate on evolution. He hypothesized that if the nisus formativus were submitted to the influence of even more powerful "resistances" than those acting on Roulin's domestic animals, then "another zoological system" than that extant today might be formed. These "resistances," he contended, had in fact occurred during the past history of the earth. Geoffroy took it for granted that all were agreed that the atmosphere and temperature of the earth were once different, and so too were the animal inhabitants. "Thus, resistances of the environment would have formerly weighed more heavily on the same organic elements [fonds organiques]." Roulin's memoir suggested to Geoffroy an analogous situation to that in which "lost animals" might have become the ancestors of animals of the present age. At this point—after he had already said more than enough—Geoffroy halted, stating that he would not continue with reflections from which Roulin had "the wisdom to abstain."113 In response to the inevitable criticism of his report on Roulin, Geoffroy continued his reflections in a second memoir on evolution published in 1828. There he refused to commit himself on whether extinct animals were the ancestors of modern animals on the grounds that geological knowledge was as yet insufficient. He claimed that he merely wished to open discussion on the matter, whereas the "book on fossil bones [Cuvier's Recherches] maintains that all minds are already enlightened on this point."114 In this second memoir, the role of Geoffroy's Deistic assumptions in leading him to entertain the possibility of evolution becomes evident. The only apparent alternative to evolution—a series of creations at different geological times—was one was that posed great difficulties to someone who envisioned God as First Cause. Cuvier, as we have seen, had not committed himself to new creations after each catastrophe, but he certainly tended toward the acceptance of this controversial doctrine. To both Geoffroy and Lamarck (and to Blainville and many other nineteenth-century naturalists), the notion of God intervening at various times in the uniform operation of nature was thoroughly abhorrent. Although Geoffroy claimed to be a good Catholic, he, like Lamarck, limited God's role in nature to the creation of the various kinds of matter and the establishment of universal laws. The notion that "the work of the six days" could have been resumed appeared to Geoffroy to be "contrary to the best historical data" and "as repugnant to the light of reason as it is to the more reflective speculations of the physical sciences."115

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Geoffroy introduced in this paper a singular argument, which reveals with particular force his divergence from Cuvier on the issue of scientific methodology. It was to reappear throughout Geoffrey's philosophical writings of the 1830s as the doctrine of "necessary facts." Geoffroy claimed that people feared to entertain the possibility of evolution because of hestitation to go beyond immediate facts. The evidence, he admitted, did not yet support evolution. But a naturalist need not be limited to facts. If the facts were supplemented by the resources of analogy and induction, then the genealogical relation between "lost" and current species could be readily intuited, given that all species fitted into the same classificatory system and were seen to be modifications of the same abstract type. Recourse to analogy and induction where facts did not suffice, Geoffroy claimed to be "the right and property of genius." He produced several examples of geniuses of the past who had divined great truths that they were as yet powerless to prove. Buffon, for example, had correctly perceived that the inhabitants of the New World always differed from those of the old world, but he supported this insight with facts none of which were now admissible. Similarly, Geoffroy judged Lamarck to have been right in his general statements regarding the influence of the environment on animal organization, yet hopelessly wrong in his examples.116 This methodological argument, raised countless times in the 1830s, was Geoffroy's rejoinder to the "positive facts." Evolution, in effect, was required by Geoffroy's world view, even though he lacked the factual data to demonstrate it. For Lamarck and Geoffroy, then, evolution provided a necessary alternative to extinction and renewed creation, but from there on, their evolutionary doctrines differed.117 Geoffroy was inclined to view evolution as the result of the direct and sudden effect of the environment acting on the fetal development of animals, while Lamarck believed that the environment exerted its effects slowly and indirectly upon the adult organism. Geoffroy could accept neither Lamarck's innate tendency to perfection nor the active response of the organism to a modified environment. The environment instead acted directly on the fetus, causing changes in the size and branching of arteries, and hence in the size and arrangement of organic elements produced by the arteries. Lamarck, a uniformitarian in geology, assumed transformation of organisms had taken place at a slow and steady pace, but Geoffroy, a catastrophist, suggested that evolution had occurred more rapidly in the past because in the past environmental change had been more abrupt. Geoffroy's most general treatment of evolution did not appear until after the debate of 1830. Fossil crocodiles were not an issue in the celebrated series of memoirs delivered before the Academie from February to April 1830. But soon afterwards Geoffroy made two trips to Caen and submitted a series of five memoirs on the subject to the Academie in 1830 and 1831. The first, second, third, and fifth of these memoirs continued the technical anatomical argumentation. He concluded from his investigation that Steneosaurus and Teleosaurus were intermediaries between Ichthyosaurus (a fossil swimming reptile) and the modern gavial. At Caen he was able to obtain an entire cranium of a Teleosaurus, and he noted with satisfaction that the animal had superimposed scales

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as in fishes and the pangolin. He still lacked evidence of the structure of the foot, but predicted that the limb, when found, would be closer to a fin than to a clawed foot.118 The fourth paper, "Memoir on the Degree of Influence of the Environment [monde ambiant] in Modifying Animal Forms," was entirely devoted to generalities and is Geoffroy's best known contribution to evolutionary theory. In his schema of the progress of zoology with which the paper began, he sought to place his evolutionary speculations in perspective and invest them with an air of inevitability. In the fourth era of Geoffroy's historical introduction, naturalists were concerned with classification and description, in the fifth era with Cuvierian comparative anatomy (i.e., the study of differences), and in the sixth with unity of composition. Now, in the seventh era, the time had come to consider differences again, but in a new light. Differences in structure were to be related to changes in the environment. Unity of composition stemmed from the original arrangement of organic substances, while variety Geoffroy attributed to the subsequent action of the "monde ambiant." Geoffroy here expanded upon his previous speculations on the role of the atmosphere in mediating organic change. Over the centuries the "energy" of the oxygen in the atmosphere had decreased, so that animals constructed so as to breathe the air of previous geological eras could no longer survive: When it happens that respiration becomes difficult of execution and finally impossible for certain systems of organs, it then necessitates and creates for itself another arrangement, perfecting or altering the pulmonary cells in which it operates, [producing] fortunate or fatal modifications which are propagated and influence all the rest of the animal economy. If these modifications bring on harmful effects, the animals that experience them cease to exist, to be replaced by others with forms a bit altered and changed to agree with the new circumstances.119

This passage may sound vaguely Darwinian, but Geoffroy envisioned no struggle for existence. Modifications of animals were produced by the environment in the same way that monstrosities were generated. Some modifications resulted in survival and propagation, others in death. No competition for resources or natural selection was involved. Geoffroy was primarily concerned with the transformation from one species or genus to another closely related species or genus, but at one point he did speculate on how a more substantial change—from one class to another— might have been brought about. Here he called into play the theory of arrests of development. The environment and, in particular, the quantity of oxygen in the air, might have acted upon the developing fetus of a lower animal, creating all at once the inheritable characteristics of the next higher class. He offered this hypothetical example: Let the pulmonary sac of a reptile in the early stages of development undergo a constriction in its middle, so as to separate all the blood vessels in the thorax from the base of the pulmonary sac in the abdomen. That would be a circumstance favoring the development of the entire organization of a bird. The air in the abdominal cells will be pushed back by the muscles of the abdomen so as to direct upon the respiratory vessels air that would then be compressed and of the quality

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of that arising from bellows, that is to say, air containing more oxygen in a smaller volume and therefore producing more energy during combustion. The remainder of the phenomena are merely effects of this first change—the greater caloricity of the blood, its more vivid color, its augmented transparency, its more rapid course, the increased energy of muscular action, the change of tegumentary tufts into feathers, etc.120

It should be emphasized once more that Geoffroy did not work out a complete or consistent theory of evolution on the order of Lamarck's. He provided no evolutionary trees of the classes, and aside from fossil crocodiles, offered few particular examples of evolution. He was more concerned with proving that the subtle fluids of the environment had the power to modify a common plan of animal organization than with insisting that all animals had been derived from one another. His followers, even his most devoted disciple, Serres, did not regard evolution as a necessary consequence of philosophical anatomy. They had no difficulty in separating the two theories, and ignoring the former. Yet the specter of transmutation of species had been raised and injected into the controversy between Cuvier and Geoffroy. Although philosophical anatomy and evolution were theoretically separable, Geoffroy had linked them, and Cuvier, for his own purposes, reinforced the linkage. Naturalists taking a morphological approach thus were careful to differentiate themselves from Geoffroy, for an unqualified adherence to unity of composition could leave them vulnerable to suspicion of the heresy of evolution. Evolution, though scarcely raised during the debate of 1830, was, because of Geoffroy's papers of 1825 and 1828, and Cuvier's indirect but agitated response to them, indeed at issue.

Two Divergent Views of Nature

By 1825 most of the French scientific community had been alerted to the growing dissension between Cuvier and Geoffroy. After that date they could scarcely help being aware of the gulf that separated the two naturalists, because from 1825 to 1829, regardless of subject, nearly every article, book review, dictionary contribution, or eloge that either of the two men wrote was replete with polemics directed at the other. Cuvier responded to Geoffroy's evolutionary speculations by his celebrated article "Nature," written for the Dictionnaire des sciences naturelles in 1825. Geoffroy replied with his own "Nature" for another multivolume dictionary, the Encyclopedie moderne, and with his published lectures at the Museum, Cours de l'histoire naturelle des mammiferes. Cuvier continued his side of the battle by devoting the greater part of the introductory material for the first volume of his monumental Histoire naturelle des poissons (1828) to a review of his objections to a host of what he regarded as false systems.121 Out of all these pages of rhetoric emerged two wholly conflicting views of nature. Of all Cuvier's writings, none is more significant for the historian than his

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essay "Nature," for it is there that he presented the very essence of his biological philosophy. This article was one of the few places where Cuvier willingly discussed the metaphysical assumptions implicit in his work, and admitted that his repudiation of certain scientific theories was motivated by religious as well as by scientific considerations. Cuvier defined nature in the light of his teleological and religious preconceptions. Then, one by one, he examined the speculative theories of his day, condemning them at a sweep as deductions, based on a dangerous misunderstanding of this definition. The single false premise that lay at the root of all these objectionable zoological theories was the belief that nature had an autonomous existence apart from God. In all common definitions of nature, Cuvier began his argument, there is presupposed a Creator and Governor of nature. Only through false application of figurative language have "puerile" philosophers attempted to endow nature itself with activity, as though nature possessed its own wisdom and power independent of the Creator. Such a mistaken conception of nature, in Cuvier's opinion, still prevails, not only in our language, but in the systems of a great number of writers who, willingly admitting the justice of the distinctions we have just made, nonetheless blindly allow themselves to be led toward doctrines based on no other foundation. Such are the scale of nature [chain of being], the unity of composition of organisms, and others similar to these, which have all been invented as the result of the belief that nature is distinct from the Creator and less powerful than He is, and find no evident support but in those fancied limits which they place on his power.122

God's freedom to create, Cuvier held, was subjected to the "conditions of existence"—it would be impossible to create a being without the proper structures to fulfill its needs—but any other restriction would needlessly and unreasonably place fetters on His activity. The first "system" that Cuvier implicated was one that he had combated since the beginning of his career—the chain of being. Intermediaries between the classes had never been found, he insisted, and moreover they could not exist. Cuvier here reiterated his functionalist credo that "Every living being possesses . . . all that which it requires for its existence." A transitional form— "a being which would have one part of its organization allied with another part suited for a different being"—was simply inconceivable.123 Doctrines like the chain of being would have the Creator form organisms not for their own sakes, but only for the sake of an abstraction. "What law is there," he inquired, "which could force the Creator to form unnecessarily useless organisms simply in order to fill gaps in a scale?"124 Unity of composition was rejected for the same reasons. For the first time, Cuvier explicitly rejected the doctrine as a whole, linking it with an extreme of evolutionary speculation. Unity of composition was depicted not as an independent doctrine, but a consequence of a belief in the development of all animals from a single form: However, to the hypothesis of a continuous scale of organic forms, other philosophers have added another, that all beings are modifications of a single one, or

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that they have [all] arisen successively by development from a single primordium. And it is upon that [hypothesis] that they graft that of an identity of composition of all [beings].125

Like the chain of being, unity of composition limited the power of the Creator to create new organs as He saw fit by forcing Him to resort to a common fund of materials in the composition of animals. Finally, Cuvier drew fire on "those who in recent years have sought to offer a new form of the metaphysical system of pantheism, which they call Philosophy of nature."126 To the first two hypotheses, Cuvier claimed, some naturalists had added a third, that of repetition of the cranium by the rest of the body. Once again he recalled to mind Oken's theory in order to ridicule the entire enterprise of seeking homologies within the parts of a single individual. All of these theories, Cuvier concluded his sermon, were based on arbitrary assumptions that would fall apart the moment they were examined in detail. In vain did the authors of these systems shroud them in figurative language, for their only rational basis was the false belief in a nature independent of the Creator and limiting His action: In fact, if we look back to the Author of all things, what other law could actuate Him but the necessity of providing to each being whose existence is to be continued the means of assuring that existence? And why could He not vary His materials and His instruments? Certain laws of coexistence of organs were therefore necessary, but that was all. For to establish others there must have been a want of freedom in the action of the organizing principle, which we have shown to be only a chimera.127

Thus, despite Cuvier's repeated protestations that natural history was a science of facts, his condemnation of the doctrines of his opponents involved metaphysical considerations quite apart from the facts. He rejected these theories in part and on his own admission because he perceived in them irreligious, pantheistic tendencies. Were the theories that Cuvier implicated logically connected? To a great extent Cuvier's confounding of them was a polemical ploy. For by insisting that each heretical hypothesis led to the next, he could render the entire package so preposterous that he could easily dispose of it all. We tend to group these theories today because what we now recognize to be true in each of them can be accounted for by organic evolution. Historically, however, these hypotheses were not always linked. For most naturalists who subscribed to these beliefs, the chain of being (as modified by Blainville), unity of composition, and the theory of arrests of development did not imply evolution. As historians, we must be careful to treat each of these theories on its own terms. In other publications, Cuvier combated the same congeries of theories by forcefully reiterating his stand that the role of the naturalist should be limited to the discovery of facts. The opening sentence of Histoire naturelle des poissons hammered home the point: "Natural history is a science of facts."128 And in October 1829, only months before the debate in the Academie, in a memoir delivered before the Academie on a subject so seemingly foreign to the contro-

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versy as the description of a new intestinal worm, Cuvier felt compelled to interject: The metaphysicians who amuse themselves by composing intestinal worms. . . from the elements furnished by the bodies of animals that they inhabit will have a fine time on this occasion. Here we have the body of the octopus which has for a parasite a worm so similar to the arm of the octopus that the illusion could not be greater. . . . Let one judge of how many systems it would be possible to found on such extraordinary resemblances. Never has the imagination been able to exercise itself on a more curious subject. For us, who have for a long time made a profession of limiting ourselves to the reporting of positive facts, we restrict ourselves today to making known, as exactly as possible, the exterior and the interior of our animal.129

In this particular case, however, the last laugh was on Cuvier. He would have been mortified to learn that researchers later in the century would discover that his "intestinal worm," Hectocotylus, was no worm at all, but the actual arm of an octopus. In certain cephalopods, it was determined, the arms or tentacles of the male become modified as sexual organs for the impregnation of the female. Here is a good example of a case in which a supposed observational fact was thoroughly theory-laden. Geoffroy later interpreted Cuvier's paper as "the prelude which lowered the barriers of the arena where the combat of 1830 was engaged."130 Geoffroy, not one to take criticism idly, countered with his own methodological precept: that any naturalist worthy of the name must look beyond the narrow circle of facts to theoretical constructs. The week after Cuvier read his memoir on the "intestinal worm," Geoffroy responded via a report to the Academie on the coming visit of the original Siamese twins to France. After explaining once more the formation of double monsters by the law of similar attraction, Geoffroy pointedly remarked: We will add that after the establishment of positive facts, it is necessary that their scientific consequences be drawn, just as after the completion of cutting stones the hour must come to work them up. Otherwise what fruit can be gathered from these materials?131

"Anatomy was for a long time descriptive and particular," Geoffroy proclaimed, "[but] nothing will stop it in its tendency to become general and philosophical."132 Cuvier's most sustained refutation of "unity of composition" and the "theory of analogues" before 1830 appeared in his lengthy introduction to Histoire naturelle des poissons. For the most part, Cuvier lashed out at his opponents in general terms, avoiding mentioning them by name. As in "Nature," he failed to distinguish their doctrines clearly and give them their due. This was not a scholarly consideration of issues, but a defensive harangue. In defining the class of fishes, Cuvier made clear his teleological bias, for he deduced the basic structure of the fish from a consideration of the functions it was intended to fulfill. Because the fish was destined to live in water, Cuvier reasoned, it had a form which minimized resistance of the body moving

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through this fluid. Since it had to respire by means of water, it did not require the same level of vitality as did birds, and was therefore provided with a smaller brain. Since the fish could only feed by pursuing and engulfing prey which also swam, a delicate sense of taste would be useless.133 The entire conformation of a fish, Cuvier boasted, could be deduced from the necessity of fishes to live in water. They had their preordained place in nature and were created with only their own needs in mind: That is their assigned place in the creation. They have been there since their origin; they will remain there until the destruction of the current order of things, and it is only by vain metaphysical speculations or by very superficial comparisons that some have sought to consider their class as a development, a perfecting, and an enobling of that of mollusks, or as a first sketch or a fetal state of the other classes of vertebrates.134

Whereas earlier Cuvier had refrained from making a categorical judgment, by 1828 he was determined to demolish the "theory of analogues" by demonstrating that there was no identity in number or arrangement of pieces, and not even the appearance of a progression from fishes to mammals. He attributed bones to fishes—namely the suborbital and the opercular bones—which he declared were nonexistent in the other classes of vertebrates.135 Moreover, he denied Geoffroy's contention that a strict comparison of the bones of the cranium in fishes with the bones of the cranium in the mammals was possible, for any such comparison would require transformations that were repugnant to common sense. Entire organ systems would have to be reversed; bones belonging to one organ in one class would need to be interpolated between bones belonging to an entirely different organ in another class. Cuvier assumed that if the transformations were not physically conceivable—if one set of bones could not be topologically remolded into the other—then homologies were meaningless. This was a restrictive view not shared by any of those who wrote on philosophical anatomy. For the first time, Cuvier explicitly repudiated Geoffroy's key homology— that between the bones of the operculum in fishes and those of the middle ear in mammals. Cuvier found absurd the supposition that the bones of the middle ear should become smaller and simpler from the mammals to the reptiles and then suddenly enlarge and take on an entirely new function, that of protecting the gills, in fishes. Such resemblances as did exist between the organs of fishes and those of other classes, he concluded, depended solely on resemblance of function.136 Geoffroy responded to Cuvier's "Nature" with an article bearing the same title, and to the Histoire naturelle des poissons with his own Cours de I'histoire naturelle des mammiferes. In both works, he argued that unity of organic composition was not an a priori construct, but a demonstrated law of nature of the same character as Newton's law of gravitational attraction. It would soon take its place, he confidently predicted, "among the small number of deductions and intellectual riches which today constitute the treasury of the human mind, and that we have dignified with the name of general laws of nature."137 To Cuvier's charge of irreligion, Geoffroy rejoined by accusing Cuvier of

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unfairly employing a religious objection against a scientific doctrine. Geoffroy's was a typical French response to explicit efforts to mix science and religion. While he reaffirmed his faith in the Christian God and the immortality of the soul, he avowed that he refused to mix mysterious matters of faith with his work as a scientific investigator: "That would be to place the data of the religious spirit, which is a matter of faith, among the discoveries of the scientist [physicien], who has to form his conclusions only by the witness of his senses."138 Truth, he warned, could not be opposed on religious grounds. Although Geoffroy held that God theoretically had the power to "interrupt and change the natural course of things," he doubted that God would in fact use this power.139 God in principle could interfere with the general plan of vertebrates to create new bones for new needs, he admitted, but only "if He wished to produce in a manner that is capricious, monstrous, destructive of His laws, in short, to do a miracle." In that case, science would be reduced to "a philosophy of exception."140 It seemed patently absurd to Geoffroy that Cuvier could number the cranial pieces of fishes from one to thirty-three, find homologies in the higher vertebrates for twenty-five pieces, and insist that the remainder of the bones were created ad hoc just for fishes. Just as there was no exception to Newton's law, reasoned Geoffroy, there could be no exception to unity of composition. Both laws were "equally a result of the will of the Creator and neither compromise nor wound any sort of liberty."141 While reiterating his belief in God's Providence, Geoffroy objected to Cuvier's practice of deducing structure from functional requirements, as exemplified in his history of fishes. Such arguments were "an abuse of the philosophy of final causes." To reason from the premise that since a bird flies through the atmosphere, it must be accorded hollow bones, feathers, and enlarged anterior extremities, he argued, would be similar to concluding "that a man who made use of crutches was originally destined to the misfortune of having one of his legs paralyzed or amputated."142 Function was for Geoffroy always dependent upon structure, and by no means sufficed to determine structure. Finally, Geoffroy undertook to defend German Naturphilosophie as best as was in his power against Cuvier's assaults. Although the German school sinned through overuse of imagination, Geoffroy found the French school of positive science to be equally at fault for its overcaution. Whatever their faults, the Germans at least had the merit of inspiring reflection on high philosophical issues. If their theories were wrong, others would refute them, and science would benefit. "If the pace of the philosophers of nature leans in general towards too much haste," he wrote, "does not the properly regular but always fearful and measured course of their rivals, devoted to the cult of positive science, likewise have its disadvantages?"143 By the end of the decade, the doctrines of Cuvier and Geoffroy had become polarized. The divergence between the two most noted naturalists of the age, it had become clear, was not limited to a simple matter of anatomy, but dealt with the most fundamental issues of science. Where Cuvier held that function determined structure and that God's freedom to create new means for new ends must be preserved, Geoffroy persisted in establishing homologies that

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were independent of function. Where Cuvier saw unbridgeable gaps in nature, especially between the four embranchements, Geoffroy found links either through homologies or through the recapitulation theory. Where Cuvier wished to retain a vital force and "pre-existence of germs," Geoffroy sought to discard all forms of vitalism from science and to demonstrate by experimental teratology that modifications in the course of development could be produced at will. By refusing to restrict God's action, Cuvier denied in effect the possibility of laws of organization (aside from correlations of function). It had been the goal of Geoffroy and Serres to determine such laws through the study of embryology and teratology. Finally, Cuvier rigidly held to fixity in nature, while Geoffroy had begun to explore the possibility of a limited transformism. Added to clash over doctrine was an equally important clash over methodology. Cuvier was fighting a rear-guard action to stave off unwanted theorizing by an insistence that natural history be limited to facts. Few shared Cuvier's fear, and most agreed with Geoffroy and Serres that facts alone were sterile, and that science, to be fruitful, required the construction of theories. Finally, the controversy was greatly exacerbated by personality clashes, institutional politics, and the uncertainties of national politics. As both men were at once vain and oversensitive, they were unwilling to defuse the conflict, and instead pushed it ever farther. Geoffroy responded with ever greater boldness and Cuvier with increasing rigidity. But to the younger generation, the battle between the two older men was not a mere matter of scientific interest, for it could affect the course of their careers. A battle over concepts had also become a battle for disciples. Younger men found themselves caught in the middle, afraid to speak their mind for fear of becoming embroiled in a feud they could not control. Through the decade, Cuvier, spurred on by the political events in France and the difficulties of maintaining his hegemony over the scientific community, as well as personal unhappiness caused by the death of his only daughter in 1827, shifted from at least a partial openness to new ideas, to a defensive stance against all biological theories which did not agree with his own. As a result, both Geoffroy and Cuvier found themselves in extreme and tenuous positions. By 1829, all the issues of the debate of 1830 had surfaced. The historian can easily reconstruct the battle lines, yet for contemporaries the issues may not yet have been brought to a clear focus. This was the role of the debate in the Academie. If it were not for the public debate, the skirmishes of the 1820s might well have been forgotten by later generations. Geoffroy had long welcomed a public battle, and on more than one occasion tried to provoke it, but Cuvier had thus far resisted temptation and avoided it, knowing that his best defense was that of indirection. But tensions had been steadily rising, and in retrospect, it is easy to see that at any time a precipitating event could turn a series of petty quarrels into an open confrontation before the Academie.

CHAPTER 6

The Debate before the Academie

In 1829 Christoph Heinrich Pfaff returned to Paris after an absence of twentyeight years. When he visited the Museum in search of his childhood friend Cuvier, he was surprised to discover a great new wing (the galleries of comparative anatomy) attached onto Cuvier's once modest residence, and an elegant valet in place of the old domestic. Cuvier, who had over the years grown portly, was now to be addressed as M. le Baron. Pfaff found that his former friend had little time for him, or for that matter, for anything else, save politics. His only topic of conversation was the newspapers.1 Cuvier assured Pfaff that France had settled down, and that there would be no more revolutions. He was mistaken. That very summer, the king, Charles X, brought about the fall of the moderate Martignac ministry and replaced it with one more to his liking, the reactionary Polignac government.2 The rise to power of a ministry with little support in the Chambers put Paris in a turmoil and led to the organization of Republican groups. While Cuvier may well have distrusted the ultra-Catholic policies of the new ministry, he nevertheless supported it, as he had supported every government in the past. In March 1830 the Chambers met and immediately lodged a protest. As a retaliatory measure, the king determined on 16 May to dissolve the Chamber of Deputies and to call for new elections in late June or early July. In the voting, the opposition won an overwhelming majority. Charles X reacted with the despotic "Four Ordinances" which would dissolve the new Chamber, revise the electoral system, and suspend freedom of the press. When the Moniteur published the ordinances on 26 July, the insurrection began in the workers' sections of Paris. The revolutionaries were soon in charge of the city. However, the deputies were able to prevent the establishment of a republic by assuming control of the movement and inviting Louis-Philippe, Due de Orleans, to come to Paris and be king. On 2 August Charles X officially abdicated and the Revolution of 1830 was over. To most Parisians, the Revolution had saved the cause of liberty. It was during this period of political upheaval that Cuvier and Geoffroy engaged in scientific debate before the Academie des Sciences. Despite the build-up of animosity between Cuvier and Geoffroy, the onset of the public debate caught contemporaries by surprise. Goethe, who wrote two 143

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articles to inform his countrymen of the significance of the event, likened it to the discharging of a Leyden jar.3 For ten years, Geoffroy had been unsuccessful in provoking the leader of French naturalists into a public discussion. Now suddenly Cuvier abandoned his former caution, rose to the bait, and attacked the doctrine of unity of composition openly before the assembled members of the Academie and its numerous visitors, among them the representatives of the press. Cuvier's jump into the fray might be attributed to a number of causes, including Lamarck's death and the machinations surrounding his replacement, the prospect of Blainville's becoming professor at the Museum, and the rise to power of Arago. But perhaps the impending Revolution was the critically important factor, for given Cuvier's anxiety about uprisings, the political instability of those months before the Revolution of July might have made it imperative that he once and for all destroy the basis of theories that he regarded as a threat to the well-being of society. The significance of the debate was not superficially obvious. It consisted of less than two months of repartee before the Academie, and appeared limited to technical anatomical issues. The arguments put forth by Cuvier and Geoffroy in their memoirs were by no means new. Most if not all of them had already surfaced during the encounters of the 1820s. However, as we have seen, behind the basic disagreement over anatomy lay a wide range of broader issues, of great significance for the future of biology in France and elsewhere. It was the function of the debate to take these issues out from the circle of savants, to bring them to a focus, and to air them before a broader educated public. It is because of the public debate that the controversy between Cuvier and Geoffroy has come down in history. The debate transformed an amorphous series of skirmishes into a clearly circumscribed event, one that could be analyzed, publicized, and politicized. Accounts of the debate usually declare Cuvier the victor.4 It was true that Cuvier had the better of the immediate logical argument. But his anatomical doctrine by no means vanquished that of his opponent. As the debate made clear to contemporaries, on the main anatomical issues neither Cuvier nor Geoffroy had chosen tenable positions to defend. On the one hand, "unity of composition" in the animal kingdom could neither be defined nor demonstrated. But on the other hand, the principle of "conditions of existence" was wholly inadequate to account for the affinities between animals. Most naturalists were willing to admit homologies that could not be explained by similarity of function. Cuvier and Geoffroy had taken extreme positions, and it was left for the next generation of naturalists to reconcile them. As the antagonists and their differences of doctrine were already so well known, and because of the emotional and personal intensity of the conflict, the events at the Academie received widespread publicity. Through attendance at the meetings and through reading weekly analyses in the newspapers and journals, the lay public was fully apprised of the issues. Geoffroy's small volume, Principes de philosophie zoologique, containing the transcript of the debate, brought the issues before savants abroad, and was read with great interest in the 1830s by such men as William Whewell and Charles Darwin. Goethe, the

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most celebrated commentator on the debate, introduced the issues to his countrymen in Germany. The official debate before the Academie closed on Geoffroy's initiative in April 1830, but the dispute by no means ended there. It was carried on unofficially by both participants in the Academie, in the journals, in the daily press, and in Cuvier's course at the College de France until Cuvier's unexpected death on 13 May 1832 closed this phase of the controversy. Finally, the debate became a focal point for those who were dissatisfied with the overly centralized, conservative, and elitist French scientific establishment. While most of the critiques of the scientific establishment appeared after 1832 and will be discussed in the next chapter, Goethe's articles are representative of many of the themes to come. He and others saw in the debate a liberalizing force that would open the Academie up to free and open discussion where once matters were "referred to secret committees, arranged and got rid of, and smothered behind closed doors."5 The debate represented to such interpreters the challenge of new and synthetic ideas to the outworn credos of the past. It is through this common link with liberty and freedom of thought that the connection between the debate and the Revolution of 1830 was more than coincidental. Academicians in Debate

The immediate cause of the confrontation between the two celebrated academicians was a paper on mollusks sent to the Academie by two unknown naturalists, Meyranx and Laurencet. Pierre-Stanislas Meyranx, the main author of the memoir in question, had received a medical degree from Montpellier and was teaching natural history at the College de Charlemagne. His coauthor was so obscure that historians have been unable to identify his first name. Following the customary practice of young men embarked on a scientific career, Meyranx and Laurencet had carried out independent research which they determined to present to the Academie in the hope of obtaining recognition and perhaps a patron. In this case, they got more than they bargained for. In October 1829 they submitted their memoir, "Some Considerations on the Organization of Mollusks," to the Academie. For six months they received no response. Then, tired of waiting to read their memoir, they requested instead that it be examined by a commission. At the meeting of 8 February 1830, Latreille and Geoffroy were named to prepare a report.6 Geoffroy had reason to be excited by the contents of the memoir, for the two naturalists had proposed a way to bridge the gap between the vertebrates and another of Cuvier's four embranchements, the mollusks. Working with the cuttlefish, a cephalopod, as a representative mollusk, they attempted to show that if a vertebrate was bent backwards so that the nape of the neck was attached to the buttocks, then the internal organs would be arranged in a manner similar to that of mollusks. The original paper of Meyranx and Laurencet is no longer extant, but there is no doubt that Geoffroy greatly distorted its contents in his report of 15 February. Geoffroy began by establishing the credentials of the

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authors: they had prepared some 3000 illustrations of mollusks and other invertebrates and had already written several brief descriptive essays. However, it was not their factual knowledge that aroused Geoffroy's excitement but their theoretical conclusions.7 Geoffroy was quick to leap upon the memoir as a support for his own doctrine of unity of composition, claiming far more for the theoretical suggestions of the two naturalists than they would have claimed for themselves. He applauded Meyranx and Laurencet for preferring the principle of connections to considerations of form, "poor counselors for philosophical comparisons." Through application of the principle of connections, Geoffroy reported, the authors had discovered in mollusks the homologue of the vertebrate diaphragm, compared the cartilaginous ring in the neck of mollusks to the vertebrate hyoid, and found elements of a vertebrate pelvis in the cartilaginous stylets which bordered on the base of the mollusk siphon. Geoffroy did not insist on these particular homologies; he was willing to admit they might be premature. Yet he assured his audience that he had no doubt that mollusks would eventually be reduced to unity of composition with the vertebrates. How could mollusks share so many organs in common with vertebrates, he asked, and yet be built on a totally different plan? Employing an argument not unlike that of Cuvier's correlation of parts, Geoffroy contended that if the organs of a mollusk were in harmony with each other, that must be because they retained the same arrangement of parts as in the vertebrates. "Just like all animal structure," Geoffroy concluded, "they cannot escape the consequences of the universal law of nature, the unity of organic composition." If that were not enough to arouse Cuvier, Geoffroy followed this statement by an even more inflammatory section, which he agreed, upon Cuvier's angry objection, to omit from the official proces-verbal of the meeting. In this retracted epilogue, which he later published on his own, Geoffroy pointed to the immense progress zoology had made since the days when naturalists concentrated only on the differences that separated the cephalopods from other animals. As an example of allegedly old-fashioned views on mollusks, he quoted from a work "rich in facts, forceful and remarkable for its knowledge and wisdom," without identifying either the title or the author's name. The source of the quotation was well known to all. Cuvier had written in "Memoir on the Cephalopods and on Their Anatomy" in 1817: In a word, we see here, regardless of what Bonnet and his followers have said, Nature passing from one plan to another, forming a whole, leaving between her productions a manifest hiatus. The cephalopods are not a transition from anything: they have not resulted from the development of other animals, and their own development has produced nothing superior to them.8

Geoffroy asserted that while it was proper in the past for zoologists to concentrate on the differences of animals, the object of zoology today had become "the knowledge of the philosophical resemblances of beings." The report ended with the recommendation that the paper of Meyranx and Laurencet be published in the Academie's journal for nonmembers. It never saw print. This time Geoffroy had gone too far, as, despite his former reluctance to

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engage in direct public controversy, Cuvier rose to his feet. Taking the floor, he vehemently objected to the section of the report that concerned him, and declared that the authors of the memoir were "completely mistaken" in their pretended homology. He promised to enlarge upon his criticism in a future memoir.9 Having agreed to retract the offensive passages from the official record of the Academie, Geoffroy believed the matter was closed. It was not so, for at the next meeting of the Academie, Cuvier came ready to fight. Meyranx and Laurencet had litle desire to become embroiled in the controversy sparked by their memoir. "I cannot find words to express how devastated I am that our memoir has given rise to disputes," Meyranx wrote to Cuvier. "We could scarcely believe that anyone could draw such exaggerated consequences from a single, simple consideration on the organization of mollusks," he protested. There was nothing in the memoir "which contradicts the admirable work that you have written and that we regard as the best guide in this matter."10 Nor did Latreille wish to become involved. He was certainly compromised in the controversy, because he had signed the report and because Geoffroy had made mention there of Latreille's ill-fated attempt to compare cephalopods and fishes in 1823. He wrote to Cuvier on 18 May 1830 to disclaim all participation in writing the report, which he claimed he did not see until just before the meeting began. He denied also any part in Geoffroy's decision to publish the suppressed portion of the report. "I had no part in this publicity," he explained, which "is very opposed to the desire I expressed to him [Geoffroy] to keep in this matter the deepest silence." While he conceded that he formerly attempted to establish affinities between vertebrates and mollusks, he had always admitted a hiatus and denied that lower animals possessed in any sense a vertebral column. He assured Cuvier that "I have since recognized the futility of these researches or of these vain efforts of the imagination."11 Latreille had every reason to be deferential to Cuvier at this time, for Cuvier had just supported his bid to become at last a titulary professor at the Museum. When the aged Lamarck died on 29 December 1829, a struggle was already underway among Latreille, Audouin, Blainville, and their advocates to obtain the chair of invertebrate zoology at the Museum. Latreille, who had for many years taught Lamarck's course, had been forced by age and illness to turn the teaching over to the young and ambitious Audouin. Blainville, considered an expert on the invertebrates, had long been awaiting this moment to become professor at the Museum. Nevertheless, Cuvier saw to it that the sixty-sevenyear-old Latreille was not passed over. After a series of complex machinations, the minister of public instruction divided the chair in two. On the very day of Geoffroy's report, the Academie chose Latreille as its candidate for the first vacancy at the Museum, and on 1 March Blainville was chosen for the second. No doubt the unpleasant prospect of Blainville's installation at the Museum reinforced Cuvier's decision to defend his scientific doctrines publicly. As the final choice of candidates by the minister had not yet been made, Latreille was in a delicate position, and he quickly acted to disassociate himself from Geoffroy.12 At the next meeting of the Academie on 22 February, Cuvier came prepared

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to do battle, with a memoir entitled "Considerations on the Mollusks and in Particular on the Cephalopods." With him he brought two large and impressive multicolored diagrams drawn by Laurillard from sketches he had provided. One showed an octopus (a cephalopod) and the other a duck-shaped mammal bent over backwards. All the organs were carefully labeled. Cuvier would show that although mollusks and vertebrates had many organs in common—namely a brain, eyes, ears, salivary glands, a stomach, a liver, a double circulation, and sexual organs—there was no reason to conclude they were built on the same plan.13 Displaying his customary clarity and precision, Cuvier first assaulted the doctrine of unity of composition by an analysis of Geoffroy's terminology. To do science, he began his argument, we must first define terms in clear and precise language. Science could not employ metaphors or rhetorical figures of speech. These criteria were then applied to the expressions "unity of composition" and "unity of plan," with the expected results. Cuvier's easy refutation of unity of composition depended on his carefully circumscribed definition of the concept. Composition, according to Cuvier, signified the arrangement among the parts—that which Geoffroy called the connections between the parts. If unity of plan and arrangement were taken in their natural sense to mean identity—that all animals were composed of the same organs arranged in the same manner—we would be led to absurdities. No one would argue that the man and the polyp had one composition and one plan. Obviously, then, unity must signify analogy. In that case, insisted Cuvier, "once divested of this mysterious cloud by which their vague meaning or distorted usage envelops them," unity of composition and plan were no longer a new banner for zoology, but principles as old as Aristotle. New homologies were discovered every day, but between one more homology and the assertion that the composition of all animals was one, Cuvier concluded, "the distance is as great as between the man and the monad." The study of "analogies" was valid only insofar as it was subordinated to a much higher principle, that of conditions of existence. "This is the true philosophical principle from which the possibilities of certain resemblances and the impossibility of certain others derive."14 Having disposed of Geoffroy's terminology, Cuvier went on to submit Meyranx and Laurencet's memoir to a detailed refutation. With the aid of his diagrams, he presented case after case of organs that were disposed differently in mollusks and vertebrates, of mollusk organs not found in vertebrates, and of vertebrate organs lacking in mollusks. For example, if a vertebrate were bent backwards, the brain would be found facing the center. For the homology to hold, Cuvier reasoned, the brain of the mollusk ought to be found on the side away from the siphon. In fact, however, it was found on the opposite side.15 Meyranx and Laurencet's argument quickly fell apart as Cuvier displayed to his audience his superior knowledge of cephalopods. Geoffroy, by contrast, knew very little about the anatomy of these animals, as Cuvier was quick to make evident. Long desiring to lure Cuvier into the arena, Geoffroy was delighted at the prospect of a public debate. He improvised a brief reply to Cuvier's memoir

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and promised to respond at length at the next meeting. While professing sadness that he and his old friend had fallen out, he welcomed the opportunity of justifying his philosophical mission. As the number of outsiders at the Academie meetings had increased, Geoffroy pointed out, the members had tended toward more reserve, and differences of opinion were rarely aired. He was pleased, he said, to return to the more lively discussions of the old days.16 Meyranx and Laurencet were soon forgotten as the debate turned into an open confrontation between the two senior academicians. On the following Monday, 1 March 1830, Geoffroy read his first rebuttal, "On the Theory of Analogues, to Establish Its Novelty as a Doctrine, and Its Practical Utility as an Instrument." From the beginning, Geoffroy wanted to enlarge the debate beyond the anatomy of mollusks. He stressed the high philosophical import of the contest: "The points to resolve are vital questions in philosophy. . . and I could only be sensitive to their influence on the moral perfection of society."17 Geoffroy's response to Cuvier's demand that he define terms clearly was at best feeble. Geoffroy had never intended unity of composition to be rigorously defined, as in fact it could not be. The resemblances Geoffroy was seeking were "philosophical resemblances" rather than obvious similarities. He claimed that he had never made a distinction as Cuvier had done between "unity of composition" and "unity of plan." Both, he contended, were shorthand expressions for "unity of system in the composition and arrangement of organic parts."18 Retreating to the less provocative label "theory of analogues" (which emphasized the method, rather than the results), Geoffroy traced the genesis of his doctrine in order to vindicate its novelty. Here Geoffroy was on more solid ground, for his concept of homology was indeed different from anything found in Aristotle. Knowing that some considered him a dreamer, Geoffroy made a point of insisting on the rigorous scientific nature of his "method." Aristotle's "analogies," Geoffroy argued, were merely instinctive comparisons because they were based on fugitive guides, namely form (shape) and function. His own "analogies," based firmly on the principle of connections, compared the individual structural elements instead of entire organs at once. His theory, he claimed, offered a scientific means of making new discoveries while Aristotle's did not.19 To illustrate the explanatory power of the theory of analogues, Geoffroy turned to the example of the hyoid bone. In man he found the bone was composed of five small pieces, while in the cat it was composed of nine. (Geoffroy noted in his third paper of the debate that he had been the first to name the pieces.) The structures had long been given the same name—hyoid—but were they homologous? Aristotelian doctrine responded in the affirmative and did not probe further, because the two organs had the same function (as supports to the voice-box). According to the theory of analogues, however, the homology was incomplete because one hyoid had more pieces than the other. To complete the comparison, Geoffroy argued, it was necessary to determine which pieces in the cat corresponded to the five pieces in man and what happened to the four missing bones. Geoffroy ascribed the reduced number of bones in man to his vertical station and sought rudiments of the missing pieces

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in ligaments which joined each side of the hyoid to the styloid process of the temporal bone in the skull. Verifying his guess by embryology, Geoffroy was able to observe in the human fetus that the styloid process was not originally attached to the skull and that it was composed of two primitive bones. It was through reasoning based on the theory of analogues that this success was possible.20 The point of the controversy between himself and Cuvier, as Geoffroy saw it, was whether one ought to restrict or enlarge the field of such "philosophical considerations."21 For the next two weeks there was a pause in the debate. On 8 March Geoffroy wrote a note to the Academie to say that he was indisposed and unable to present his memoir. Cuvier, for his part, informed the Academie that he would read his memoir only in the presence of Geoffroy. On 15 March the large crowd gathered at the Academie was disappointed because Cuvier was unable to attend.22 The controversy was resumed on 22 March with Cuvier's memoir, "Considerations on the Hyoid Bone,"23 followed by Geoffroy's memoir, "Application of the Theory of Analogues to the Organization of Fishes."24 Cuvier went over much of the same ground with even greater forcefulness. Again he demanded a definition of unity of composition that would render the theory verifiable: But how can one discuss a question when one cannot pose the terms of it? In this matter I have made clear and positive requests. You insist on the elements [i.e., the individual bones rather than entire organs]. Well then, do you mean that there are always the same elements, do you mean that these elements are always in the same mutual disposition"? Finally, what do you mean by your universal analogies? If our colleague had made a clear and precise response to my requests, that would be a fine point of departure for our discussion. But in his lengthy deduction he has not responded, because it is no response to say that all animals are the product of the same system of composition. It is to say the same thing in other terms, and in much more vague, much more obscure terms.25

He declared the question at issue was to determine whether the so-called theory of analogues was universal or if there existed only limited "analogies" of all sorts. Confronting Geoffroy on his own ground, Cuvier took up the proffered example of the hyoid bone. He proposed to examine systematically the hyoid in mammals, birds, and reptiles to show that it changed in number of parts even from genus to genus, that it changed in arrangement of the pieces, and furthermore, that many animals had no hyoid at all. No matter how one defined the vague terms "analogy" and "unity of composition" and "unity of plan," Cuvier promised to demonstrate that they could not be applied in a general manner. A particular example, the hyoid of the howler monkey, may serve to illustrate the points at issue. Geoffroy had argued that the enormous hyoid of this animal, which resulted in its peculiar howling cry, was simply a modification of the hyoid of other mammals. Cuvier, on the contrary, insisted that the hyoid of the howler was essentially different from that of other mammals. In the

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hyoid of the howler, he found no trace of the anterior horns, the stylohyoid ligament, or the styloid process. This absence he explained in teleological terms: since the function of the hyoid of the howler was substantially different from that of the hyoid of other mammals, different numbers of parts and attachments were called for. In order to produce the howling sound, the hyoid took the form of a huge drum rigidly attached to the lower jaw. There was no need of an attachment to the cranium, as in other mammals.26 In his commentary on Cuvier's paper (in the footnotes of Principes de philosophie zoologique), Geoffroy denied any new attachment but did not contest the change in the number of elements. He offered a materialistic explanation for the same facts. Appealing to the law of balance of organs, he argued that the hypertrophied hyoid body developed at the expense of the bones making part of the styloidian chains, and therefore only a thin ligament remained. "Thus," Geoffroy concluded, "that which the theory of analogues does not encounter in number of parts . . . it finds in justifications, in compensations which it can discern, in rudiments which give the how and the why of the disappearance of certain materials."27 Cuvier particularly objected in his memoir (as he had in the past) to those of Geoffroy's homologies which presupposed complicated transformations or metamorphoses of parts. In birds, Cuvier and Geoffroy both agreed, the hyoid was no longer suspended as it was in mammals from the temporal bone, the posterior horn was lacking, and the tongue was osseous. To Cuvier these facts required no explanation beyond noting that new needs had called for new structures to accommodate those needs. Seeking a nonteleological explanation for the apparent change, Geoffroy had in an earlier memoir contended that the hyoid bone rotated from its position in mammals and that the posterior horns had been transformed into the osseous structure of the tongue. Such a supposition Cuvier claimed to find incomprehensible: Without a doubt, this is a somersault possible to conceive in a skeleton whose bones are held together only by brass wires, and where there are only bones [to consider]. But I ask of anyone who has the slightest idea of anatomy: is it admissible when one considers all the muscles, all the bones, all the nerves, and all the vessels which are attached to the hyoid bone! It would necessitate—But I stop myself] The very idea would frighten the imagination!28

All that Cuvier was willing to grant with respect to the hyoid bones of the higher vertebrates was a certain resemblance of structure, with the degree of difference depending on the different uses to which the organ was put. In a final warning to his audience, Cuvier made a brief reference to the religious issue. Repeating the argument of his 1825 article "Nature," he insinuated that "unity of composition" would place unnecessary restrictions on the Creator and was moreover detrimental to the progress of science: But, if one neglects all these considerations in order to see only these pretended identities, pretended analogies, which, if they had the least reality, would reduce Nature to a sort of slavery, into which, fortunately, her Author is far from having enchained her, then no one will know anything about beings, either in themselves or in their relations. The world itself would become an indecipherable enigma.29

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Geoffroy was not able to respond immediately to this charge, for the Academie allowed him only to read the memoir he had already prepared on the theory of analogues applied to fishes. Geoffroy first attempted to justify his shift of the focus of the debate from mollusks to vertebrates, to hyoid bones, and now to the fishes. Cuvier had complained earlier that Geoffroy had abandoned the subject under discussion, namely the mollusks. Geoffroy now excused himself by arguing that the study of homologies between mollusks and vertebrates was not yet sufficiently well advanced to allow for fruitful discussion. He accounted for the lack of many vertebrate organs in mollusks by reference to the theory of arrests of development. Mollusks had been permanently arrested at a lower degree of organic development before many of the organs of vertebrates made their first appearance. Although he believed that affinities between mollusks and vertebrates would eventually be discovered, Geoffroy claimed that for now he preferred to defend his doctrine with respect to the fishes, animals which occupied an intermediate rank between higher vertebrates and invertebrates, and whose structure it was necessary to understand before proceeding to mollusks.30 Cuvier, in the course of the debate, made little response to this appeal to the recapitulation theory, except to remark in passing in his second paper that the principle of "degradation, an insensible simplification of beings," was absolutely contradictory to "identity of composition," yet that the doctrines were often allied, such were the "bizarreries" in some minds.31 Refusing to take the recapitulation theory or embryology into serious consideration as explanatory tools in philosophical anatomy, Cuvier continued his argument as if unity of composition would stand or fall depending on whether one could find the same parts in the same arrangement in adult animals. Geoffroy's presentation on fishes contained much justification and no new science. He traced the entire history of his research on the homologies of the bones related to respiration in fishes and those in mammals, reptiles, and birds. The skeleton of fishes, he once more asserted, was the same in essence as the skeleton of mammals, but modified to accommodate itself to the aquatic environment. In conclusion, he again hailed the advent of a new scientific era. Should we continue to pursue the beaten paths of anatomy, he asked, or should we not attempt to open new paths under the guidance of recent discoveries?32 Both rivals were now prepared for a drawn-out battle. That neither was fully in control of the situation is shown by the petty way in which they behaved. The following meeting, on 29 March, began with an argument over who should be permitted to speak first. Geoffroy claimed the right to respond to Cuvier's memoir on the hyoid, since he had not had his say at the previous meeting. Cuvier insisted that since Geoffroy had presented the last memoir, it was now his turn. The dispute scarcely redounded to the credit of either participant. The result was that Geoffroy first presented his memoir, "On Hyoid Bones," and Cuvier followed with improvised comments.33 Geoffroy stressed in his memoir, devoted entirely to self-justification, that the disagreement was not over facts but over the "scientific appreciation" of facts: "It is a question of philosophy that divides us."34 He had little quarrel

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with Cuvier's long-winded recitation of facts concerning the hyoid in mammals and birds. He himself had shown that the number of parts of the hyoid varied (however, he denied that the connections were altered). Nor was there anything surprising in the fact that lower animals lacked a hyoid, for a hyoid was incompatible with their degree of development. Considering the rapid progress that had been made in the science of organization since he had first introduced the theory of analogues, by German Naturphilosophie, by the studies of monstrosities, and by the work of Serres on animal development, he felt he ought to be excused some errors in his early work on the hyoid.35 In his opinion, Cuvier had failed to come to grips with the real issue. Cuvier had not understood the importance of seeking resemblances that were masked by apparent dissimilarities. The value of the theory of analogues was that it offered a scientific explanation for differences of structure. In this, his final contribution to the debate, Geoffroy widened the scope of the controversy to include a number of the broader issues that had surfaced in the 1820s: final causes, facts versus theories, evolution, and Naturphilosophie. Geoffroy contrasted the old method of comparison, that of Cuvier and Aristotle, which was forced to resort to final causes, to his own method, which eschewed final causes in favor of the belief that "such is the organ, such will be its function."36 Once more he brought up the curious argument of his 1828 memoir on evolution, repeated time and again in his philosophical writings of the 1830s, that the great geniuses of the past had often intuited a general truth while the facts that they brought forward in its defense were found to be false. As an example, he cited Lamarck's contention "that there are in the environment [le monde exterieur] causes of sufficient influence and activity to modify by their action the organization of animals." One should not be deterred from employing one's faculty of judgment by the often and deliberately repeated proposition "that natural history is the science of particular facts " he warned. While the German Naturphilosophen might have abused the a priori method and allowed their imagination to lead them to the point of poetry, the school of Cuvier, in Geoffroy's estimation, had gone to the opposite extreme in demanding that science be limited to registering positive facts. He portrayed his own work as part of a European movement which was threatening to leave France behind.37 In his last contribution to the debate, read on 5 April, Cuvier continued his enumeration of counterexamples of unity of composition in the hyoids of vertebrates, but he too enlarged the bounds of the dispute by injecting several of the themes from his article "Nature."38 In reviewing the hyoids of vertebrates living in water, he particularly focused on Geoffroy's hypothesis that the hyoid of fishes was composed of a mixture of pieces, some of which were found in the hyoid of higher animals, and others of which were found in the sternum. To combat this unusual determination of the hyoid in fishes, which he declared to be "one of the most singular hypotheses that have ever been proposed in anatomy," Cuvier first examined the sternum in mammals, birds, and reptiles. He found uniformity neither in the number (there were anywhere from one to nine pieces) nor in the connections of the pieces. Sometimes he found both

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sternum and ribs, sometimes ribs without a sternum (in serpents), and sometimes a sternum without ribs (in frogs). Cuvier paused in the argument to reaffirm his belief in the sufficiency of teleological explanations. "Why, indeed, would nature have acted otherwise?" he inquired. "What necessity would have constrained her to employ only the same pieces and employ them always?" Continuing in his investigation, Cuvier demonstrated to his satisfaction that even with all the translations, rotations, and interpolations of parts required by Geoffroy's hypothesis, it fell hopelessly apart. If fishes had a larger hyoid, this was because because their branchial respiration required it. Nature was not obliged to borrow fragments from the sternum to develop the hyoid in fishes, for "her creative powers suffice to give to fishes, as to her other productions, all that was necessary to each one of them."39 Here, as in "Nature," Cuvier linked together Geoffroy's theory of analogues with other hypotheses which he opposed, namely Naturphilosophie and evolution. Resorting to his usual strategy, he merely insinuated the connection rather than demonstrating it in any meaningful way: I know very well that, for certain minds, there is behind this theory of analogues, at least confusedly, another theory, very ancient and long ago refuted, but which some Germans have revived for the benefit of the pantheistic system called nature-philosophy, that of the production of all species by the successive development of originally identical germs.40

Although the major issue at controversy in the debate was the validity of the theory of analogues or philosophical anatomy, both Cuvier and Geoffroy had injected the issue of evolution. Both had related evolution at least obscurely to philosophical anatomy. Thus while evolution was not the central focus of the debate, naturalists later in the century were not entirely mistaken in their belief that the debate concerned evolution. Indeed, in the two years after the debate at the Academie ended, evolution became increasingly an issue of contention between Cuvier and Geoffroy. By the beginning of April, the debate was no doubt straining the patience of the members of the Academie and threatening to get out of hand. It was clear that Cuvier and Geoffroy were arguing at cross-purposes, and could come to no resolution of the issues. Both planned long series of memoirs that would only have reiterated the points they had already made. Geoffroy proposed to examine Cuvier's objections with repect to monstrosities, the bones of the cranium, and homologies between vertebrates and articulates, while Cuvier planned to examine "several other laws advanced by various naturalists" as well as to go through the entire animal kingdom organ by organ to prove that unity of composition was nonexistent.41 Moreover, the debate threatened to degenerate into a free-for-all. Because of the stature of the combatants and the press coverage the debate received, the seats for visitors were filled to overflowing each week. The presence of a large and noisy audience tended to make the debate more like a theatrical event than a serious scientific discussion. Geoffroy had complained on 29 March that "a meeting of the disciples of the Portico" was becoming transformed into "a pit applauding the outrageous come-

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dies of Aristophanes."42 In the preliminary discourse to Principes de philosophie zoologique, Geoffroy regretted also that the issue of religion had been brought into the debate against his will: "I wished to rely on the arguments of naturalist to naturalist; the arguments [instead] became theological."43 It was partly for these reasons that the public confrontation came to an abrupt halt at the close of the 5 April meeting of the Academie. Geoffroy took the initiative in the matter by announcing to the Academie that he would not reply to Cuvier's memoir. He would put an end to a polemic which, instead of clarifying issues, had altered his friendship with Cuvier. Instead, he proposed to publish a series of works defending his doctrines against Cuvier's objections and distributed a "prospectus" to the members of the Academie.44 For his part, Cuvier was also satisfied to end the debate. In his analysis of the work of the Academie for 1830, he commented that the discussions "had ended by becoming too specialized, too detailed, for the authors to continue to demand for themselves the time and the attention of the Academie."45 Geoffroy immediately set to work to publicize the controversy to a wider audience. Principes de philosophie zoologique was rapidly written. By 15 April 1830 the preliminary discourse was sent to the printer and in May the work was ready for sale. The work contained the report on Meyranx and Laurencet (including the suppressed portion) and the texts of five of the six papers presented by Cuvier and himself, together with introduction and commentary. Here in one small volume, in easily assimilated form, were most of the matters under dispute in the 1820s. Though supposedly printed in only a very limited edition, the book was of immense importance in focusing the attention of naturalists, especially those outside Paris, on fundamental issues for the future of biology.46 The Debate Publicized

The Cuvier-Geoffroy debate might have been long ago forgotten if it were not for Geoffroy's book and the extensive coverage given to the debate in contemporary journals and newspapers. Principes de philosophie zoologique reached a primarily scientific audience, but through the many other publications, the debate entered into the common intellectual milieu. It was through the reports in journals and newspapers that the public was alerted to the controversy and came to fill the seats at the Academie meetings. Publications followed the arguments from week to week, printed long excerpts of the papers presented to the Academie, and did not hesitate to take sides on the issues. Reviewers differed widely on the significance of the controversy, some interpreting it as a simple matter of anatomy while others pointed to philosophical issues of major importance. Those in the latter category frequently linked Geoffroy with Naturphilosophie and saw him as the leader of a progressive and synthetic school of natural history which was displacing the conservative and outmoded school of Cuvier. In general, the more conservative journals favored

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Cuvier, while the more liberal periodicals, those that especially welcomed the Revolution of 1830, were sympathetic to Geoffroy. It is significant that despite the references to evolution in the course of the debate, none of the reviewers were under the impression that the debate concerned transmutation of species. Regardless of the larger philosophical issues involved, all saw the primary scientific issues as having to do with philosophical anatomy. A broad view of the events of the spring of 1830 was taken by the most famous of all contemporary commentators on the Cuvier-Geoffroy debate, the German novelist, poet, and amateur naturalist Johann Wolfgang von Goethe. We are interested here in Goethe's commentary, not in order to understand Goethe or German science, but because the two articles written by Goethe on the debate in September 1830 and March 1832 entered forcefully into the French context of the controversy. These essays, which recounted for his countrymen the course of the debate and its significance, appeared in translation in such major French journals as the Revue encyclopedique and the Annales des sciences naturelles. Whenever Geoffroy's followers wished to underline the momentous nature of their cause, they never failed to note that the eminent Goethe had devoted the last article of his career to the debate.47 Of the journals favoring Geoffroy, one of the most widely read was the Revue encyclopedique, published in four volumes a year from 1819 to 1833, and directed to a nonspecialized educated audience. One of the earliest issues contained Flourens's laudatory review of Philosophie anatomique, and the Revue had ever since shown partiality to Geoffroy. It presented weekly accounts of the debate, including lengthy extracts of Cuvier's and Geoffrey's papers. A note from the editors indicated that since the journal generally published Geoffroy's views, it was now satisfying a sense of justice in publishing Cuvier's first memoir on the debate as a separate article. Yet Cuvier was not given too great an advantage, because Geoffroy was given the opportunity to append a reply.48 Most partial of all the journals to Geoffroy was the Gazette medicale de Paris, a medical weekly begun in 1830. It frequently published letters from Geoffroy, most of them describing malformed fetuses. In the weekly accounts of the meetings of the Academie des Sciences and the Academie de Medecine (where Geoffroy, an elected member, also aired his views), the editor, Jules Guerin, a physician with a special interest in orthopedics, often inserted his own opinions. The favoritism shown Geoffroy is illustrated by the account of the meeting of 22 February at which Cuvier read his first memoir of the debate. Cuvier was depicted as forced to defend his outworn doctrine against "new ideas" introduced by Geoffroy and "adopted by the greatest naturalists of Germany": M. Cuvier has perceived that such a sanction given to opinions which are no longer in agreement with his could well cause a complete revolution in zoology. It is in order to combat this imminent tendency that he has armed himself with all his talent.49

Guerin published Cuvier's objections to unity of composition, but he judged them to be "neither solid nor direct," and confidently awaited Geoffroy's response.

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In his review of Principes de philosophie zoologique in the 30 November issue, Guerin interpreted the debate as concerning the validity of the doctrine of unity of composition in the animal kingdom, and linked Geoffroy with the course of scientific advance. During the debate, wrote Guerin, Cuvier seemed to win his case in the eyes of the majority, but as time passed, Geoffroy would regain the advantage. Geoffroy's defense, "lacking art and obscure and unintelligible at first," has become better appreciated now that the public has come to realize that Cuvier represented the "status quo" while Geoffroy represented the "urge for progress": But this is the history of all scientific progress. Criticism prevails at first, especially when it comes from a powerful mind, but it soon gives way before the natural force of truth, because the human mind, once it comes to understand [the truth], in the end [always] ranges itself on its side.50

Guerin's joyful reception of the Revolution of 1830 may not be unrelated to his championing of Geoffroy. Both Geoffroy and the Revolution represented to him liberation and progress. Hailing the "glorious era" that was commencing for France, he expected that the "return of liberty ought to contribute as much to the advancement of the sciences as to the progress of political institutions."51 Some publications made a special effort to avoid partiality. The liberal daily paper Le Globe gave a full but evenhanded coverage of the debate. It had in the past frequently published accounts of Geoffroy's memoirs to the Academie. On the other hand, the paper gave publicity to Cuvier's conception of science through publishing the lectures of his popular course in the history of science at the College de France.52 The Annales des sciences naturelles, the major professional journal for naturalists aside from Museum publications, also followed a scrupulous policy of keeping above the fray, but gave much space to the events of February to April 1830. The journal, controlled by the proteges of Alexandre Brongniart, had always made a point of publishing articles of all shades of opinion, including many that Cuvier might likely have found disconcerting. The reports of the meetings of the Academie in the journal's supplement, the Revue bibliographique, contained excerpts from Cuvier's clear and logical memoirs and paraphrases from Geoffroy's more diffuse memoirs. Annales des sciences naturelles published not only Cuvier's first memoir of the debate, but also one of Goethe's pro-Geoffroy articles on the debate and Antoine Duges's controversial effort at a compromise.53 The periodical that most avidly supported Cuvier was the Journal des debats, whose science editor, the physician Alfred Donne, was, according to Geoffroy, a disciple of Cuvier. One of the most important daily newspapers in Paris, the Journal des debats represented Catholic, royalist opposition to the Polignac ministry. Unlike the more liberal papers, the Journal des debats obeyed the king's order not to publish during the July Revolution. Donne printed long extracts of Cuvier's contributions to the debate, but failed to give Geoffroy equal coverage. He explained to his readers that he had intended to give an analysis of the memoirs of Geoffroy, but found it difficult to do so: "We ought to avow that the theory of analogues, which is founded on general

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and somewhat abstract considerations, is much less easy to follow than the precise and positive facts of comparative anatomy developed with such art by M. Cuvier."Geoffroy's doctrine, he implied, was by comparison, a philosophical system that was impossible to demonstrate in a rigorous way, and could be accepted by Geoffroy and his followers only as an article of faith.54 Geoffroy reprinted in Principes de philosophie zoologique two flattering reviews of the debate from daily newspapers, Le National and Le Temps, both representing the liberal opposition. While Le Temps, though insisting on Geoffroy's originality, retained some measure of impartiality, the editors of Le National rallied to Geoffroy and offered an exceptionally broad interpretation of the controversy.55 The main points of the commentary were incorporated into at least two reviews of Geoffroy's book, one by Gaspard-Joseph Martin Saint-Ange for the Revue encyclopedique and the other by J.B.G.M. Bory de Saint-Vincent for the staid Moniteur.56 The controversy, according to the review in the 22 March 1830 issue of Le National, concerned nothing less than the fate of zoological philosophy as taught by Aristotle and the naturalists of twenty-two centuries. Since Aristotle, wrote the reviewer, most naturalists believed that each animal was provided with special organs conforming to the final cause of the organism. However, for the past thirty years other principles had been introduced into Germany by Kielmeyer, Oken, Spix, Tiedemann, and J.F. Meckel, and into France by Geoffroy. Would the old zoological philosophy, championed by Cuvier, but now demonstrated "insufficient and incomplete," yield to the new and progressive ideas, of which Geoffroy's laws—the theory of analogues and the principle of arrests of development—were representative? The reviewer was concerned to underline the great relevance of the controversy to the nonspecialist. When scientific questions touched on the highest generalities, the reviewer wrote, the interest of the public must necessarily be aroused. The questions raised by the dispute are "of a nature to seize the imagination of every man who thinks, and to forcefully occupy all minds for whom the spectacle of animate nature is a fertile source of poetic, philosophical, or religious emotions." Geoffroy's doctrine, the review implied, was entitled to public favor because it would remove final causes and the arbitrary will of the Creator from science and would be more conformable to a subjective, aesthetic view of nature than Cuvier's.57 The public stake in the philosophical issues raised by the debate was further elaborated upon by Goethe in his two essays on the debate.58 In the singular conversation with Frederic Soret on 2 August 1830 which was quoted in the introduction to this book, Goethe was more astonished at the news of the debate than at the fall of Charles X. "The volcano has come to an eruption; everything is in flames," he exclaimed to Soret.59 What, we may ask, led Goethe to become passionately involved on Geoffroy's behalf in a French scientific controversy? One obvious reason for Goethe's interest in Geoffroy and the debate was similarity of views (even though, as has been pointed out earlier, the foundations of Goethe's and Geoffroy's views were different). Like Geoffroy, Goethe

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placed primary emphasis on animal and plant structure (though he did not reject teleology). In fact it was Goethe who first popularized the term "morphology." Well before Geoffroy embraced philosophical anatomy, Goethe, convinced that all vertebrates were built on the same underlying plan, which he called the "archetype," was led to search for the intermaxillary bones (premaxillae) in man, thus refuting Peter Camper's suggestion that man differed from the apes by the absence of these bones. Although the bones were invisible in the adult, Goethe demonstrated that they existed as separate bones in the fetus and as partially detached bones in the infant. To explain modifications of the intermaxillary bones in different animals, Goethe advanced a principle of compensation similar to that of Geoffroy's law of balance of organs. His discovery, made in 1784 and recalled in his second article on the debate, was reported at the time to Camper, Blumenbach, and others, but was not published until 1820. In other letters to friends, predating Oken's publication of 1807, Goethe sketched out the vertebral theory of the skull.60 Goethe's most celebrated application of the archetype concept, however, was to plants. In Metamorphosis of Plants (1790), he traced in poetical language the development of an annual plant from the seed of one generation to the seed of the next, reducing all plant structures to a single ideal type, the leaf. The cotyledons, which Goethe regarded as imperfect leaves, were successively metamorphosed to form the calyx, corolla, stamens, and pistils under the generative influence of the ever more refined sap.61 Despite the superficial resemblance to Geoffroy's morphological mode of thought, Goethe's biological writings were most likely unknown to Geoffroy before the 1820s. But once he learned of Goethe's work and interest in the debate, Geoffroy was happy to become his champion. Having arranged for the Revue encyclopedique to publish Goethe's first article, Geoffroy appended an article of his own presenting Goethe's credentials as a naturalist and insisting on his right to judge the debate.62 Similarity of doctrine is, however, insufficient to account for Goethe's intense excitement concerning the debate. For Goethe the debate raised a number of much larger philosophical and social issues, among them the reception of Naturphilosophie in France, the relative roles of analysis and synthesis in science, and the control of scientific ideas by scientific elites. Goethe's two essays served as vindication for himself and his countrymen against assaults from the French, and in particular from Cuvier. His own work finally receiving recognition in the 1820s, Goethe took great pride in the achievements of his fellow Naturphilosophen. Like many of his contemporaries, he believed the so-called synthetic science of Geoffroy to be derivative from Germany rather than linked to the older philosophe science in France. He was therefore particularly incensed by Cuvier's innuendos against the Germans in his memoir of 5 April, and endeavored to defend German science from Cuvier's veiled charge of irreligion. Goethe saw the debate as primarily a conflict between the analytic view of nature represented by Cuvier and the synthetic view of nature introduced from his homeland into France by Geoffroy. "What is all intercourse with nature," he asked Soret, "if by the analytic method, we merely occupy ourselves with

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individual material parts, and do not feel the breath of the spirit, which prescribes to every part its direction, and orders, or sanctions, every deviation, by means of an inherent law!"63 In his second article, he drew a parallel between Daubenton and Buffon on the one hand and Cuvier and Geoffroy on the other. While not quite so dry and uninspired as Daubenton, Cuvier, in Goethe's estimation, nevertheless shared Daubenton's preoccupation with the details of natural history. Geoffroy, like Buffon, "seeks to penetrate the cause of the universality of things." Cuvier and Geoffroy, he concluded, represented two poles in perpetual conflict. Although he believed that both the analytic and synthetic approaches were necessary to ensure the progress of science, it was clear that his sympathies lay with Buffon and Geoffroy.64 Finally, Goethe was aroused by the debate and its immediate aftermath because he hoped that it signaled that science had at last been liberated from the shackles of those who conspired to stifle novel ideas. "This sanctuary of the sciences," Goethe wrote of the Academie des Sciences in his first article on the debate, is a place "where one is perhaps still more concerned with covering obstacles by a veil and eluding them rather than dealing with them openly."65 His conversation with Soret made allusion to a momentous meeting of the Academie that had taken place on 19 July 1830. The outcome of the battle that arose at this meeting, Goethe excitedly informed Soret, was that "the synthetic manner of treating nature, introduced by Geoffroy into France, cannot be held back any longer": The affair has now become public, through the free discussion of the academy, and that in the presence of so large an audience. It is no longer referred to secret committees, and arranged and got rid of, and smothered behind closed doors. From the present time, mind will rule over matter in the physical investigations of the French. There will be glances of the great maxims of creation, of the mysterious workshop of God!66

What had happened at the 19 July meeting to give Goethe hope that the Academie had changed course? Goethe's optimism was occasioned by a heated clash between the two permanent secretaries of the Academie, Cuvier and Arago, over access to the proceedings of the meetings. Goethe explained in his second article on the debate, "I have been informed that the debates of 1830 have modified the customs of the Academie and that was especially manifested in the meeting of 19 July." At the meeting of 12 July, Cuvier had presented a short paper on the extinct bird of Mauritius, the dodo, which was followed by an argument among Cuvier, Blainville, and Geoffroy as to the proper classification of the known remains. On 19 July, when Arago read the proces-verbal of the previous meeting, instead of just noting the title of Cuvier's paper as was customary, he read an extract from it. Cuvier interrupted Arago's reading with a vehement protest. Accusing Arago, who had just been elected secretary on 7 June, of abandoning the customs of permanent secretaries for the past century and a half, Cuvier contended that to give a detailed account of the memoirs would lead to great inconvenience. The secretaries would not be capable of giving an accurate, brief analysis of the memoirs not in their immediate area of expertise, and if,

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instead, the authors themselves wrote the extracts, they would be too long. Taking the opposing viewpoint, Geoffroy congratulated Arago on his happy innovation, which, he claimed, was moreover conformable to what was practiced by all other learned societies, including the Societe Philomatique and the Academie de Medecine. Dumeril supported Cuvier, arguing that if Arago had his way, a major proportion of every Academie meeting would be devoted to the reading of the proces-verbal. Blainville sided with Geoffroy and Arago, and suggested the matter be taken up in a secret committee. This was agreed to. When Arago finally was allowed to finish reading the analysis of Cuvier's memoir, Cuvier complained once more that Arago had not supplied all of his reasoning, and warned that other authors would no doubt lodge similar objections and thus take up the valuable time of the Academie with unprofitable arguments.67 The incident seemed to suggest that the former secretive and elitist policies of the Academie which were supported by Cuvier would now give way to more liberal policies, and that in the future there would be more open discussion of differences of opinion. Goethe declared to Soret that in this affair, they had "a powerful and permanent ally" in Geoffroy.68 Goethe's interpretation of the debate as ideas versus facts, or analysis versus synthesis, was echoed in the 1840s by Isidore Geoffroy. The philosophical and political aspects of the controversy, brought to the attention of the public by Goethe in the immediate aftermath of the debate, were for the most part ignored by later scientific commentators. But they were expanded upon by a number of nonacademicians of the 1830s and 1840s. They, like Goethe, saw in Geoffroy a champion of freedom of expression in science, and they gave weight to their demands for the reform of science by appealing to the authority of the illustrious naturalist/poet. Duges Attempts a Compromise

For naturalists the major issues were anatomical. Was there unity of composition in the animal kingdom? To what extent could one establish links between embranchements? How far could philosophical anatomy be extended? Cuvier and Geoffroy offered extreme and opposite answers to these questions. The debate thus offered a challenge to those who wished to find some middle ground. Although most of the leading naturalists eventually did arrive at a compromise position, it was only later that they publicized their views. Knowing that it was unprofitable to become involved in the controversy, most of the core group of naturalists prudently avoided taking stands on these issues at this time. It is not surprising that there was little immediate reaction to the debate among members of the Academie. With less experience in the pitfalls of the politics of Parisian science, Antoine Duges of Montpellier, stirred by the memoirs of the debate, brashly presented to the Academie in 1831 an attempt to reconcile the two adversaries. His effort is indicative of the continuing attraction of philosophical anatomy, and also of the real confusion, highlighted by the debate, concerning the status of homo-

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logies. His failure suggests that the years 1830-1832 were not the time for an open discussion of the issues. Born in Mezieres in the north of France in 1797, Duges had studied medicine in Paris, and was in 1831 professor of external pathology and operative medicine at the Faculte de Medecine of Montpellier. Since 1826 he had been writing papers on zoology and comparative anatomy, several of which were submitted to the Academie and reported upon by commissions. His "Memoir on Organic Conformity in the Animal Scale," read at the Academie on 17 October 1831, nearly succeeded in bringing about a resumption of hostilities.69 Although he thought of himself as offering a compromise, Duges was in fact solidly in the camp of philosophical anatomy, and can be considered one of Geoffroy's most important disciples in France. With the help of his friend and fellow Montpellier-trained naturalist, Alfred Moquin-Tandon, he developed the fruitful idea of conceiving of invertebrates as colonies of simpler "organisms," each of which contained actually or ideally all the organs necessary for individual life. This concept was expanded upon by invertebrate zoologists later in the century, especially by the evolutionist and Museum professor Edmond Perrier. In place of "unity of organic composition," Duges proposed to substitute a new and more flexible formula, "organic conformity in the animal scale." In his opinion, Geoffroy in the recent debate had insisted in too absolute a manner on unity of plan in the animal kingdom. Convinced by Cuvier's arguments that it was not possible to compare all details of organic structure, Duges still believed that "the mutual disposition of principal parts" could be traced from class to class. To establish "organic conformity," Duges proposed a revision of Geoffroy's laws of animal organization. In particular, he argued that the "law of complication of organisms"—the gradual increase in complexity as one ascended the scale—must be supplemented by a more important law, that of the "repetition or multiplicity of organisms."70 Duges's concept of "organism" was a generalization of the concept of "zoonite" advanced by the philosophical botanist and anatomist Moquin-Tandon. In his thesis on the family of Hirudinidae (leeches and related annelids) presented to the Faculte des Sciences of Montpellier in 1826, Moquin-Tandon showed that the leech could be divided into a series of similar segments, each corresponding to five external annulations (anneaux) and each provided with its own stomach, nervous ganglion, swelling of the vascular system, lungs, and seminal vesicles. Each segment thus had a complete system of organs and, as experiments demonstrated, was capable, more or less, of living on its own. These "elementary individuals" he called "zoonites." In some invertebrates, he suggested, zoonites could live independently, while in others they lost their liberty through modification and more intimate association with other zoonites.71 Duges defined "organism" in a specialized sense as an animal or portion of an animal which was actually or ideally capable of an independent life, and expanded the concept to include all invertebrates and even the vertebrates (which Moquin-Tandon had regarded as unitary animals). As one ascended the

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animal scale, Duges explained, the "organisms" fused together ("law of coalescence") and became modified in order to centralize functions ("law of modification and complication"). The lives of the individual "organisms" thus increasingly became subordinated to the life of the animal as a whole. In the case of vertebrates, the coalescence was so complete that it was difficult to discern the separate "organisms," yet their presence was indicated by the segmentation of the vertebral column. The "laws of disposition" regulated the various modes of association of "organisms": they could be joined in a circle as in radiates, in a cluster as in animals which Duges called racemes (racemiaires), or biserially as in articulates and vertebrates. Based on these laws of organization, Duges offered a new classification of the animal kingdom, in which seven primary divisions took the place of Cuvier's four embranchements, and proposed to show how one might pass by gradations from one group to the other. He diagrammed two parallel paths by which one might establish a transition from the unitary animals (monodaires) to the biserials: one passing through the radiates and mollusks and the other through the racemes and helminths (worms). Tackling the more difficult problem of comparing vertebrates and articulates, Duges rejected from the outset Geoffroy's homologies between the exoskeleton of insects and the vertebrae of vertebrates, since he believed that invertebrates possessed at most a rudimentary skeletal system. On the other hand, he readily adopted Geoffroy's notion, which he supported by embryological evidence, that an articulate crawled with its back facing the earth and its ventral surface facing the sun. On the assumption that every "organism" of biserial animals was theoretically provided with a pair of appendages, Duges developed an elaborate system of homologies between mouthparts and legs in articulates and vertebrates. His most outrageous hypothesis, no doubt, was his comparison of the five thoracic legs of decapod Crustacea with the arm of man, which he regarded as resulting from the fusion of five primordial appendages corresponding to the five cervical nerves. The fusion of these five original appendages, he claimed, was more pronounced as one ascended the arm from the five free fingers and five encased metacarpals to the two bones of the forearm and the single humerus.72 Idealistic theories such as these were hardly likely to please Cuvier when he was called upon to judge them. Although Geoffroy claimed to disagree with Duges on minor points, he was in general sympathy with a work which "enters . . . upon the highest questions of natural philosophy." He and Blainville had been named by the Academie to present the report on Duges's memoir, but soon afterward, Geoffroy decided to excuse himself on the grounds that he was both "judge and party" and because he feared "a new collision" at the Academie. Instead, he published some remarks on the memoir in a letter to the Gazette medicale. He felt that Duges had misunderstood "unity of organic composition," which he asserted to be a more general doctrine than that of Duges. Despite his criticisms of the memoir, Geoffroy believed that it deserved the utmost attention of the Academie. He recommended that a commission of as many as five members be selected to deal with it.73

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Cuvier had a very different perspective on the memoir, as the naive young naturalist was chastened to learn. When Geoffroy resigned from the two-member commission, Magendie was named in his place, but according to Duges, arguments over setting up of a commission did not cease until Cuvier himself was named to write the report.74 Cuvier's review of the memoir was drafted, but he died before he could formally submit it to the Academie. In the report, which is extant in manuscript, Cuvier applied his usual common-sense analytic approach to divest Duges's doctrine of all its transcendental aspects. From his analysis of Duges's definition of "organism," Cuvier concluded that once Duges admitted that "organisms" could coalesce to the point of losing their ability to carry on an independent life, the concept could mean no more than resemblance of parts. A resemblance among segments in certain invertebrates Cuvier readily granted, as long as it was recognized that the resemblance was never complete except in truly composite animals such as corals. Mollusks, Cuvier noted, did not fit Duges's "system" at all, since they were extraordinarily complicated animals with no trace of segmentation. Still deeply engaged in battle with Geoffroy, Cuvier took pleasure in pointing out that Duges's system had "nothing in common with that of unity of organic composition," since Duges allowed the number and arrangement of parts to vary.75 As in the past, Cuvier was most alarmed by the attempt to link embranchements. To establish "conformity" between classes, he asserted, "one is obliged to deliver oneself to unproven hypotheses and to improbable interpretations." How were comparisons such as that between the five thoracic legs of Crustacea and the arm of man to be understood? he inquired, and insisted that if they could not be understood in a real and physical sense, they were meaningless: Was there once a time when men had twenty feet? Was there a moment during pregnancy when these twenty feet existed in the embryo? Or else, if that has never taken place, . . . in what physical or intelligible sense is it possible to say that these members are the result of a fusion?76

How could even the boldest imagination transform in his mind the legs and mouthparts of articulates into vertebrate arms, legs, and jaws once all the muscles, nerves, and vessels were considered? An animal body could not be divided and molded as if it were "made of pate." The differences between vertebrates and articulates, Cuvier reiterated, were "primitive and essential."77 Disappointed at Cuvier's response, Duges nevertheless published his memoir in book form in 1832 and hoped for a better reception from the Academie now that Cuvier was gone. In the preface, he admitted that, despite his best efforts, he had not succeeded in distancing himself from the quarrel before the Academie, nor had he been able to counteract Cuvier's repugnance for any generalization in natural history that was the least bold. "He announced a severe judgment to me," Duges wrote, "and I do not know to what point I managed to soften the harshness of it in a long conversation in which I did not disguise either the strengths or the weaknesses of the doctrine submitted to his examination."78 When the book was presented to the Academie in 1832, Blainville was named to present a verbal report. Five months later, Duges was still

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requesting that the report be made. Geoffroy, then president of the Academie, reminded Blainville of his obligation, but no action was taken.79 Undaunted, Duges continued to elaborate his system of philosophical anatomy until his untimely death at the age of forty in 1838. His more descriptive work in comparative anatomy, which he continued to submit to the Academie, finally earned him the sought-after place of correspondent of the section of anatomy and zoology in 1835. Geoffroy and Serres were among those who warmly supported his candidacy.80 As this minor episode illustrates, the debate raised problems that were both pressing and challenging, but this was not the time for clear-headed discussion of biological issues. Well aware of the politics of the situation, the core group of naturalists at the Academie tried to stay out of the controversy. Those who came forward in support of Geoffroy at this time were mostly naturalists on the fringes of the community. Duges represented just such a supporter, a young physician and naturalist from the provinces, genuinely excited by the debate, and eager to come to a resolution of the issues. But as an outsider, he was naive concerning the workings of the Academie and the response that his paper would generate. Both Cuvier's and Geoffroy's reactions were by now rigid and predictable. This was no time for a compromise, even if less far-fetched than Duges's, to be welcomed with any enthusiasm by anyone.

The Controversy Continues

The official debate at the Academie was called to a halt on 5 April, but the battle fires were by no means extinguished. If anything, in the two years that followed, the flames burned with an ever greater frenzy. Both opponents continued to wage battle through all that they did: in their published papers, in occasional clashes at the Academie such as that provoked by Duges, in Cuvier's course at the College de France, and in his notorious eloge of Lamarck. Once brought to the attention of the public by the debate of February to April, 1830, the issues in all their ramifications remained in public view all through 1830, 1831, and 1832. The continual parrying may have excited the public and made for good press copy, but was hardly likely to have been welcomed by the Academie. For those who aspired to leading roles in Parisian science, the unending controversy produced an uncomfortable tension which had the effect of limiting free discussion. Geoffroy's behavior from 1830 on increasingly alienated him from the members of the Academie. His grandiose claims for his work, his constant demands on the Academie's time, his hypersensitivity to anything that might be considered a slight—all this must have sorely tried the patience of his colleagues, even those with some degree of sympathy for his views. Colleagues no doubt found equally unbearable Cuvier's repetitious pontifications on scientific method. The disingenuous, superficial, and satirical manner in which Cuvier treated the theories of his opponents probably seemed to his fellow aca-

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demicians as unjust as it was unwarranted. The two adversaries had long ago closed themselves off to any modification of doctrine or of habit. The Revolution of July 1830 brought about little change in the controversy. Not long after Geoffroy's book appeared, the uprising long feared by Cuvier took place, resulting in the replacement of the reactionary reign of the Bourbon King Charles X by the more liberal regime of Louis-Philippe, due de Orleans. Geoffroy applauded the event, seeing in it the re-establishment of "our national liberties." A humanitarian rather than a political partisan, he sheltered for two weeks the deeply compromised Archbishop of Paris.81 On his way to England when the Revolution broke out, Cuvier decided to continue his journey. In Britain the report was spread that Cuvier had deliberately fled from France.82 Once the Revolution was accomplished, however, Cuvier was not unhappy with the change of regimes. He retained all his positions and added to them that of peer of France. By late August he was back in Paris ready to resume battle against Geoffroy. At the Academie tensions flared up again in October when Geoffroy's brand of transmutation of species was finally introduced full-blown into the conflict. On 4 and 11 October, having recently traveled to Caen in Normandy, Geoffroy read two memoirs in which he again broached the possibility that the fossil crocodiles of Caen were the ancestors of modern crocodiles.83 Angered by Geoffroy's insinuation that he had failed to recognize a supposedly distinguishing feature of crocodile anatomy, Cuvier took the floor to protest. The point at issue was technical: whether the petrous portions of the temporal bones were modified in crocodiles, as Geoffroy maintained, to become separate bones found above the brain, united and sutured in the form of an arch. In the heat of the argument, Cuvier promised to prepare a memoir for the next meeting in order to respond to the "false allegations" of his colleague. Geoffroy took umbrage at this and petulantly declared that "since on all points he found himself opposed by such demanding sensitivities," he would refuse to present his memoir on Steneosaurus in which he planned to examine whether current species descended from extinct species. The editor of the Gazette medicale, who reported on the meeting, added, "Let us hope that this hasty resolution will be changed!"84 A resumption of the debate was only narrowly averted. On 18 October the hall was full of curious spectators, but Cuvier disappointed them by refusing to speak. The next week the crowd returned. This time the president declared that if Cuvier recommenced his debate with Geoffroy, he would call a closed meeting of the Academie. Cuvier may well have arranged this decision with the president ahead of time. In any case, he agreed to table the reading of his memoir indefinitely.85 It was clearly Cuvier's policy not to be drawn into another direct collision with Geoffroy. Geoffroy, not one to shrink From Further confrontation, had prepared a rebuttal, which contained the first of many references to Goethe's commentary on the debate. When Cuvier did not speak, Geoffroy had his paper—a short, mostly polemical piece on the petrous portions of the temporals—printed in the Gazette medicale. In it Geoffroy alluded to "the first authority of Ger-

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many . . . the celebrated Goethe . . . who has just accorded my work the greatest honor that a French book can receive."86 One can assume that the members of the Academie for the most part did not share Geoffroy's high estimate of Goethe as a scientist, and that Geoffroy did little to further his cause among academicians by his continual references to the German poet. In 1831 Geoffroy was probably the most frequent speaker at the Academie, presenting or communicating papers on at least nine occasions between January and August. On 3 January he read a note "On the Physiological Theory Designated Under the Name of Vitalism," in which he avowed "that there could be no other laws than the general laws for the explication of affinities of bodies, whether brute or organized." Despite his earlier resolve not to continue his paleontological papers, he returned to the fossil crocodiles in February and read or communicated four more papers, including, on 28 March, the highly controversial "Memoir on the Degree of Influence of the Environment in Modifying Animal Forms," which was discussed in the previous chapter.87 All of Geoffroy's papers were intended to provoke his opponents, but Cuvier remained silent in the face of these assaults. Cuvier, by contrast, presented few papers and reports to the Academie, but he was every bit as preoccupied as Geoffroy with carrying on the struggle. The only papers he presented to the Academie between his note on the dodo in July 1830 and his death were a memoir on the formation of the bones of the sternum read on 2 January 1832 and a memoir on the development of cephalopods read on 2 April. Having neglected embryology during the debate, Cuvier now presented two embryological studies, written in a seemingly descriptive style, but obviously aimed at Geoffroy. In the former he showed that the sternum of birds did not invariably arise from the same number of points of ossification. Sometimes there were two points of origin, and sometimes five. Although the obvious intent of this paper was to undermine the embryological argument for unity of composition, Cuvier made no reference to Geoffroy. Nonetheless, Geoffroy and Serres were up in arms, Serres to defend epigenesis and Geoffroy to accuse Cuvier of reopening the debate. Cuvier calmly responded that he had said nothing for or against epigenesis, and had no interest in discussing general questions. He had merely brought up a point in descriptive anatomy. Cuvier published the memoir in the Annales des sciences naturelles, to which Geoffroy responded by his own memoir defending the "theory of analogues."88 Cuvier's final memoir took on Geoffroy's and Serres's contention that mollusks could be linked to vertebrates through the theory of arrests of development. In this paper, ostensibly a description of the fetus and fetal membranes of a cuttlefish, Cuvier pointedly noted that the umbilical cord attached to the embryo neither by the ventral surface as in vertebrates, nor by the dorsal surface as in articulates, but below the mouth in a place peculiar to cephalopods.89 The moral, never explicitly stated, was nevertheless clear. On other fronts, Cuvier planned a careful strategy. His plan of attack was two-pronged. On the one hand he accumulated facts that contradicted unity of composition and indefinite variability of species, and on the other hand he continued to speak out against all speculation in science, while not mentioning

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Geoffrey specifically by name. On the factual side, he spent the last years of his life preparing a new edition of his Lecons which was to contain the anatomical refutation of unity of composition. To combat Geoffroy's memoirs on evolution, Cuvier planned to write a work entitled "On the Variety of Composition of Animals," for which he drafted part of an introduction. In this polemical work, he promised to examine one by one the works of authors who joined "positive observations" to rational generalizations. He would disentangle truth from error, and reduce the generalizations to their just proportions "in making known the limits Nature has assigned to them." Two theories singled out for special scrutiny were the theory of arrests of development and Lamarckian evolution.90 Cuvier's repeated attacks on speculation in science served to reinforce in the public mind Goethe's contention that the debate concerned facts versus theories or analytic versus synthetic science. Extraordinarily defensive to the point of paranoia in his last years, Cuvier seized upon the assignment to write an eloge of Lamarck as a golden opportunity to present an object lesson on the evils of generalizing in science. Historians who have referred to this eloge have typically focused on Cuvier's treatment of Lamarck's ideas and methodology.91 But the abusiveness of Cuvier's language in what was after all supposed to be a eulogy can only be fully understood in the context of recent events. His barbs were directed as much at current opponents—Geoffroy, Blainville, Serres, the Naturphilosophen, and their supporters—as they were at Lamarck. Cuvier opened his eloge with a disparaging comparison of the work of Lamarck with that of another recently deceased scientist, Alessandro Volta. Volta, Cuvier wrote, was among that small number of men "endowed at once with an elevated mind and perfect judgment" who have produced only "certain truths," given only "evident demonstrations," and deduced only "inescapable consequences, never allowing themselves to advance anything hazardous or doubtful." To this paragon, Cuvier contrasted another group of scientists: Others, with minds no less vibrant, no less capable of seizing new insights, were less severe in the examination of evidence. To the venerable discoveries with which they have enriched the system of our knowledge, they were unable to restrain themselves from wedding fantastic conceptions. Believing themselves capable of dispensing with experiment and calculation, they laboriously constructed vast edifices on imaginary bases, similar to those enchanted palaces of our old novels that one can cause to vanish by breaking the talisman on which their existence depends.92

In such a category of thinkers fell Lamarck. Only his botanical classification for the Encyclopedie methodique and his work on invertebrate taxonomy were judged by Cuvier worthy of him. Throughout the eloge, Cuvier emphasized that Lamarck's theories, each one linked to the next, were peculiar to him and accepted by no one else. Without experimenting, Lamarck had invented his own chemistry in opposition to Lavoisier, and from 1802 on, he had invented his own physiology to match.

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Spontaneous generation with the aid of heat produced the lowest organisms, and as for the more complicated beings: Nothing, said he, was so easy to comprehend. Let the orgasm excited by the organizing fluid be prolonged. It will increase the consistency of the containing parts, it will render them susceptible to react on the moving fluids that they contain. There will be irritability, and irritability will lead to sensation. The first exertion of the being thus beginning to develop ought to tend to enable it to subsist, to form itself a nutritive organ. Voila, an alimentary cavity! Other needs, other desires, produced by the environment [les circonstances], will lead to other exertions which will give birth to other organs . . . It is by virtue of wanting to swim that membranes appear on the feet of water birds; it is by virtue of going to the water and not wanting to get wet, that the legs of shore birds are lengthened [etc.].93

Thus, in this eloge, the only place where Cuvier discussed Lamarck at length, his critique was satirical, superficial, and mean-spirited. Lamarck's family was appalled by it. Asked by others in the Academie to make changes, Cuvier refused. The eloge created such a stir that it was not read until after Cuvier's death, and then in a truncated form. The Academie's response to the eloge suggests that Cuvier was straining the patience of his colleagues nearly as much as Geoffroy.94 Yet another front on which Cuvier carried on the battle against speculation was in the Nouvelles annales of the Museum. This series commenced in 1832 with an "Advertissement" in which Cuvier dedicated the journal in no uncertain terms to the exposition of fact alone. Experience, Cuvier wrote, showed that only well-observed facts—descriptions and figures of new species, new characters which will facilitate classification, new facts on the history of animals, and details of animal anatomy—were durable. Such contributions to science retained their value over time, while hypotheses fell by the wayside: Everyone can see, on the contrary, that pure conceptions of the mind, theoretical dissertations, hypotheses [are] as variable as the imagination which creates them, and change back and forth from year to year. Whatever luster they can throw, whatever stir they can make at the moment they appear, they soon fall into the same oblivion as have fallen all the hypotheses or theories that have preceded them.95

The new journal, Cuvier asserted, would contain only observed facts and their immediate consequences. Those who wished to give vent to their theories or "meditations" were welcome to publish them in separate works, "but not in this collection, which will be composed only of positive facts."96 Cuvier's colleagues at the Museum were not altogether in sympathy with this harangue, for despite the severity of the prospectus, Geoffroy was able to publish in the same volume his final determination of the bones of the hyoid in the four classes of vertebrates, complete with labeled colored figures. Geoffroy did, however, make a special point of observing that his results were so clearly exhibited that even the supporters of positive facts would be satisfied.97 The same theme of facts versus theories appeared in the final arena where

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Cuvier took up the cudgels against his opponents: his course in the history of science at the College de France. He had scarcely taught a course for fifteen years when, in 1829, he took the platform to begin a series of lectures which traced the progress of science from the Egyptians to the present day. What better vehicle to exhibit the fate of undue speculation! The opportunity to hear France's premier naturalist discoursing on subjects that everyone could understand brought an immense crowd to the lecture hall. Le Globe published the lectures as soon as they were delivered, and a disciple prepared his notes for later publication in book form. From the careful observer, Aristotle, and the a priori theorist, Plato, Cuvier pointed to example after example of the supposed triumph of good scientific method over bad. He had in his lectures, he avowed, "put the human mind to the test," and observed which works survived the passage of the centuries and which had passed on "without utility for science." "All the hypotheses of which I have spoken have fallen, and probably there are many others which will disappear in their turn. The facts, on the contrary, the real truths, founded on experience, have remained and will remain immutable."98 Not everyone approved. Alexander von Humboldt whispered criticism of the lectures into the ear of his young protege Louis Agassiz, seated beside him. He defended unity of composition and his countrymen against Cuvier's slurs. Ampere confronted Cuvier publicly in his own course at the College de France.99 Cuvier continued lecturing through 1830 and 1831 until just five days before his death. On 8 May 1832, after an interruption necessitated by the cholera epidemic, Cuvier reached a high point of his course in a general lecture on Naturphilosophie, evolution, and unity of composition. The baron Pasquier, who was in attendance, found it a religious experience: There was in this last part of the lecture, a calm and a justice of perception, an open revelation of the intimate and complete view of the religious observer, recalling involuntarily the book which speaks of the creation of all humankind, Genesis. This affinity, rather avoided than sought out, which was not found in the words, but in the ideas, came to light suddenly when the professor pronounced these words: Each being contains in itself, in an infinite variety and an admirable predisposition, all that is necessary for it. Each being is perfect and viable, according to its order, its species, and its individuality.100

It was a fitting finale to Cuvier's career. The very next day Cuvier felt the onset of a painful and paralyzing malady. With Cuvier's death a few days later on 13 May, one phase of the controversy over philosophical anatomy was at an end. The Debate and the Community of Naturalists

If the debate were judged solely as an academic contest, there is little doubt that Cuvier would emerge the victor. His manner of argumentation—cool, controlled, crisply clear, and clever to the point of being patronizing—was calculated to produce conviction in his audience. By contrast, Geoffroy appeared excitable and defensive, the organization of his arguments was diffuse, his sen-

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tences were convoluted, and his reasoning often obscure. As Cuvier made fully evident, Geoffroy had failed to give a clear definition of "unity of organic composition" or of the "theory of analogues," let alone prove their applicability to the entire animal kingdom. Insofar as the debate concerned unity of composition or evolution, Cuvier could be said to have won the day. Yet, while even the supporters of Geoffroy agreed that Cuvier had had the upper hand in the debate, there were many who felt that judgment ought not be so hastily rendered. While unity of composition in the entire animal kingdom might be untenable, yet there was more to the theory of analogues, or philosophical anatomy, than Cuvier was willing to admit. Cuvier had oversimplified and too quickly dismissed Geoffroy's doctrine. The debate had little effect in stemming interest in philosophical anatomy. Perhaps the most important outcome of the debate was that it polarized anatomical doctrine and presented a challenge: that of reconciling the opposing claims of morphology and teleology within a nonevolutionary framework. It left hanging the question of whether there were unalterable "laws" of animal organization, and how these laws might flexibly accommodate the evidence of homologies both between and within animals with the evidence of adaptation to seemingly predetermined ends. The removal from the scene in 1832 of the most uncompromising critic of the entire enterprise of philosophical anatomy allowed for a softening of viewpoints, and at last made possible attempts at a compromise. The doctrines of the major zoologists and anatomists in France in the 1830s and 1840s—Blainville, Flourens, Isidore Geoffroy, and Milne Edwards—all combined elements from both Cuvier and Geoffroy. Their work will be taken up in the final chapter of this book. Here we shall be concerned with the reactions of naturalists in the immediate aftermath of the debate. How did they pronounce upon the arguments brought forth by Cuvier and Geoffroy? We have already noted the sympathetic responses of several of Geoffroy's supporters. Gaspard-Joseph Martin SaintAnge, a young physician and embryologist who collaborated on a paper with Isidore Geoffroy, and Jean Baptiste George Marie Bory de Saint-Vincent, an eccentric naturalist-voyager, army officer, and early evolutionist, both wrote glowing reviews of Principes de philosophie zoologique. Like the reviewer in Le National, they interpreted the debate as concerning the fate of a new scientific method introduced by Geoffroy and the Germans to replace the inadequate Aristotelian method that had held sway for twenty-two centuries. Duges, as we have seen, was sufficiently stimulated by the debate to revise philosophical anatomy to take into account some of Cuvier's criticisms. Two of Geoffroy's major allies also came forward to back him. Geoffroy's chief disciple, Serres, did not write directly on the debate, but in an article in 1834, he did reiterate his view, in support of Geoffroy, that the vertebrates could be linked to the invertebrates through the theory of arrests of development. When invertebrates were compared to adult vertebrates by the method of "descriptive comparative anatomy," the hiatus between vertebrates and invertebrates appeared insurmountable. But once preformation of

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germs was abandoned and recourse was had to "the rules of transcendental anatomy and embryogenesis," Serres claimed, the true affinities between the two groups would be readily uncovered. If invertebrates "are considered for what they appear to us to be—permanent embryos—and if their organization is compared to the embryogenesis of vertebrates, their differences are effaced and from their analogies a great many hitherto unperceived affinities emerge."101 Dutrochet supported his protector, Geoffroy, through a paper presented at a provincial society in August 1830, "On the Spirit Which Ought to Prevail in the Study of the Natural Sciences." He defended the controlled use of reasoning in the natural sciences, as well as the necessity for the scientist to have a knowledge of all branches of science, not just a single specialized area. In 1834 he presented a paper to the Academie des Sciences on the relations between vertebrates and invertebrates, in which he adopted and expanded upon Geoffroy's views on the rotation of the body of articulates with respect to the surface of the earth.102 While some of Geoffroy's supporters tended to think the whole future of science was at stake, Cuvier and his disciples minimized the entire affair. Cuvier made light of the debate in his report on the work of the Academie for 1830. "It was an amicable polemic between naturalists who have for each other a just esteem," he said. The issues were purely anatomical: whether the resemblance of plan and composition that everyone agrees to exist among the vertebrates extends to the other embranchements of the animal kingdom, and if, among the vertebrates themselves, this resemblance holds to the point where it can be called identity of composition, or as M. Geoffroy first expressed it in absolute terms, if the same parts repeat indefinitely in animals.103

Cuvier's main followers wrote no articles on the debate in the 1830s. Laurillard, Duvernoy, and Flourens, while they passionately defended Cuvier's doctrine, scarcely mentioned the recent debate in the hagiographies they wrote shortly after Cuvier's death. Most of the leading naturalists at the Museum and the Academie des Sciences—Latreille, Audouin, Dumeril, Milne Edwards, Flourens, Frederic Cuvier—avoided publicizing the conflict or explicitly taking sides in the 1830s. Only much later did a few of these naturalists venture their opinions in print: Isidore Geoffroy in his 1847 biography of his father, Flourens in an article written in 1864, and Blainville in a book Cuvier et Geoffroy Saint-Hilaire, written in the 1840s but not published until 1890. Since these commentaries were written after Geoffroy's death, in the light of the last years of Geoffroy's career and the popular response to the debate in the 1830s and 1840s, it will be best to leave them for the final chapter of this book. By 1835, the major nonevolutionary interpretations of the debate in the period before 1859 had already been brought to light. On one level, as Cuvier stated, the controversy was a question of anatomy. Was animal structure or animal function paramount? Was the concept of homology new and how far could it be extended? Were there any homologies at all between embranchements (or major groups of animals)? Within an embranchement, to what extent was there a unity of plan?

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The debate was also viewed from a wider perspective. The two naturalists could not agree on the anatomical problems because they differed as to what constituted a scientific theory. On this level the debate was concerned with the scope and aims of natural history. Was natural history to be the study of the whole of nature in order to find general laws governing animal generation and organization, even though those laws were not yet subject to complete verification, or was it the detailed enumeration of species? What were the relative roles of facts and ideas, analysis and synthesis in science? Related to the search for laws of organization was the religious issue. Did laws of organization exist aside from the principle of "conditions of existence," or did God's freedom to create preclude the possibility of any generalizations on animal structure? Was recourse to final causes desirable in natural history, and were final causes sufficient to account for the phenomena? If not, how were organs that changed function and useless organs to be reconciled with Providence? Did the Creator also manifest Himself in establishing ideal patterns of organization that could be perceived by the rational mind? Was transcendental anatomy necessarily pantheistic and hence irreligious? Evolution, though raised by both sides in the debate, was only a secondary concern. Although the causes and extent of variation, and whether new species could be produced over time, were old and continuing problems, no one saw them as the focal point of the debate. Most of the partisans of philosophical anatomy understood homologies in ideal terms, as patterns imposed by the Creator upon nature, and not as evidence for the transformation of species. There were also the institutional and professional issues raised by Goethe. Should scientific ideas be controlled in the interests of a small elite, or should everyone be free to contribute his ideas and receive a fair hearing? If science was a public concern, what responsibilities did scientists by profession have to the public? How much access ought there to be to what went on in the Academie? Finally, there was the relation of the debate to the philosophical and social concerns of the Revolution of 1830. The relationship was indirect, but nevertheless real. To a segment of the educated public, the Revolution of 1830 represented political liberty, freedom of speech and the press, and freedom from domination by the Church. Geoffroy's doctrine and his right to promulgate it could be interpreted in the light of the rhetoric of liberty. Philosophical anatomy also represented a form of liberty to contemporaries: the liberty to progress beyond outworn formulas based on traditional religious beliefs and to ask broad and fundamental questions about nature. The broader philosophical, religious, and social concerns were amplified in the 1830s and 1840s when the Cuvier-Geoffroy debate passed beyond its immediate context to become a romanticized historical event laden with significance for the future. It was in large part because of Geoffroy's activities of the 1830s that the debate became entrenched in French popular culture. Freed by the end of the repressive Restoration and the death of Cuvier, Geoffroy could indulge himself, as he once had in Egypt, in searching for the fundamental materials and forces of the universe. He fueled the growing interest in the

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debate of 1830 by his philosophical writings directed to the public and offering a progressive and optimistic view of man and nature, attractive to a variety of reformers. In the hands of novelists and popular and political writers, Cuvier, Geoffroy, and the debate became symbols of what was wrong with French science and its institutions. It is to the debate in this larger setting outside the Academie that we now turn.

CHAPTER 7

Beyond the Academie: The Many Uses of the Debate

In 1835 the French writer Honore de Balzac sent to Geoffroy a copy of his new philosophical novel Louis Lambert, the story of a student of life. Among Lambert's friends, Balzac introduced a friend of his own youth, "Dr. Meyraux," the Meyranx of the Cuvier-Geoffroy debate. In the story Lambert and Meyraux were drawn by their mutual interest in unity of type to the course of comparative anatomy at the Museum. As a philosopher rather than a scientist, Lambert attended the lectures not for the details of animal structure, but because the study of comparative anatomy threw light on "the real relation that may exist between God and man."1 In a second novel, Un grand homme de province a Paris (1839), Dr. Meyraux appeared once again, this time as a former colleague of the hero, and was identified in his relationship to the Cuvier-Geoffroy debate: Meyraux died after stirring up the famous controversy between Cuvier and Geoffroy Saint-Hilaire, a great question which divided the whole scientific world into two opposite camps, with these two men of equal genius as leaders. This befell some months before the death of the champion of rigorous analytical science as opposed to the pantheism of one who is still living to bear an honoured name in Germany.2

Yet a third Balzac novel, one of his most famous, Pere Goriot, was dedicated in its second edition (1842) "To the great and illustrious Geoffroy Saint-Hilaire as a tribute of admiration for his labors and his genius."3 Balzac was but one of many writers and propagandists of the 1830s and 1840s who came to see Geoffroy as a heroic figure, Cuvier as a paltry fact collector, and the debate as a major event in French intellectual history. Why, we may ask, did a novelist such as Balzac introduce the debate into his books, and why did he and others outside the scientific community choose to champion Geoffroy over the former revered chief of French science, Cuvier? What meanings did the debate have for nonacademicians, especially for those who were not in a position to judge for themselves the technical anatomical arguments of the 1830 memoirs before the Academie?4 175

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After 1832, the debate, as perceived by a new audience in a new milieu, became transformed in character. No longer were the strictly scientific issues— unity of composition and the theory of analogues—predominant. No longer were scientists and physicians the primary audience. Other issues present in the debate of 1830 and its immediate aftermath, but not emphasized at that time, emerged with new importance: philosophy and religion, the conflict between the professional zoologist and the philosopher of nature, and the institutions and politics of science. Concerns such as these, and not philosophical anatomy in itself, brought the debate to the attention of intellectuals such as Balzac. That the controversy between Cuvier and Geoffroy touched many outside the Academie in these decades was in large part due to Geoffroy himself. In the years after 1832, Geoffroy made a deliberate effort to reach a sympathetic public, both through the style and content of his writings and through personal communication. His pronouncements of the 1830s, depending very little on new research, took him further and further away from the zoology of the Museum and Academie and into a new world peopled by philosophers, writers, social reformers, and political activists. The more Geoffroy insisted on his unconventional speculations, the more the academic scientists disowned him, the more he turned to other avenues of support. He came to see himself as engaged in a mission to convert his countrymen to his vision of a progressive, Newtonian universe. He succeeded in only alienating the scientists, but he found others outside the scientific community who were ready to respond. The young intellectuals who gravitated toward Geoffroy in the 1830s formed no unified group and had no single reason for their interest in his work. Some were unknowns—a cleric who wrote a review of Etudes progressives, a medical student who wrote a dissertation on soi pour soi—who were mentioned briefly in Geoffroy's works. Many of the writers and political activists who attended Geoffroy's Sunday night salons held at his home in the Jardin des Plantes in the 1830s may have been familiar names in their day, but are now all but forgotten.5 Still others have become major figures in French history: the writers George Sand and Balzac, the historian Edgar Quinet, and the physician, scientist, and politician Francois-Vincent Raspail. All of these literary talents, finding Geoffroy's doctrines appealing for philosophical or political reasons or both, employed their pens to further his cause. Perhaps the major attraction of Geoffroy's writings was his romantic and progressive vision of the universe. With the advent of the more liberal reign of Louis-Philippe, with Cuvier gone, and with the realization that he was nearing the end of his own career, Geoffroy returned to the grandiose speculations of his youth. He freely indulged himself in the search for the fundamental laws of the universe, uniting organic and inorganic matter, and epitomized by the universal law of soi pour soi. Geoffroy's universe, created by God and continually evolving to an ever greater perfection, presented a welcome alternative to the traditional religious world-view, an alternative that offered the possibility of social reform. As commentators interpreted him, Geoffroy saw nature not in static terms and limited by the arbitrary will of an autocratic God, but as essentially plastic and progressive.

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Nonspecialists also admired Geoffroy for his scope of vision. Geoffroy did not shrink from dealing with the broad issues raised by natural history, the issues that were obviously of vital importance but that other savants deliberately ignored. Those who protested the division of science into specialties presided over by professional researchers found in Geoffroy an ally. He was lauded as the leader of a new synthetic movement in science, allied to Naturphilosophie, that would return science to what it was meant to be. Philosophical motives for interest in Geoffroy and the debate could easily shade into political ones. If Geoffroy's scientific doctrine was of such worth to mankind, why was it not more commonly accepted? It could be argued that Geoffroy failed to convince the other savants because the institutional and political structure of official French science was stacked against him. Critics denounced the Academie as riddled with factions and opposed to all innovation in science. The government, by exerting too much control over science, and the Catholic Church, on which the government relied for support, were also assigned a share of the blame. It seemed to many as if the savants, in the employ of the government, had accepted without question the religious worldview, and now went about their timid business of classifying and describing, refusing to examine any further the fundamentals of their science. They were depicted by would-be reformers as mere fact-collectors, more interested in fame and fortune than in the search for truth. Cuvier was seen as the chief villain responsible for the supposed decline of science since the eighteenth century. In short, by championing Geoffroy, the underdog in the debate, one could protest the advent of modern professional science, its loss of scope and relevance, and the elitist institutions that perpetuated it. The many uses of the debate will be explored in this chapter by looking at a number of individuals, all nonacademicians, who took up Geoffroy's cause in the 1830s and 1840s. The philosophical and religious appeal of Geoffroy's doctrine is illustrated by the writings of Sand, Balzac, and Quinet. The trenchant critique of official science, its institutions, and its system of patronage is represented by Balzac's satire "The Ass's Guide for the Use of Animals Who Wish to Achieve Honors," by the writings of Raspail, and finally by a clever and obscure satire on the professors of the Museum published by a journalist, Bertrand-Isidore de Salles, and a scientific popularizer, Frederic Gerard. The responses of nonacademicians are more than a side issue to the study of the reaction of the scientific community to philosophical anatomy and the debate. When the leading zoologists of the following generation wrote on the debate, they did so in part because the debate was already a part of the common culture. A familiarity with the contest between the two great rivals was presupposed by Blainville and Isidore Geoffroy in their books and by Flourens and Milne Edwards in their articles and eulogies. Their treatment of Cuvier and Geoffroy and their interpretation of the controversy were conditioned by the prior public response of the 1830s and 1840s. To understand with any depth the reasons why writers such as Balzac became interested in Geoffroy and the debate, one would have to study not only the careers of the various individuals involved, but also the political, intellectual, and social milieu that engendered their writings. We propose here

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only the more modest goal of suggesting how a scientific controversy, removed from its immediate context, takes on a life of its own. It can be endowed with many meanings and many uses, each appropriate to a given time and place. A Natural Philosopher and His Audiences

In 1835 Geoffroy predicted that it would be one day said of him that he had rendered to society two great services: (1) the revelation of the all-encompassing law of soi pour soi, and (2) bringing Cuvier to Paris.6 This remark captures in a nutshell the conflict that permeated all of Geoffroy's career between the need for professional recognition and the desire to pursue science according to his own lights. The second "service" was one for which the entire scientific community would have rewarded him. The first was one that nearly the entire scientific community repudiated. Earlier in his career, Geoffroy had restrained his penchant for speculation in order to maintain the support of his colleagues and the possibility of professional advancement. Even during the debate and its aftermath, Cuvier's presence had at least a moderating influence on Geoffroy. But after 1832, there was no one to take Cuvier's place as "legislator of science." Geoffroy, nearing the end of his career, no longer felt the need to abide by professional restrictions in his writings. Freed from the usual constraints imposed by a scientific community, Geoffroy set out to develop further the system of the universe that he had sketched out in Egypt in 1801, combining his old ideas with what he had learned from the study of teratology and teratological evolution in the 1820s. The result was the delineation of a universe set in motion by God, and then allowed to progress according to the interplay of a single system of laws for the physical and biological sciences. It was a nonvitalistic system, but also nonreductionist, for the primary law, that of soi pour soi, derived from biology. Through the 1830s, Geoffroy persisted in promoting his ideas at the Academie, but his erratic and unprofessional behavior increasingly alienated him from his colleagues. He had lived long enough to achieve the status of grand old man, one of the few survivors of the Golden Age of French science, and for that he was given some measure of indulgence. Yet the Academie members had only limited patience with his frequent rambling and polemical presentations. To Cuvier, Geoffroy's ideas had posed a danger to science and society. To his colleagues of the 1830s, Geoffroy was less a threat than an embarrassment and a nuisance. When Cuvier died, Geoffroy had hopes of assuming some of the mantle of Cuvier's authority in the scientific community, but in this he was only marginally successful. Soon after Cuvier's death, on 28 May 1832, Geoffroy announced in a note to the Academie his candidacy for the post of permanent secretary. In support, he presented his 1832 collection of funeral eulogies, Goethe's two articles on the debate, and a work on the fossils of Normandy, presumably dealing with evolution. His bid to inherit this most powerful of posts fell flat. In the first of two ballots against Flourens and Pierre Louis

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Dulong on 9 July, he received only seven of forty-five votes and in the second ballot a mere two. When the victor, Dulong, resigned the post in 1833 because of ill health, Geoffroy was no longer a candidate, and Flourens was elected in a three-way contest with Dumas and the philosophical botanist Auguste de Saint-Hilaire. Retaining the post until his death in 1867, Flourens eventually assumed Cuvier's role of guardian of scientific orthodoxy.7 Geoffroy was compensated in some measure, however, by his close victory over the geologist Pierre-Louis-Antoine Cordier, to become vice-president of the Academie for 1832. It was a fitting recognition of his status as an elder statesman of science. When in 1833 he automatically assumed the presidency for the year, he gloried in the exercise of this largely honorific post.8 Geoffroy's output of books and memoirs in the 1830s was enormous. In addition to a set of eulogies and a biographical account of Buffon, he published two major works of natural philosophy, Etudes progressives (1835) and Notions synthetiques (1838).9 He barraged the Academie with papers, filling the newly created Comptes rendus of the Academie with extracts as well as with additional communications. Sometimes Geoffroy would insert a piece a week for a period of months. The common theme running through almost all these publications was that there was only one set of laws in nature for organic and inorganic bodies, for physiology and psychology. Much of this work was not science at all, but rather a defense of synthetic philosophy and his right to pursue it. Often Geoffroy took someone else's research as a starting point for the elaboration of his own theories. One can imagine the reaction of Flourens when Geoffroy converted his study on the beating of arteries into an example of soi pour soi, or of the rigidly antitheoretical Magendie when Geoffroy used his work as a point of departure for a discussion of whether repulsion and attraction were truly antagonistic principles.10 Despite numerous provocations, none of the academicians undertook to answer Geoffroy in print. Instead they adopted the traditional Cuvierian strategy of ignoring what they disapproved of. Much to Geoffroy's disappointment, no scientist wrote a review of Etudes progressives or Notions synthetiques. The only response he received was that his thinking was a priori, that his philosophy tended toward irreligion, and that he had no business dealing with subjects outside his field. These criticisms were made orally; no savant attacked him in writing. On the occasion of one of Geoffroy's papers, however, the academicians registered their disgust by means of a mass exodus from the hall. This was a paper read in September 1834 on variation in plants entitled "Power of the Environment [monde ambiant] Governing the Generation of Causes to Which Are Related the Different Forms and the Multiplication of Species of Plants."11 Geoffroy admitted that in discussing botany he was leaving his field, but he excused himself on the ground of having obtained the examples he cited from a respectable source, the botanist and zoologist Moquin-Tandon, who was then on a visit to Paris. The point of Geoffroy's paper was that changes in the environment produced monstrosities in plants that were inheritable and could lead to the formation of new species. Amused by Geoffroy, but not entirely sym-

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pathetic to his foibles, Moquin-Tandon described the presentation as a "long and tiresome homily" read in "a heavy and emphatic manner." The reading of the memoir, he reported in his personal journal, "was the signal for the departure of several members of the Institut and a part of the public." Even Geoffroy's son Isidore walked out on him. Those who remained engaged in private conversations. No one listened with much attention.12 According to Moquin-Tandon, by 1834 the savants were beginning to feel that Geoffroy was losing his grip on reality. His son and Serres did their best to cool the old man's ardor, but there was little they could do to stop him from embarrassing them. As Moquin-Tandon observed, "The intellectual activity of M. Geoffroy is astonishing, but his aptitude for observation diminishes from day to day; he has the fire of youth, but not the reserve of maturity." He sympathized with Isidore Geoffroy, who, he reported, suffered greatly from his father's "bizarreries."13 Other friends suffered as well. Geoffroy was at this time part of a group of savants—the botanists Auguste de Saint-Hilaire, Pierre-JeanFrancois Turpin, and Adolphe Brongniart, the physiologist Dutrochet, the embryologists Serres and Victor Coste, and the two Geoffroys—who planned to form a society of physiology for men of broad views. One difficulty, according to Moquin-Tandon, held them back: "the mania of M. Geoffroy SaintHilaire senior to compose memoirs overnight. They are afraid he would want to fill the journal of the Society with his memoirs."14 Blainville, the reigning zoologist of the day, set off a crisis in Geoffroy's career in late 1834 by refusing to publish one of Geoffroy's reports to the Academie in the Nouvelles annales of the Museum. In his review of a memoir of philosophical anatomy by the physician, editor, and translator AntoineJacques-Louis Jourdan, Geoffroy had as usual exploited the occasion to promote his own research: his work on teratology and the necessity to advance beyond the "timid classifiers and describers" who wanted to remain at the "ABC of the science of the naturalist."15 Having sympathy with neither teratology nor speculation, Blainville set a precedent by not allowing the report to be printed.16 In a rage, Geoffroy declared his collaboration with the Museum journal at an end, and announced in January 1835 a subscription to his own series of memoirs.17 He gave his hastily thrown-together book the pretentious title Progressive Studies of a Naturalist During the Years 1834 and 1835, Continuing His Publications in the 42 Volumes of the Memoires and Annales of the Museum d'Histoire Naturelle. In the introduction, Geoffroy mourned the end of friendly cooperation among the professors of the Museum and vowed to write in the future for others: "Other times, other ways! The Annales have come to be considered an AFFAIR." He had been told, he said, that the contents of the journal ought not to "offend the general taste." "However," he avowed, "I believed myself engaged in a mission." Resolutely he had decided to publish his own volume of the Annales, and if "naturalists of the current age" did not appreciate him, he was "determined to write for those of times to come."18 He filled the book with his recent memoirs to the Academie, all of them written in a polemical style and promulgating unorthodox views: the anatomy

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and physiology of monotremes (platypus and spiny anteater), lactation in cetaceans, the review of Jourdan's memoir on vertebral processes serving as teeth, the reproductive system of the mole interpreted in the light of teratology, paleontological evolution, and finally soi pour soi. Having begun his opus with another scheme of the progress of science, culminating in "its philosophical disposition in the future,"19 he ended with the announcement of his law of laws, key to the phenomena of the universe: a vague, general law which was intended to extend Newtonian attraction to the smallest particles of matter. Exhausted by the publication of Etudes progressives, Geoffroy left Paris later in 1835 for his estate in the country, and without books or notes, wrote a second work of natural philosophy, Synthetic, Historical, and Physiological Notions of Natural Philosophy (Notions synthetiques). Although composed soon after his first book, Notions synthetiques was not published until 1838.20 As was to be expected under the circumstances, the book contained much personal justification and little science. Notions synthetiques was to deal with the application of soi pour soi to physics, but Geoffroy never got beyond a few vague generalities. Most of the work concerned Geoffroy's right to formulate soi pour soi at all. In these works Geoffroy passionately defended his right to indulge in natural philosophy rather than restricting himself to the areas of his professional expertise, namely, zoology and comparative anatomy, as expected by his scientific colleagues. The goal of science, he declared, was to seek "the harmony, the affinities, and the true reason of things."21 He regretted that this type of investigation was considered "philosophy" rather than science by "the naturalists, our adversaries, enclosing themselves in the duty of Descriptions and Classifications." But, he assured his readers, to "render manifest the sublime harmonies of organized bodies," was indeed "to do science and the best science, no doubt."22 On several occasions during the decade Geoffroy cited a line from Balzac's philosophical novel Louis Lambert—"Science is one and you have divided it"—to justify going beyond the confines of his field. Geoffroy lamented that each science was proceeding on its separate course, with its own principles invented to explain phenomena. No longer did anyone attempt to link the sciences together. "The savant of each specialty has taken the decision to isolate himself in a canton, as if living at the bottom of a well, built for their use alone," wrote Geoffroy. No wonder such specialists failed to understand him, for if they were isolated in a pit, how could they know anything of what occurred on the earth's surface?23 "Would it be rational," he asked, "if from a purely zoological proposition, consequences unfold which invade the domain of sciences that are considered distinct, as, for example physics proper, to refuse to deduce them"? To expect him to limit himself to zoology, he argued, would be to require him "to impose limits on logic itself."24 The law of affinity of soi pour soi, literally "self for self," marked the culmination of Geoffroy's search for unitary laws. Soi pour soi was to be an extension of Newton's law of gravitation (which was believed by many to be applicable only to astronomy) to the smallest particles of matter. A generalization

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of the attraction of similar parts applied to monsters in 1827,25 Geoffroy's muddy and confusing law was annouced to the Academie des Sciences in late 1834 and introduced to the public in the final memoir of Etudes progressives, entitled "Universal Law (Attraction of Soi pour Soi), or Key Applicable to the Interpretation of All Phenomena of Natural Philosophy." The memoir was to be the introduction to a projected work called Genesis of Things.26 Geoffroy claimed that he had initially discovered his universal law at Alexandria in 1801 and only rediscovered it in 1827. At great length he recounted for his readers the fateful episode in Egypt: the finding of the two electric fishes, his turbulent intellectual effort, his blindness, and the reluctant decision not to pursue his conclusions further. To underline the importance of the law, he appealed in several of his writings to Napoleon, whom he continued to admire at a time when he was largely out of favor as a revolutionary, a despot, and an opponent of the Church. Notions synthetiques, dedicated to Napoleon's memorable meditation "The World of Details," recalled Napoleon's conversation with Monge on the eve of his departure from Egypt. Geoffroy believed that by the promulgation of soi pour soi he had succeeded in fulfilling Napoleon's ambition to discover the universal laws governing the action of the smallest particles of matter.27 As an innate property of matter and the sole source of its activity, soi pour soi supposedly obviated the need for any other inherent qualities of matter, and in particular, the law was intended "to be substituted for the explanations of the vitalists."28 According to Geoffroy, matter was in essence homogeneous, and became diversified only through modification in time and space. "Imponderous" fluids (as opposed to "imponderable"—Geoffroy denied the possibility of absolute weightlessness) such as light and caloric were produced from homogeneous matter by the process of combustion, and they were returned to the state of "concrete matter" by the reverse process of electrification. The latter process entailed the attraction of soi pour soi. Combustion, according to Geoffroy's system, separated matter into solids and elastic "imponderous fluids," while electrification gathered the fluids together, returning them to the solid state, whenever identically similar substances confronted each other. A steady state was maintained through alternating acts of these two processes.29 One would have to agree with Geoffroy's adversaries that he never gave a clear explanation of what soi pour soi meant. Geoffroy adduced few facts on which to base his extension of the law of attraction of similar parts to the inorganic world. Moreover, he refused to acknowledge the need for experiment or mathematization. It was no wonder that the academicians ignored or ridiculed the aged naturalist turned philosopher.30 Geoffroy admitted that a number of readers had found his law both confusing and exasperating, but persisted nonetheless in further argument: Soi pour soi: I understand that these words which often appear in my explications fatigue those of our readers of the Comptes rendus who expect no clarity from them, nor any resolution.

He reproached himself only for having delayed so long—since his return from Egypt—in announcing his law.31 His colleagues seemed to take the position

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that the best way to deal with Geoffroy's law was to ignore it. Scientific journals refused to review his works of natural philosophy. Even his son wished to limit soi pour soi to physiology.32 Deeply hurt that the crowning achievement of his life should go almost unnoticed, Geoffroy complained in Notions synthetiques that "my idea received neither praise nor criticism, but [only] indifference and silence concerning that which I cannot help but regard as the most considered and vast of my enterprises."33 Although the historian may be tempted to laugh at soi pour soi, the law was on Geoffroy's part a serious effort; it put the finishing touch on a system of nature that had a philosophical and religious appeal to him since the outset of his career. Geoffroy's was a Deistic universe, which, while differing in details, was not essentially different from that envisioned by his predecessors Buffon and Lamarck. Geoffroy believed that God was coeval with matter. Primitive matter was a true chaos until God intervened to give it "a single, an all-powerful, a wholly efficacious property, to regulate for all time the order and administration of all creations in the universe." That property was soi pour soi. "But beyond that," Geoffroy hastened to add, "without dispute the cause of causes is GOD!"34 The universe as conceived by Geoffroy was materialistic but not machine-like or reductionist. He did not attempt to reduce all phenomena to a pre-existent set of physical laws considered to be the fundamental laws of nature. Although organic and inorganic bodies could be distinguished by their distinctive forms, they both functioned as special cases of the same universal laws. Of these laws, none was more fundamental than soi pour soi, derived from biology, but applicable alike to molecules and planets and to arteries, veins, and muscles. Thus there was no gap between biology and physics, but rather "a single science, embracing all ages, all space, and all bodies great and small."35 Extraordinarily sensitive to the veiled charges of irreligion that his work engendered, Geoffroy was at pains to defend himself from impiety. On one hand he lashed out against those who would criticize him on religious grounds. How could it be impious to observe that which is? "To contemplate the work of God—that is what those in high social positions in the government or the Church forbade."36 On the other hand, whenever possible, he referred to religious personages who supported his works. His efforts to refute charges of materialism and pantheism only added to the confusion of his doctrine. He assured his readers that although he rejected vitalism, he was not to be considered a materialist, for he believed soi pour soi acted, so to say, as "the universal soul of nature" or immaterial principle.37 In a note to the Academie in 1837 on the identity of physiology and psychology, Geoffroy promised that he would discuss further the corporeal soul of Saint Augustine.38 The Academie was happily spared. Geoffroy's doctrine continued to be equated in the public mind with Naturphilosophie, considered by many to be pantheistic. Geoffroy recalled that in 1820, Oken's Isis had proclaimed him a Naturphilosoph ["philosophe de la nature"] and the appellation had stuck. The association with Naturphilosophie, he claimed, had greatly compromised him in France: "I was, so to speak, affiliated with a sect, or so-called sect, against which war was being waged in

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Paris."39 By the 1830s, Geoffroy took a more favorable view of Naturphilosophie than he had in the previous decade. In an article titled "Pantheistic Heresies," written in 1836 for the Dictionnaire de la conversation, Geoffroy argued that pantheism in its original and proper meaning was the religious doctrine of the first ages of humanity. If its proponents tended to place the Creator and the created at the same rank, this was a fault of ignorance, not of deliberate impiety. Pantheism was never the equivalent of atheism. With the aid of revelation and the light of science man had now progressed far beyond this early understanding of the universe. It was therefore unjust to equate modern Naturphilosophie with the pantheism of another era. Whereas in the 1820s Geoffroy had tried to distance himself from Naturphilosophie, in the 1830s he had become an ardent defender of Germany, "the nation the furthest advanced in philosophical matters." In the late 1830s he made a journey to Germany, where he was treated with honor.40 Perhaps one of the most appealing features of Geoffroy's universe to supporters outside the scientific establishment was its progressivism. This aspect derived not from an inherent tendency to progression, as in Lamarck's theory of evolution, but rather from unidirectional changes in the atmosphere and temperature of the earth, which in turn required animals to become modified in order to survive. Thus Geoffroy insisted on progression of animals in the geological record. When an anomaly appeared, such as the discovery of marsupial bones from the secondary era when all previous mammalian bones had been restricted to the tertiary era, Geoffroy countered in 1838 with the proposition that marsupials and monotremes should not be considered mammals at all, but rather a separate class lower in the scale.41 A theory of evolution was almost a necessary corollary of Geoffroy's general views on God, nature, and natural law, for the alternative was the entirely unacceptable proposition that God had intervened from time to time to create new animals. Throughout the 1830s Geoffroy continued to present papers on fossil vertebrates and their evolutionary implications. He made several trips to Auvergne and Normandy to survey recent fossil discoveries, particularly new remains of crocodilians. These fragments of a former world were, he declared, "the witnesses of so many links in a progressive series and of a continuous filiation."42 In place of "three distinct and isolated creations" corresponding to the three major divisions of the geological record, Geoffroy advocated "only one system of creations" that was progressively altered "under the all-powerful influence of the environment [des milieux ambiants]."43 The "two zoologies"—the animals of former times and those of the present day—"follow each other without lacuna or interruption, engendering one from the other" through modifications produced by the environment.44 After 1831, Geoffroy downplayed the teratological component of his evolutionary doctrine, emphasizing instead the general action of the environment on a basic fund of materials of animal organization. At times he wrote of modifications as accidentally produced. At other times, he wrote of modifications as preordained by the Creator. "Each thing arrives at its pre-fixed moment," he declared in Etudes progressives. "It is in this sense that one can say that the birth of man was from all eternity in the plan of Providence, yet to appear only

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at an anticipated day" when the environment had acquired the proper conditions for man's development.45 The gradual perfection of animal structure was followed by a gradual perfection in the moral development of man. The history of man, according to Geoffroy, was that of the progressive development of humanity from darkness to intelligence. In some places, Geoffroy indicated that God had intervened so that man, without any further progression in material structure, would be provided with the conditions for "a perfection which is more psychological than physiological." At that moment in history, he said, man became capable of penetrating the designs of God.46 In other places, Geoffroy insisted on the continuity of man with the animals. In a series of notes in the Comptes rendus in 1836, Geoffroy contested the common practice of placing man in a separate order of mammals from the primates. His point of departure was the orangutan of Sumatra, then on exhibit in the Museum menagerie. Neither monkey nor man, the orangutan, according to Geoffroy, disrupted the standard classificatory system separating Bimana (man) and Quadrumana (monkeys and apes). He argued that neither differences in anatomy nor differences in moral qualities sufficed to set man apart from the apes, for the moral qualities of man could be reduced to his physical conformation. In past ages, man and orangutan were similar in conformation and habits, but in the course of time the two species had developed inversely, the orangutan gaining in physical force while man gained in intellectual function.47 The contradictions inherent in these two examples were the result of a nearly hopeless effort to reconcile a fundamentally materialist philosophy with some remnant of Providence.48 In writing on the relations of God, man, and nature, Geoffroy was breaking the unwritten rules of the scientific community. Like Lamarck before him, Geoffroy, by refusing to act in an appropriate professional manner, invited rejection by his colleagues at the Museum and Academie. They were writing monographs of systematic zoology and comparative anatomy, not, like Geoffroy, forever straying out of their fields of study or continually re-examining the fundamentals of their science. After 1833 Geoffroy ceased to publish in the journals of the Museum and the Memoires of the Academie des Sciences. Between 1832 and 1840, he published only one paper in the Annales des sciences naturelles. That he could continue to publish in the Comptes rendus only substantiates the claim made by the many critics of this series that it was hastily thrown together, with no adequate review, and included many items that were not fit to print.49 It suffices to note the mere titles of two of his last communications to the Academie in 1839, published in the Comptes rendus, to convince one that Geoffroy was lost in a world of his own: "There is Only a Single Physics in the Universe in Which the Worlds Weigh Against Each Other, Communicating by an Immense Molecular Diffusion, Sublime Attenuation of Matter (Imponderous Elastic Gases), and Are Regulated by Means of the Principle (Attraction of Soi pour Soi)." "On a New Argument of Intrastellar Physics, Under This Title:—That There Is in Nature One Substance or Fundamental Essence, LIGHT, Filling All Space in the Universe; Formed of Seven Principles, Susceptible Virtually of Direct and Indefinite Transmission, Transforming Incessantly . . ."50

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As the decade wore on and Geoffroy received less and less encouragement from his colleagues, he came to present himself as a martyr engaged in a providential mission. Believing himself the victim of persecution, he aired at length in print his real and imagined misfortunes. Although he persisted in trying, he unwillingly realized that he would not succeed in convincing the savants of the worth of his ideas and that he would have to look for response elsewhere. By 1837 Geoffroy was showing signs of fatigue and depression, and indulged himself in such self-pitying passages as the following: Posterity, if it deigns to occupy itself with the battles of this age, will have to decide between my adversaries and me. I have a body inclined toward the grave. . . . I have not long to wait.51

In 1838 the professors at the Museum demonstrated their lack of confidence in him by taking control of the menagerie away from him and giving it to his longtime assistant Frederic Cuvier. The loss of the menagerie was a terrible blow to his ego. Encouraged by Geoffroy, the publicist Jean Reynaud wrote a piece on Geoffroy and the founding of the menagerie for a popular journal, Magasin pittoresque, and Geoffroy himself detailed his woes in an essay entitled "Outraged Old Age."52 Adopting the role of an outcast, Geoffroy identified himself with the repudiated natural philosophers of an earlier generation—with Lamarck and especially with Buffon. Both were, like himself, thinkers in the grand style, and their reputations had therefore suffered at the hands of the current generation of academicians. At Lamarck's tomb in 1829, Geoffroy had already acknowledged his colleague to be a kindred spirit, a victim of the same repression that he experienced in his own career: A man of powerful, profound, and most often admirably generalized thoughts, de Lamarck conceived them with a view toward public utility. If he often encountered great resistance, he spoke of it as a condition imposed on every man who begins a reform.53

Geoffroy predicted that posterity would give to Lamarck, whom he described as "one of the greatest zoological talents of our times," his just reward.54 In 1838 Geoffroy once more recalled in print the abuses that Lamarck had suffered "with dolorous patience." "Lamarck lived for a long time poor, blind, and forsaken," he avowed, "but not by me; I loved him and venerated him always."55 Geoffroy's defense of Lamarck, however, was timid and reticent compared with his vigorous efforts on behalf of Buffon. Buffon's reputation had been tarnished in the eyes of Cuvier and many naturalist-academicians because he had catered too much to the popular taste, indulging in speculation instead of adding to the store of so-called positive knowledge. Deeply offended by this verdict, Geoffroy took it upon himself to restore the author of the Histoire naturelle to his former glory. When in 1837 the professors decided that instead of placing Augustin Pajou's statue of Buffon in the new hall of mineralogy, they would order a statue of A.-L. de Jussieu, who had recently died, Geoffroy raised a storm of protest.56 In 1838 he wrote an introduction to a new edition

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of Buffon's works, which he published separately along with a new edition of his eulogies. He dedicated this, his last book, "To the memory of the glorious progress of natural philosophy . . . commencing with the discovery of the astronomical system BY KEPLER AND NEWTON and continuing under the authority of unitary conceptions on the harmonious life of the universe BY BUFFON AND GOETHE."57 In addition, he penned a notice on Buffon for the Encyclopedie nouvelle, an article on his statue in the Comptes rendus, and numerous digressions on Buffon in his other works of natural philosophy.58 Misunderstood for a half-century, Buffon, Geoffroy predicted, would come to be honored as a prophet of the new age. He traced the theory of analogues to Buffon, as well as the doctrine of "mutability of species," and regarded Buffon as the founder of'histoire naturelle generale."59 "His glory, with the aid of the immense knowledge of the nineteenth century, will be to have directed and concentrated the elevated meditations of the current age upon the life and essence of things."60 While Geoffroy never completely renounced his mission to convert the savants of his day, he directed his work more and more to readers outside the Academie. The episode of the orangutan of the menagerie illustrates this tendency to appeal to the public rather than to the academicians as the final arbiter on scientific matters. Geoffroy accused the savants of being motivated by false pride when they separated the orangutan from man, but suggested that the people who visited the menagerie every day, because they were not hampered by the prejudices of the savants, knew better. He declared that having considered the opinions of both the academicians and the populace, he had finally decided to favor the latter: I have for some time wavered and if I now pass into the opposite ranks, it is because I have faith in the solidity of popular judgments. The masses are endowed with an instinctive sense, which renders them shrewd and makes them very skillful at seizing the synthetic point of questions.61

His correspondence, he claimed in Notions synthetiques, had placed him touch with "a nucleus of thinkers who will adopt my generalization of the Newtonian idea of attraction."62 Purposely, he had ceased to write in a technical style directed at "savants de profession," and instead he would reach the savants through "another class of thinkers": For the latter, scientific works only come bristling with technical difficulties, so much so that they neglect them. These men with synthetic views differ from those who pose as the sole and true interpreters of the great phenomena of nature only because, with the titles of philosophers, poets, and theologians, they study differently, in a more general manner, the rules, the effects, all the phenomenal manifestations of the universe.63

Geoffroy deliberately sought out "men with synthetic views" and gave them credit for their support in his works. He sent them his books and read theirs, and in imitation of Cuvier instituted a salon at his home in the Jardin des Plantes. It was in response to Geoffroy's active encouragement, then, that critics of science and the scientific establishment came to write about Geoffroy and the debate.

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Until 1840, when he became too ill to attend the meetings of the Academie, Geoffroy persisted in the promotion of his scientific and public mission. In 1841 he went blind, and unable to continue teaching, he resigned his chair at the Museum in favor of his son. He would gladly have left his chair at the Faculte des Sciences to Isidore as well, but was prevented from resigning by the dean, Dumas, who avowed that Geoffroy was too important a name for the Faculte to lose. But while Dumas honestly admired Geoffroy as the founder of philosophical anatomy, perhaps the more pressing reason for his refusal to let Geoffroy go was that he wished to reserve the chair for the man who eventually received it in 1844, his longtime friend and co-editor of the Annales des sciences naturelles, Milne Edwards.64 Removed from active participation in scientific affairs for three years, Geoffroy expired peacefully on 19 June 1844 at the age of 72. He died a popular hero. Nearly two thousand people attended the ceremony at Pere Lachaise cemetery: most of the leaders of science, physicians, artistic and literary figures including Victor Hugo, Jean Reynaud, David d'Angers, and Edgar Quinet, employees of the Jardin, and other "persons of all classes of society."65 The Debate and the Moral and Social Implications of Ideas

Literary authors and publicists, when they came to write about the controversy between Cuvier and Geoffroy, interpreted it in terms of Geoffroy's writings of the 1830s. They focused not on the anatomical arguments, but on the metaphysical assumptions behind the anatomical disagreement. They were unconcerned with transmutation versus fixity of species and did not understand the complexities of morphology versus teleology. Geoffroy's doctrine of unity of composition mattered only insofar as it called for a progressive series of links between animals rather than a series of arbitrary gaps. Style and world-view, rather than science, were primarily at stake. Cuvier represented a narrow enterprise of fact-gathering within the confines of the traditional world-view, while Geoffroy represented a bold and intuitive attempt to create a new and more harmonious science with greater promise for mankind. The traditional static world assumed by Cuvier seemed to critics to place restrictions on the questions man could ask of nature. Many features of the organic world could only be attributed to God's seemingly arbitary choice, rather than to universal laws that might be rationally sought out. Geoffroy's doctrine, as expressed in the 1830s, offered instead a lawful and progressive world, one that seemed to grant man the right to investigate nature to the limit of his capacities. It was this optimistic view of man's ability to comprehend the universe in Geoffroy's work that appealed to writers of the 1830s and 1840s such as Sand, Balzac, and Quinet. Whereas the savants, even his own son, rejected Geoffroy's soi pour soi as too speculative and metaphysical, the romantic novelist George Sand, just beginning her career, was sufficiently impressed by the grandeur of this uni-

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versal law that she tried to incorporate a discussion of its momentous consequences into one of her novels.66 Geoffroy, no doubt having heard of Sand's interest, deliberately took the initiative to enter into correspondence with her. From 5 June 1835, the date of Geoffroy's first letter, they carried on a correspondence of mutual admiration. She placed one of his letters in her famous "hieroglyphic chimney," and he, in turn, had a bust of her made.67 When the novelist finally met the old naturalist face-to-face in the menagerie of the Jardin des Plantes in 1836, she thought him a trifle odd, but nonetheless a great man: The old Geoffroy is for his part a rather curious beast, as ugly as the orangutan, as talkative as a magpie, but for all that full of genius.68

Geoffroy sent Sand copies of his works, notably his article "Pantheistic Heresies," and parts of his as yet unpublished Notions synthetiques, and begged her to employ her pen in order to disseminate and defend his scientific theories. She protested that she was not capable of doing justice to the subject: "I would surely write a laughable work if I tried to explain how all that arranges itself in my head; thus I will not undertake it." Yet she assured Geoffroy that he was "on the path of the prophets of truth" and that the doctrine espoused in Notions synthetiques was "broad and magnificent" and "throws to the ground the Genesis of Cuvier [Cuvier's view of creation] for anyone who detests meanness in the arts." She informed Geoffroy that she had already attempted unsuccessfully to express her thoughts on his philosophy of nature. These pages, intended for a new edition of her novel Lelia, she sent to Geoffroy. In the future, she told him, she hoped that she might write something on behalf of his cause, but despite Geoffroy's repeated solicitations, she never published anything on the matter.69 The fragment intended for Lelia, entitled "Soi pour soi, la science," suggests that Geoffroy's appeal rested not upon his science but upon his romantic world-view. Believing the traditional God to be a violent, hateful, and vindictive Deity, who created the world at His caprice, Sand sought a more satisfying philosophy that would give more scope to man's rational mind. Although nature, in Cuvier's system, had been providentially arranged for man, man was forbidden by his religion to question nature's mysteries. "Never could the Creation be interrogated by this stupid master whose proud ignorance was his religion, his law, and his virtue."70 Cuvier's world-view, as interpreted by Sand, thus left incomprehensible gaps of knowledge. Nature was left fragmented: He [Cuvier] seems to break the unity of the universal system and place inexplicable lacunae in the thought of God. To say that man ought to halt before certain mysteries, deny his intelligence before the caprice of a coarse and jealous God of centuries past, that is what no one can explain or wishes to explain on the part of the great Cuvier.71

In the doctrine of Geoffroy, "venerable worshiper of the universal plasticity of God," she found, on the contrary, continuity, order, and harmony. She went so far as to read into Geoffroy the chain of being reaching from the stone to

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God. Despite the imperfections of Geoffroy's style, his work revealed a grand vision: A style that is hardy, strange, harsh, obscure, thoughts as vast as the world, formulated with a profound scorn for form, and [yet], in the midst of his vagueness of expression, of his ignorance of and disdain for words and phrases, the transports of biblical thought, the cries of the mysterious and grandiose soul, as one finds only at the dawn of religions: this is what I have found, as poet, in this brief formula of a universal system. But as man. . . I have found in this new perception of Creation that which is most worthy of faith, that which is most satisfying to the human spirit, and that which is the most conformable to man's noble instincts and to his inexhaustible thirst for order and harmony: the universal unbroken chain, the balanced and harmonious joining by innumerable links and insensible gradations of inert nature to animated nature, of the stone to the plant, of the insect to the bird, of the brute to man, of man to the all, and of the all to God.72

Sand predicted that the future would occupy itself more and more with the debate of the two greatest naturalists of the age. The majority, she recognized, had, under the sway of Cuvier's reputation, concluded that Cuvier "nearly silenced for a time the voice of his rival." But Geoffroy, who would "pursue until his death" the development of his ingenious perceptions, was among those "whom today's generations do not understand, but whom tomorrow's will rehabilitate."73 Balzac's preference for Geoffroy was based on many of the same motives. Geoffroy's relations with the great French novelist were also, it appears, initiated by the naturalist. The earliest letters are lost, but by 1835 Balzac was already in correspondence with both Geoffroy and his son, exchanging books and expressions of admiration.74 In 1831 Balzac had written in praise of Cuvier in his novel Le peau de chagrin. Balzac's enthusiasm for Cuvier was awakened by the contemplation of the vast stretches of time revealed by Cuvier's paleontological discoveries. Ironically, he compared Cuvier, the promoter of a science of facts, to a poet. Is not Cuvier the greatest poet of our century? Lord Byron reproduces moral throes in verse, but our immortal naturalist has reconstructed worlds from a whitened bone; rebuilt like Cadmus, cities from a tooth. . . Cuvier is a poet by mere numbers. He stirs the void with no artificially magic utterance; he scoops out a fragment of gypsum, discovers a print-mark and cries out "Behold!"—and lo, the trees are animalized, death becomes life, the world unfolds.75

As he read more in the sciences and began to develop his own philosophy, influenced by Gall, Lavater, and Swedenborg, the implications of Cuvier's "doctrine of fatalism" became evident to him. A proclaimed Catholic and monarchist, Balzac was nonetheless, like Sand, irresistibly drawn toward Geoffroy's philosophy by its romantic, pantheistic, and unified vision of the world. In Balzac's case, the social dimension of these philosophical ideas was made explicit. From Livre mystique, written in 1832, and containing the novels Louis Lambert and Seraphita, emerged Balzac's own unitary philosophical system. Bal-

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zac, like Geoffroy, rejected the separation of science from philosophy. The purpose of science was not just to accumulate facts about nature, but to reveal the relations of man to God and society. Balzac took his effort to reform philosophy seriously and hoped the scientists would benefit from it. He wrote to his sister: Louis Lambert has cost me so many labors. I was obliged to reread so many works to write this book. Perhaps it will lead science into new paths some day. If I had written a work purely for savants, it would have attracted the attention of thinkers who will not glance at it. But, if chance places it in their hands, they will perhaps speak of it!76

In the novel, Louis Lambert explained to his fellow student, Dr. Meyraux (the Meyranx of the debate), that as a philosopher, he was studying comparative anatomy because of its importance in clarifying man's relations with God: Is not this a need of the age? Without the highest assurance, it is impossible to put bit and bridle on the social factions that have been let loose by the spirit of scepticism and discussion, and which are now crying aloud: 'Show us a way in which we may walk and find no pitfalls in our way!' You will wonder what comparative anatomy has to do with a question of such importance to the future of society. Must we not attain to the conviction that man is the end of all earthly means before we ask whether he too is not the means to some end? If man is bound up with everything, is there not something above him with which he is again bound up? If his is the end-all of the unexplained transmutations that lead up to him, must he not be also the link between the visible and invisible creations? The activity of the universe is not absurd; it must tend to an end, and that end is surely not a social body constituted as ours is!77

Through analyzing the similarities among animals—"the animal type as a whole"—Lambert expected that "the secret of the various moral zones through which man passes will be discovered." "In these days," the hero asserted, "science is one; it is impossible to touch politics independent of moral questions, and these are bound up with scientific questions."78 Lambert lamented that the Institute, which "might be the central government of the moral and intellectual world," had been ruined by its subdivision into separate academies.79 When Balzac sent Livre mystique to Geoffroy in 1835, Geoffroy immediately recognized its relevance to his ongoing battle against scientific specialization. One reason for shelving Notions synthetiques, written in 1835 and not published until 1838, was, he wrote to Balzac, in order to incorporate Louis Lambert into it. From Balzac, Geoffroy derived the motto of his new book, "Science is one and you have divided it."80 In the "Avant-propos" of La comedie humaine (1842), Balzac's most widely read discussion of the debate, Balzac explicitly modeled his conception of "social species" on Geoffroy's doctrine of unity of composition. A grand scheme embracing all segments of French society in the nineteenth century, La comedie humaine provided Balzac with a superstructure to almost all of his literary works. Balzac declared that Geoffroy had triumphed over Cuvier in the debate of 1830. Unity of composition, according to Balzac, was not "a sci-

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entific innovation," but had occupied the minds of "mystical writers" such as Swedenborg and the "finest geniuses of natural history" such as Leibniz, Buffon, and Bonnet for the preceding two centuries. In the writings of all these men one could find "the rudiments of the beautiful law of soi pour soi" on which "unity of composition" depends: There is only one animal. The Creator has used only a single pattern for all organized beings. The animal is a principle which takes its external form, or, to be more exact, the differences in its form, from the milieus in which it is obliged to develop. Zoological species are the result of these differences. The proclamation and defense of this system, which is, moreover, in harmony with our ideas of divine power, will be the eternal glory of Geoffroy Saint-Hilaire, the victor over Cuvier in this point of higher science, and whose triumph has been hailed by the last article written by the great Goethe.81

In La comedie humaine, Balzac formulated a parallel between the notion of zoological species and that of "social species." As there was in essence only a single animal, there was likewise but a single form of humanity. In both the zoological and the social world, differences between species could be best explained by the action of the environment. Pere Goriot, the work that Balzac dedicated in its second edition of 1842 to Geoffroy, was one of Balzac's best known studies of the effect of milieu on the development of character. Both Sand and Balzac preferred Geoffroy's pantheism, mysticism, and breadth of vision to the narrow, analytic science of his rival Cuvier. Indeed, Sand and Balzac saw in Geoffroy more of a philosopher and mystic than he really was. Geoffroy's science was, to Balzac, superior to that of Cuvier because it was not amoral and objective, but instead lent support to the notion of the progressive nature of humanity. Geoffroy himself had shown only a limited concern for the moral and social implications of his works, but he was flattered to have others use his ideas toward that end. The moral significance of Geoffroy's work was also the subject of the funeral eulogy by the celebrated French historian Edgar Quinet. A romantic, republican, and ardent anti-Catholic, Quinet taught at the College de France in the years of republican agitation preceding the revolution of 1848. He and his friend Jules Michelet, also a professor at the College de France and a partisan of Geoffroy,82 promoted the ideals of liberty, democracy, and nationalism. In 1843 they collaborated on Des jesuites, an inflammatory history of the Jesuit order. As a concession to Church power, both were removed from their posts before 1848. In his interpretation of Geoffroy, Quinet naturally saw him in terms of his own battles against king and Church. When a savant died in nineteenth-century France, it was customary for a representative of each institution of which the deceased was a member to give a funeral oration. Thus at Geoffroy's funeral on 22 June 1844, Dumeril spoke on behalf of the section of anatomy and zoology of the Academie, Michel Eugene Chevreul for the Museum, Dumas for the Faculte des Sciences, and Pariset for the Academie de Medecine. Following these speeches, all but that of Dumas polite rather than enthusiastic, came three more from devoted friends: Serres, the octogenarian Lakanal, and Quinet. In his eloquent address,

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the last to be presented, Quinet expressed the attraction and relevance of Geoffroy's thought for him. Having a firm belief in the power of ideas, especially religious ideas, to mold society, Quinet saw Geoffroy's work as essentially religious: as an attempt to find the harmony in nature which presaged the effort to establish harmony in the social order. He compared Geoffroy to an Archimedes removed from all earthly care, serenely seeking out the mysteries of creation. Unknowing, Geoffroy had produced a science of great import to man in society: How does it happen that with so little love of reputation and fame, this man, wholly withdrawn into science, should become popular among us? It is because the idea to which he has given birth is, in many respects, at the very heart of our time. The desire, the presentiment, the necessity for a vast unity—that is what the world is working toward. M. Geoffroy Saint-Hilaire, a true prophetic genius, has established in nature and in science that harmonious principle that we still seek in the civil, political, and religious world. That is how the works of this creative mind are linked to the current work of the whole human race, and since he first arrived at this foundation of unity that everyone is seeking for in every manner, he has, without knowing it, caused the world to concern itself with his glory.83

Quinet linked Geoffroy with Naturphilosophie and recalled that "the greatest writer of Germany, Goethe" undertook to popularize Geoffroy's victory in France. Geoffroy had been accused of being a poet, admitted Quinet, and he was indeed one, like all great men who are able to intuit the truth. Concluding his eulogy on a lofty moral note, Quinet exclaimed: "Aid me with your light and your virtue! The best thing of my life will always have been to have obtained your friendship."84 The Debate and the Politics of Science

Philosophical interest in Geoffroy and the debate cannot be easily separated from political interest. Those who admired Geoffroy for his breadth of vision were often the same people who railed against official science for turning the study of nature into the gathering of details irrelevant to larger human concerns. As representatives of those who referred to the debate in the context of a critique of official science, we include in this section a Balzac short story satirizing the debate, Raspail's angry assault on scientific coteries, and an obscure satire of the professors of the Museum, Histoire naturelle drolatique, A common picture emerges from these works. True science, these critics presupposed, asked meaningful questions about man's relation to God and nature. But because of the structure of modern science in France, creative philosopher-scientists were ignored while mediocrities were showered with benefits at public expense. The Academie had become filled with narrow, analytic drudges who knew nothing of real philosophy but only how to play the game of rewards and sinecures. In such a system, science was no longer directed to the people but was clothed in obscure language understood only by specialists. Within this exaggerated picture of the French scientific establishment, Cuvier

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was stereotyped as the conniving villain, and Geoffroy as the philosopher par excellence and man of the people. The debate as an illustration of unsavory politics in action is especially well depicted in a little-known satire by Balzac. Entitled "The Ass's Guide for the Use of Animals Who Wish to Achieve Honors," it appeared in 1842 in a collection of animal tales edited by Pierre-Jules Hetzel and illustrated by the talented engraver J.-J. Grandville.85 Its narrator was an ass, whose owner, one Adam Marmus, was an impoverished primary school instructor residing in the suburbs of Paris. Marmus was struck by the reflection that, while he struggled to teach children to speak, read, and write, his ass could quickly teach the little asses all they needed to know in life. The idea of a new science—the comparative study of instincts—came to his mind. Although ignorant of natural history, Marmus resolved to go to Paris to request a position from the minister of public instruction in order to pursue his discovery. In the capital, Marmus chanced to meet an equally impoverished journalist who offered to act as his public relations man. He convinced Marmus that real status was irrelevant so long as one had the appearance of somebody important, and he held out the appealing prospect: "You have invented, my dear man, instinctology, and you will have a chair of Comparative Instincts."86 Together they concocted a scheme. The ass would be painted so as to resemble an extraordinary zebra, given a cow's tail, and taught to walk like a giraffe. Marmus would be billed as an intrepid explorer recently returned from the Mountains of the Moon in Africa where he had discovered an amazing beast (the ass) which destroyed all the classificatory categories of the celebrated Baron Cerceau (readily identifiable as Cuvier). Both the new species and the new science were to be offered as proof of the unitary theory of Cerceau's rival, "the great philosopher" (Geoffroy). They would argue that just as there was in essence only a single animal, there was only one instinct, modified according to the environment. The motive behind all this trumpery was to force concessions from the powerful baron: We will intervene by a new science against the Baron Cerceau in favor of the great naturalist-philosophers who believe in zoological unity, and we will obtain from the all-powerful baron good conditions for selling him our science.87 The savants live by nomenclature; let us turn the nomenclature upside down. They will become alarmed, they will come to terms, they will seduce us, and like so many others, we will let them seduce us.88

Their plot to sell out to the baron succeeded royally. Fearful of the consequences of the new science, the savants sent to Marmus an academician (illustrated as an ass in the costume of the Institut) who did not hide his concern for saving "the fatalist doctrine of the Baron Cerceau." The resources at Cerceau's command were impressive: "Behind Cerceau there were ranged the academicians, the University, the legions of professors, and the government supporting a theory presented as the only one in harmony with the Bible."89 The great philosopher's system had to be nipped in the bud, "because, after all, . . . it is pantheism." With the assistance of a disciple of the great philosopher,

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Marmus wrote a notice of his discovery and its importance for the unitary system. At the Academie the affair soon became so grave "that no academician dared to give an opinion" on it. Finally, the minister of public instruction succeeded in striking a bargain with Marmus by offering him a sum of money from the funds allotted for the encouragement of science. In return, Cerceau took control of the new science of instinctology and had one of his henchmen, represented in the illustration by a parrot, rework the field in accordance with his views. The baron's disciple, who took over Marmus's proposed course at the Athenee, taught that instincts corresponded to the classificatory categories of the baron. He "played variations on instincts exactly as Paganini played variations on the fourth string of his violin." The "zebra" was treated as "a monstrous exception." Named an assistant librarian at the Jardin des Plantes, the journalist decried the great philosopher in the periodicals. "He was regarded as a dreamer, as an enemy of savants, as a dangerous pantheist, and his doctrine was made fun of."90 Cerceau had Marmus named "professor of something somewhere," but his legislative duties made it necessary to delegate his teaching to a suppleant, the journalist turned librarian, who paid a man of talent 20 francs from time to time to prepare the lectures. Showered with honors, master and ass lived happily ever after. As for the great philosopher, he was powerless to prevent this treason, for he had "neither places to give, nor any government for him but the government of science, at the head of which Germany has placed him." The moral of the ass's tale was that there exists in the budget a powerful contribution paid by imbeciles to intriguers, that every chair can be pimped [une marmite], that the public is a vegetable, that he who knows to be silent is more clever than he who speaks, that a professor is named less for what he says than for what he does not say, and that it is less a matter of knowing than having. My former master has placed his entire family under the shelter of the budget. . . . Dare say that I am an ass, I who have given you here the method to succeed and the epitome of all the sciences.91

In this exaggerated and cynical story, the truth or falsity of scientific theories was irrelevant in the choice between Cuvier and Geoffroy. Cuvier and his disciples prevailed because they were allied with conservative forces in society and because they controlled the political power to make their ideas accepted. Others with opposing points of view either were converted because it was to their career advantage to be converted or were rendered ineffective. An even more trenchant account of the debate as political intrigue appeared in the polemical writings of the scientist and social reformer Francois-Vincent Raspail. Born into a poor family in 1794, Raspail received a clerical education. After the Restoration he came to Paris, an avowed Bonapartist and republican, and soon an anti-Catholic as well. No mere theoretical democrat, Raspail had fought at the barricades in 1830, and for his subversive activities spent several years in jail. Opposed to authoritarianism of any kind, whether in government or in science, Raspail was less concerned with the intellectual than with the political dimension of the Cuvier-Geoffroy debate.92 Raspail at one time aspired to a scientific career but soon ran afoul of the

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academicians. In the 1820s he taught himself science and medicine, but did not take a medical degree, nor, as far as the records go, any formal courses. Choosing for his subject of investigation the grasses, in part because they were the proletariat of the plant kingdom, he submitted his first paper on their embryology to the Academie in 1824. In retrospect, Raspail's research has been judged as thoroughly competent and highly prophetic; he is now heralded as a precursor of the cell theory, a first-rate microscopist, and the founder of histochemistry. That his work did not find favor among the academicians was due less to its controversial nature or to dirty politics than to Raspail's abrasive personality.93 In the 1830s Raspail recalled that the only academician who appreciated his first paper was Geoffroy, who supposedly came up to him afterward and said, "Don't be discouraged, young man, you are twenty years ahead of them."94 Nonetheless, the memoir was successful enough to be printed in the Annales des sciences naturelles. Raspail presented other papers at the Societe d'Histoire Naturelle which were commented upon favorably by Biot. But in 1827 a quarrel with the botanist Adolphe Brongniart, one of the editors of the Annales des sciences naturelles, put an end to cooperation between Raspail and the academicians. It did not help that Raspail accused Adolphe's father, Alexandre Brongniart, of refusing him entry into a public meeting of the Academie. The elder Brongniart, who claimed that Raspail had tried to enter in the middle of a paper, implored Raspail in a letter to "discuss scientific matters graciously;. . . scientists wish to be approached in a considerate manner. They have the right to expect it."95 From then on, except for Geoffroy, he was generally hostile to the academicians, and especially to the Brongniart clan.96 Raspail became convinced that the affairs of science were nothing but the machinations of elite cliques. In 1829, with his friend J.-F. Saigey, Raspail launched his own scientific journal, Annales des sciences d'observation, as a vehicle for a continued assault on official science.97 In a piece entitled "Scientific Coteries," he lashed out against the repression of innovators by leading factions. Though not mentioned by name, he no doubt had in mind men like Brongniart and Cuvier: Places, favors, competitions, eulogies in scientific journals, all is in the end at the disposal of the privileged coterie, and misfortune to any recalcitrant head! His very talents will only hasten his fall and make it harder to bear.98

He contrasted the fate of a young savant willing to court the favor of the leading faction with that of one who will not. If the young man "attaches his glory to the bandwagon of the intrigue in favor," he will perforce limit his evidence, his intelligence, and the subjects he can treat. If, however, he tries to remain independent of the factions, he will be opposed at every turn: First, he will not be attacked to his face, but he will be ridiculed in secret. His discoveries will be bantered about in soirees, banquets, and concerts. . . The scientific or literary journals that might be at his disposal will be prohibited from occupying themselves with the work of a recalcitrant, of this obscure innovator! Attacked from all sides, he will be cited by no one.99

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Let the innovator persist, continued Raspail, and he will then be attacked head-on, his work destroyed through sarcasm, gross omissions, and mutilated quotations. The Cuvier-Geoffroy debate furnished Raspail with tailor-made ammunition for his critique of the scientific establishment. In an article on the debate entitled "Theory of Analogues, and Discussion between MM. Cuvier and Geoffroy" for his 1831 polemical work Nouveaux coups de fouet scientifique [New Scientific Lashings], he interpreted the debate as one more example of academic "intrigue." According to Raspail, Cuvier entered into the debate for political reasons, because he perceived the partiality of young French savants toward Geoffroy and could not bear such "scientific defection."100 This time Cuvier received more resistance than he expected to his attempt to score an easy victory over Geoffroy. Although Raspail found Geoffroy guilty of uncritical thinking and strategic blunders, Cuvier was far more to blame for resorting to fallacious arguments, ridicule, and manipulation of his audience. Moreover, Cuvier had told his protege Alfred Donne what to write in the reports in the Journal des debats. Raspail admitted that Meyranx and Laurencet's comparison of cephalopods and vertebrates would not hold (as Cuvier showed), but he was certain that the presence of so many common organs in mollusks and vertebrates attested to real homologies. He hailed the theory of analogues as "a generality which, by its simplicity of expression and the extent of its application, threatens to engulf as a grand law all the pompously fastidious apparatus of a nomenclature so easy to create and so troublesome to learn."101 Like most of the critics of official science, Raspail pictured Cuvier as an uninventive fact-monger, who, he claimed, collected 45,000 francs a year in sinecures, and whose vaunted genius consisted solely of a prodigious memory and clarity of speech: But among us savants, let us avow it. Has M. Cuvier really advanced science a single step?. . . What has M. Cuvier produced? Descriptions and nothing but descriptions. But who does not know how to describe nowadays? Who does not know how to count fins and fish bones and designate their colors? The men of the world will perhaps cite to me his Theory of the Globe. According to the journals of the time, M. Cuvier seemed to have been present at the work of the Creation. Well! This theory, it must be said, was nothing but a gross irrelevance, and as such it has fallen so that nothing more remains of it. Poor Frenchmen, how intrigue snares you! You pale before these geniuses created by complacency and sanctioned by servility. But if, in a more liberal system of education, the monopoly of sinecures is destroyed, and all those called to science whom nature has fitted for intellectual work, and the cabinets which are now open only for the curators are no longer closed, I will bet that out of a hundred students, in three years 102 of study, there will be found ninety-nine Cuviers and one genius of the first rank.

The debate to Raspail was thus an unequal contest between a powerful member of a ruling faction and an imaginative innovator who did not have the same resources at his disposal.103 Despite Raspail's acerbic personality and near paranoia, Geoffroy was ready to encourage him and in 1833 he began a correspondence. As president of the Academie in 1833, Geoffroy, inflated with his temporary importance, believed

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that he could help Raspail get his due recognition by having the Montyon prize in experimental physiology awarded to him. He told Raspail that he thought he could convince his colleagues, if only Raspail could be persuaded to cooperate.104 Geoffroy was then called in by the minister of public instruction, and, in one version of the story, was warned, "I forbid you to line the rioters' pockets."105 It is unlikely that by 1833 the academicians would have voted for Raspail in any case. At this time, Raspail's New System of Organic Chemistry (1833) had just reached Geoffroy, who worried lest the reference to nepotism in Raspail's inflammatory introduction might be construed as applying to his own son, recently made a member of the Academie.106 Raspail assured Geoffroy that had he known of Isidore Geoffroy's election, he would have rephrased his insinuation (which was obviously directed at the Brongniarts). He wrote to Geoffroy on 9 July 1833: I respect the affections of a father, but I will never respect the machinations of an intriguer. But you, monsieur, have not gained your stripes by intrigues. In truth, who would not blush that the sons and relatives of M. B—are to be found in all vacant places, not by right of conquest, but by right of birth.107

Raspail intimated that Isidore Geoffroy would have done better had he refused the seat and turned his back on the Academie. Geoffroy himself was only a partial opponent of the institutions of science—he intrigued for positions with the rest of the academicians and certainly did not disapprove of nepotism when his son was concerned—yet he naively but deliberately associated himself with an angry dissident like Raspail. It was because of such actions that Geoffroy was depicted by Raspail and other critics as more of a rebel against the system than in fact he was. A more clever and somewhat less vitriolic assessment of Cuvier and Geoffroy appeared in 1847 in a small volume with the strange title Histoire naturelle drolatique et philosophique des professeurs du Jardin des Plantes. Both Isidore de Gosse, the author of the work, and Frederic Gerard, who wrote the introduction and several of the articles, were, like Raspail, outsiders exasperated by the control of scientific thought by a centralized establishment. De Gosse was the pseudonym for Bertrand-Isidore de Salles, a literary and scientific journalist who later became an administrator in his prefect. Gerard, an unsuccessful naturalist turned popularizer, was one of the few French advocates of Lamarck and evolution in the period before Darwin.108 After the manner of a natural history text, Histoire naturelle drolatique undertook the history of the professors of the Museum and their aides, assigning to each a Latin binomial name and a characteristic description. The resulting menagerie was presented in a series of hilarious "episodes scientifiques et pittoresques" which are extraordinarily valuable for the insights they offer into the popular perception of official science. The main theme of the book was that science and the scientific community had declined from their former eminence. Where once there had been great inventive geniuses, now there were only mediocrities who hid behind jargon

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and their sinecures. Concern for broad and relevant questions had been abandoned as science had become fragmented into specialties, each with its own specialist language. The centralized institutional system promoted nonentities rather than true savants, for access to a professorship was controlled by factions whose leaders were concerned primarily with patronage and money rather than with the discovery of truth. As in the case of Raspail, the main offender in this regard was the nepotistic Alexandre Brongniart (given the name Porcelainianus imperiosus) and his clan.109 Cuvier was depicted as the leading force behind this retrograde movement of science. For the Jardin des Plantes, wrote Gerard, he was the robe of Nessus. Given the Linnaean name "Analyticus diplomaticus, Lacepede," he was satirically described as follows: The leanings of his Protestant education inspired in him an aversion to encyclopedic philosophy. From his entry into the temple of science, he did not understand all the splendor of the edifice. He saw only lines to reproduce, capitals to sketch, an architectural arrangement to describe, without seeing that from all of it there was a general idea to deduce. He undermined the broad views that presided in education and substituted the cold data of his Tableau du regne animal. More courtier than naturalist, he elevated himself to the level of the great by his submission to the will of successive powers, and consecrated in science a deplorable error: that natural history is a science of details, and that the method is the first and last goal of the naturalist. He has done nothing for the success of studies proper to emancipate the mind, and he has prevented universal men who have understood science differently than he from letting their doctrines predominate.110

Cuvier was thus despised both as a repressor of what was most meaningful in science and as a flatterer to men in high places. As in other assaults on scientific elitism, Geoffroy emerged as a scientist of the people, one of a handful of recent scientists who appreciated the true importance of the investigation of nature. He was portrayed as the last in a long line of true savants. The authors of Histoire naturelle drolatique had a special interest in Geoffroy. Gerard had prepared a forthcoming book on him entitled "Geoffroy Saint-Hilaire and His Era: History of the School of Natural Philosophy." The work was advertised, but there is no indication that it ever appeared.111 Named "Transcendentalus honestus, Serres," he was lauded as follows: Geoffroy is no longer, and all eulogy rendered to his memory is no longer flattery but a just homage. Good, honest, an enthusiast of elevated and profound thoughts to whose research he devoted his life, Geoffroy, man of heart and full of sensibility, understood Lamarck and loved him. To love one who is abandoned would already be a noble action, if to have opened to science a path that has in vain been denied, had not been a sublime task sanctioned by 50 years of assiduous work, and had not acquired for him the right to the respect of posterity.112

Other heroes among the professor-naturalists were Lamarck ("Philosophus clarissimus, Buffon"), Lacepede ("Ichthyologus affabilis"), Serres ("Anatomicus philosophus, Geoffroy") and Isidore Geoffroy ("Teratologus, Serres").113 Contrasting Geoffroy to the mediocre talent of the day, one skit satirized the sorry battle for Geoffroy's seat at the Academie. The sun, pale and weeping at

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the news of Geoffroy's death, called the planets together and said, "The Earth, my daughter, has lost an estimable man . . . and who will replace him?" However, the planets Vesta and Juno (representing Arago and Brongniart, the leaders of factions at the Academie) could think of nothing but promoting the causes of their respective candidates, the two nonentities Valenciennes and Duvernoy. Valenciennes, previously described as one who "believes that it suffices to know how to distinguish a carp from a pike to be a great man,"114 was awarded the seat of the great Geoffroy, and for his reception speech prepared a discourse on "comparative fricassee of rabbit." The sun groaned and at last consoled himself with the thought that academicians were scarcely worth the bother.115 In an especially noteworthy skit that satirized the introduction of esoteric terminology into natural history, Geoffroy appeared as the scientist of the common people. The skit poked fun at Blainville, who in his effort to overthrow the classification of Cuvier, had created a new and complicated terminology. In the story, entitled "A Conjuration," Blainville was made to say: The common people in all these parts are invading the sanctuary of science. The language that we speak is no longer unknown to them, and already there are among this mob men who are judging our writings, picking at our style, and pronouncing on the merit of each and every one of us. However, what would happen if we placed between the people and the savant a triple barrier so that indiscreet eyes could no longer penetrate the mysteries of the temple?116

To Valenciennes's inquiry of what remedy might be found for such an evil, Blainville suggested that naturalists create for each specialty a language full of mysterious terms derived from Greek and Latin "which strike with stupor the ignorant herd and impart to our persons a sacred character.117 All of the professors were enthusiastic, except for Geoffroy who alone defended the public interest: I protest that a proposition as odious as this should be executed. We are here to teach the people in the language that they speak those things that can be of use to them. We should not forget the goal for which this establishment has been instituted, and it would be lacking in honor, duty, and reason to follow a path that would render us all contemptible and ridiculous. Science gains in grandeur when it is popularized. . . . Let us say simply in French that which we know, and let us not hide our ignorance. A barbarous language is nothing but a convenient veil for mediocrity.118

Needless to say, Geoffroy's plea fell on deaf ears and the "conjurers" in the end agreed to the introduction of Blainville's Monodelphia, Didelphia, Pilifera, Celerigrada, Gravigrada, etc.119 Indeed, Blainville deserved his reputation for elitism, for he firmly believed that professional naturalists should be judged only by their peers. The portrayal of Geoffroy as a savant concerned for the welfare of the people, and even of the animals in the menagerie, was surely overstated. In the final episode of the volume, entitled "Morality," a young woman symbolizing Truth told two young men that she would reveal who the true savants really were. All the professors at the Museum were taken into a dark room

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where an immense globe of gold-beater's skin was hung from the ceiling by a thread. On the globe were written the words "Modern Science." The professors hastened to inscribe their names, a difficult process because the globe was incessantly revolving. Finally La Verite became disgusted and punctured the balloon. She lit the hall with her mirror, reversed the busts decorating the room, and produced a great book bearing the title "Philosophy." In it were inscribed the names of Bernard de Palissy, Rabelais, Montaigne, Newton, Franklin, Buffon, La Mettrie, Erasmus Darwin, Linnaeus, Adanson, A.-L. de Jussieu, Laplace, Lamarck, and Geoffroy.120 The men and women cited in this chapter are only a sample of those who became ardent defenders of Geoffroy in the 1830s and 1840s. Much more might be uncovered concerning the many obscure individuals who are mentioned in Geoffroy's works of natural philosophy or who attended his salons. Those discussed here do reveal, however, some of the main reasons why in the 1830s and 1840s Geoffroy was thought to have triumphed over Cuvier. His defenders, taking their image of Geoffroy from his writings of the 1830s, were attracted to his vision of a universe continuously and progressively evolving according to the interplay of universal laws. They appreciated the broad scope of his science and the boldness of his ideas. They admired his taking an independent stand against his fellow scientists. And they sympathized with him as the underdog in an unequal contest against a powerful and well-connected rival. Yet for the most part Geoffroy's advocates among the nonacademicians had only a superficial grasp of his scientific doctrine. They understood few of the fine points of anatomy brought out in the debate, and they had no scientific experience on which to base an informed and independent judgment. Instead they saw only what they wished to see in Geoffroy. They made him into a mystic, a pantheist, and an anti-Catholic. They portrayed him as an active rebel against official science and its institutions. They hailed him as a scientist of the masses. Some saw him as removed from all politics, an Archimedes concerned only with the advance of pure knowledge, while others depicted him as motivated primarily by the moral and social implications of his doctrine. The image became more important than the reality. Geoffroy and his works, as inevitably happens with all far-thinking scientists, were distorted and exaggerated, and exploited for ends that Geoffroy could not have foreseen. For the leaders of the generation of naturalists after 1830, who understood Geoffroy even if they did not agree with him, the debate posed a different set of problems. Geoffroy's doctrine could not be accepted as a whole, and yet he had raised too many important issues to be ignored. How the major naturalists in France and Britain came to terms with philosophical anatomy and the debate is the subject of the final chapter of this study.

CHAPTER 8

Teleology, Morphology, and Evolution: The Debate and the Future of Zoology

"The world was divided," exclaimed Flourens of the Cuvier-Geoffroy debate in his 1852 eloge of Geoffroy for the Academie: The austere and regulated thinkers, those who are most concerned with the severe and precise course of the sciences rather than with their rapid bursts, took the part of M. Cuvier. The bold minds ranged themselves on the side of M. Geoffroy. From the depths of Germany the old Goethe applauded his arguments.1

In his 1847 biography of his father, Isidore Geoffroy wrote of the inevitable confrontation between two great schools—the school of facts and the school of ideas—and of "the celebrated discussion, which, in 1830, was to resound in all of learned Europe."2 To read these two accounts, one by a disciple of Cuvier and the other by a disciple of Geoffroy, one might conclude that French zoologists after 1830 were long divided into two sharply opposed camps. It might well be supposed that the diametrically opposite positions articulated during the confrontation of 1830 might have set the stage for a continued battle of disciples. Far from being split apart by the debate, however, French naturalists in the 1830s and 1840s reached an extraordinary degree of unanimity on the issues brought forth by the controversy. The confrontation of two opposing views in 1830 served not so much to divide naturalists into two camps, as to bring to a focus a fundamental problem in biology, and to emphasize the need for a resolution. To be sure, Cuvier had his loyal disciples—Duvernoy, Frederic Cuvier, Dumeril and Valenciennes—who attained high positions but did not deviate substantially from Cuvier's theoretical views. Likewise, Geoffroy could boast of two successful followers, his son and Serres, who continued to promote the doctrines of unity of composition and arrests of development. But even the closest of disciples modified their stance in the light of the debate. This was so in the case of Duvernoy, who, unlike his mentor, willingly recognized the significance and originality of philosophical anatomy.3 And this was especially true in the case of Isidore Geoffroy, who backed away from many of the positions taken by his father. Most of the other leaders of zoology after 202

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1830—Flourens, Blainville, Audouin, and Milne Edwards—could not be said to be strict disciples of either Cuvier or Geoffroy. Rather they borrowed parts of their doctrines from both in order to formulate a compromise between two extreme and untenable views. With increasing knowledge of comparative anatomy, the strict teleology imposed by Cuvier—the belief that every structure was created by God to fulfill a specific function—was difficult to maintain. As there was no place in his doctrine for rudimentary organs, Cuvier had little to say concerning them. Any perceived unity of plan within an embranchement was to be interpreted as a secondary result of similarity of function. Thus Cuvier had to ignore or explain away very real evidence of structural unity underlying diversity. Taken at his word, Cuvier would seem to have precluded the possibility of biological laws of nature. The Creator, constrained by no other law than "the conditions of existence," was to remain free to create a new part whenever it suited His design. In order to combat the chain of being, a "law" that would supposedly limit the Creator, Cuvier had had to argue that there was not a semblance of a scale of perfection in nature. Every animal was perfect in itself, and comparisons of higher and lower were meaningless, except in the most general sense at the level of the embranchement. Geoffroy, for his part, had undertaken to defend an even more unworkable doctrine than had Cuvier. He persisted in the belief that if the same number of pieces did not exist in the same arrangement in all animals, one could always supply physiological explanations for the disparity. Most naturalists after 1830 recognized that strict accounting for each element of organization was an impossibility. Unity of composition in Geoffroy's sense did not exist. It was also clear to most observers that the diversity of animal structure could not be explained by reference to morphological considerations alone. Even Geoffroy had failed to banish functionalism from his writings, because he was unable to define structures in completely abstract terms without reference to their function.4 Geoffroy's homologies between embranchements were especially difficult to maintain, dependent as they were on the controversial theory of arrests of development, which implied a return to the largely discredited linear series of animals. Isidore Geoffroy and the intellectual heirs of Cuvier might still dispute the relevance of final causes in biology. They might still argue over the meaning of the debate and who had triumphed. But by and large the leaders of French zoology at the Museum and Faculte des Sciences—Isidore Geoffroy, Flourens, Blainville, and Milne Edwards—all arrived at a similar compromise solution. On the one hand they all accepted the Cuvierian dogma of distinct embranchements. On the other hand, they all agreed that homologies were an essential part of natural history and that one could trace homologies between elements of organization whose function had clearly varied. All believed that ideas were as important as facts in the advance of science. And all agreed that the task of the zoologist was to seek regularities in nature, whether called "laws," "principles," or simply "tendencies," that could not be explained solely by the "conditions of existence."

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Cuvier's apparent success in the debate and the subsequent rejection of Geoffroy by the scientific community in the 1830s therefore did not result in Cuvier's doctrines thenceforth holding exclusive sway. Geoffroy had made a decisive impact upon French zoology. Thanks in large part to the insights gained from the work of Geoffroy and his followers, Cuvier's framework for zoology was considerably broadened by the next generation of naturalists. Philosophical anatomy, divested of its more speculative and rigid trappings, entered the mainstream of natural history and comparative anatomy as an indispensable tool for clarifying the natural system of classification. In a very real sense, neither Cuvier nor Geoffroy "won" the debate. The synthesis that emerged in the 1840s was more flexible and sophisticated than the doctrines of either combatant. Other aspects of the debate—professionals versus amateurs, philosophy and religion, and the politics of science—were largely ignored. The leaders of science at midcentury had no quarrel with the scientific establishment, with professional standards, or with the judgment of scientists by their peers instead of by the public. They had little desire to examine at length the metaphysical commitments of their science. The trenchant critique of scientific conservatism and elitism was left to outsiders. Scientists dealt with the debate explicitly only at the levels of anatomy and methodology, although other considerations influenced their interpretations of the controversy. Although much weight can and should be given to the accumulated evidence of animal structure in promoting a settlement—that is, to factors internal to science—social factors were also involved. With Geoffroy's death, the last of the great men presiding over the early years of the Museum was gone. The next generation of naturalists were never public figures in the manner of Cuvier and Geoffroy. As institutions matured, science became more bureaucratized and insulated from public and political pressures. It was in keeping with the increasing professionalism of French science that the leading naturalists should wish to minimize unseemly controversy. With professionalism and insulation came also a tendency to increase the sphere of natural knowledge, that is, to enlarge the domain of explanation by natural law. Philosophical anatomy when properly circumscribed could lead to theoretical generalizations that lent luster to natural history and its practitioners without seriously undermining political and religious orthodoxies. For most naturalists (with the exception of Blainville), the prestige of natural history was better assured not by denigration of Cuvier or of Geoffroy, but by looking back on them both as heroes of the Golden Age of French science. Accord was reached at the expense of reinterpreting Cuvier and Geoffroy, in some cases by significantly refurbishing their personalities and their doctrines. Cuvier, as depicted by his disciples, lost his rigidity. Often his disciples, not wishing to contradict their master, attributed to Cuvier a more liberal attitude toward philosophical anatomy than he had in fact possessed. Duvernoy even gave Cuvier credit for being one of the founders of philosophical anatomy.5 In Flourens's interpretation, Cuvier did not object to unity of composition within an embranchement, only to homologies between embranchements, and thus he

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interpreted the debate as a matter of four plans or one. Geoffroy, in his son's biography, was divested of his brashness and his tendency to flout the scientific establishment. His evolutionary theories were toned down and his excursion into physics with the doctrine of soi pour soi was conveniently ignored. All of this revisionist history served to bring Cuvier and Geoffroy closer together and to dissolve the battle lines of 1830. Whereas in France naturalists did not always admit that they were taking a compromise position, for they were not fully aware of the very basic conflict between morphology and a strict teleology, in Britain, where naturalists were steeped in the British Protestant tradition of natural theology, the acceptance of philosophical anatomy took place in a more self-conscious manner. It was an English naturalist, Richard Owen, who most clearly recognized the disparity between Geoffroy and traditional Cuvierian teleology. To Owen, who had become immersed in the Cuvier-Geoffroy debate on a visit to Paris in 1831, neither teleology nor morphology alone was adequate to explain animal structure. The laws of nature had thus to encompass both a morphological and an adaptive principle. The interpretation of the debate as a conflict between a morphological and a teleological view of nature, an interpretation suggested by Owen, derives not from the French, but from the British context. As Dov Ospovat has pointed out in his study of the development of Darwin's theory of evolution, the rejection of teleological explanation as sufficient for the understanding of animal structure was a major achievement of naturalists of the 1830s and 1840s. In France, Britain, and Germany, the leading naturalists discarded strict functionalism—the belief that every organ was useful to its possessor—but without rejecting teleology itself. They came to hold that purpose was revealed not just in the adaptation of means to ends but also in the system of laws of organization.6 A fruitful synthesis of teleology and morphology depended upon a more extended knowledge of embryology. One serious weakness in the anatomical theories of both Cuvier and Geoffroy was their ignorance of embryology. Cuvier had based his classification of the animal kingdom on the adult animal alone, ignoring the data of physiology and embryology. Despite accumulating evidence to the contrary, he retained throughout his career a preference for the antiquated theory of preformation over epigenesis. Geoffroy, who believed in a true process of development, an epigenesis, and recognized the importance of embryology in determining homologies, nonetheless applied the embryological criterion in only the most superficial manner. He examined, in practice, only the final stages of the formation of the fetus, counting centers of ossification as bones. Cells, germ layers, histological and morphological differentiation, and gill arches—fundamental concepts of German embryology in the 1820s and 1830s—were all but unknown to him. Moreover, he insisted upon the recapitulation theory, which to most of his colleagues was an untenable oversimplification of embryological observations. A revised understanding of embryology was discovered to be the key to the unraveling of homologies. That knowledge came primarily through Germany. With the work of the German embryologist Karl Ernst von Baer, embryol-

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ogy took on a new relevance and became the paramount criterion for the determination of homologies.7 In his celebrated study, Ueber die Entwicklungsgeschichte der Thiere (1828), von Baer subjected recapitulation to a severe critique. In its place, he proposed a branching conception of development which was later taken over by Darwin into evolutionary theory. Von Baer's study of embryology revealed no unity of plan among all animals except at the very commencement of development. All animals began life as a single cell, but almost immediately thereafter their development diverged into four totally separate paths, corresponding to Cuvier's four embranchements. Development, according to von Baer, was a progression from the generalized to the specialized. The chick, to which von Baer devoted particular attention, first revealed the attributes of the vertebrate type, then those of the class of birds, and then the characteristics of order, genus, and species. At no time in its development did it take on the characteristics of an adult reptile or fish. While a higher animal might in the course of development appear similar in some respects to the adult state of animals lower in the scale, it would never take on all the characteristics of a lower animal.8 The power of German embryology is especially revealed in the resolution of the long-debated problem of the homologies of the opercular bones in fishes. An astonishing number of naturalists took up the problem of the operculum in the wake of Philosophie anatomique. A few, like the British naturalist Robert Edmund Grant, agreed with Geoffroy that the opercular bones corresponded to the tiny bones of the middle ear in mammals. Most other embryologists and comparative anatomists, including von Baer, Richard Owen, the Swiss naturalist Louis Agassiz, and the British anatomist Rymer Jones, admitted with Geoffroy that the opercular bones could not have been created entirely de novo for fishes, but offered alternative and less controversial homologies than Geoffroy's.9 But it was German embryology, and in particular the study of the development of gill arches, that eventually led to the currently accepted homologues in mammals of opercular bones, and the homologues in fishes of the bones of the middle ear. In 1825 Martin Heinrich Rathke discovered gill slits in the embryos of mammals and birds. By following the development of the gill arches, German embryologists could determine homologies with an accuracy that was impossible by the study of the adult and the close-to-term fetus. Rathke observed in the embryo of birds that the membranous operculum grew out of a flap at one edge of the hyoid arch. In the corresponding membranous operculum in fishes, there later developed the opercular bones and the branchiostegal rays. Thus Geoffroy's comparison of the hyoid and the branchiostegal rays in fishes was invalidated. In Rathke's determination, opercular bones were in one sense peculiar to fishes, while at the same time they were indicated in rudimentary form in the development of the higher vertebrates.10 By a similar study of the gill arches, Karl Bogislaus Reichert was able in 1837 to trace the homologies of the bones of the middle ear. He showed that these bones in mammals developed from the first vertebral arch and were the equivalent of primitive structures of the cranium of reptiles and other lower verte-

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brates. Mammalian ear ossicles, he found, were homologous to bones concerned with fish jaw attachment.11 This result, implying no less striking a change of function than that of Geoffroy's original homology, is often cited today as a classic example of the concept of homology.12 While Geoffroy's particular homology was discarded, his contention that homologues must exist was in the end vindicated. In France with Milne Edwards and in England with Owen, von Baer's embryology was incorporated into comparative anatomy, resulting in a new synthesis reflecting the strengths of both Cuvier and Geoffroy. It was this new and richer understanding of unity in diversity in the animal kingdom that Darwin encountered when he came to formulate his theory of evolution by natural selection. Philosophical anatomy not only provided some of Darwin's strongest evidence for evolution, but as Ospovat has argued, it also contributed directly to the formulation of Darwin's theory. Darwin at first rejected teleological explanations but still retained a belief in final causes, and particularly in perfect adaptation: the belief that within the constraints of the Creator's laws, all animals were perfectly adapted to their surroundings. By his effort to incorporate into his evolutionary theory the morphological laws worked out by Owen, William B. Carpenter, Milne Edwards, and others, Darwin eventually came to reject not only teleological explanation but also perfect adaptation. The evolutionary theory that resulted was one in which there was ample variation, and in which animals were only approximately adapted to their environment. His evolutionary theory of 1859 was therefore significantly different from that which he had articulated in his sketch of 1842 and essay of 1844.13 With the advent of the battle over evolution raised by Darwin's publication, the Cuvier-Geoffroy debate was given a new life in France. As Geoffroy's ideas were reinterpreted by Darwin, so too was the Cuvier-Geoffroy debate reinterpreted in the light of evolution. Both those who were opposed to evolution and those who favored it hearkened back to the debate as a momentous battle over evolutionary theory. The progressives were inclined to see the debate not as a conflict between Cuvier and Geoffroy but rather as a conflict between the conservative Cuvier and France's founder of evolution, Lamarck. Thus, the debate not only provided a point of departure for biological theorizing in France and England, but also served as an important chapter in the history of evolutionary biology. Before 1859 it emphasized the importance of theories in biology and the need to search for morphological laws. It led to the abandoning of teleological explanation as sufficient in explaining animal structure, and gave encouragement to new approaches to embryology. The new understanding of philosophical anatomy in the works of Owen, Milne Edwards, and Carpenter, all influenced by the debate, provided ammunition for Darwin and also led him to rethink his theory of evolution. In France it served nonacademicians as a focal point for disenchantment with the scientific establishment. The debate thus took on many guises, as it was interpreted by people with different interests, in different decades, and in different national contexts.

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Interpretations and Reconciliations: Isidore Geoffroy, Flourens, and Blainville

In the 1840s and early 1850s, three of the major figures in French zoology took up in their writings the subject of the Cuvier-Geoffroy debate. In some ways they differed as much as any three men of science might. Pierre Flourens was an ardent apologist for Cuvier, while Isidore Geoffroy Saint-Hilaire was an equally devoted admirer of his father. Henri de Blainville, on the other hand, had nothing positive to say about either rival. Flourens subscribed to final causes, Isidore Geoffroy to unity of composition, and Blainville to the animal series. These substantial differences of doctrine were reflected in their differing assessments of the significance of the controversy. Nevertheless, the scientific doctrines of all three men were shaped by the combined influence of Cuvier and Geoffroy. Each, in his own way, reconciled the two extremes represented by the two men in the debate. Say what they would, Isidore Geoffroy, Flourens, and Blainville were strict disciples neither of Cuvier nor of Geoffroy. Born at the Museum in 1805, Isidore Geoffroy was given all the assistance required to make an early start on a successful career as a zoologist.14 At the age of eighteen, he was publishing on mammals, and shortly afterward, in 1824, he was appointed an aide-naturaliste. By 1829 he had received his medical degree and was acting as a temporary suppleant to his father at the Museum. His three-volume classification of malformations of the fetus of 1833 gave teratology its name, secured its place among the sciences, and obtained for its author a seat in the Academie at the exceptionally early age of twentyseven.15 In 1838 Isidore Geoffroy left Paris to become dean and professor of zoology at the newly created Faculte des Sciences at Bordeaux. Three years later, he was back in the capital, having obtained his father's chair at the Museum. When Blainville died in 1850, Isidore Geoffroy, following once again in the path laid out by his father, acquired a chair at the Faculte des Sciences as well. His research centered on teratology and the effects of domestication upon animal structure and behavior. All contemporary biographers were struck by the marked contrast of character between father and son. Nicholas Joly, a disciple of Isidore Geoffroy, compared the two Geoffroys in typical terms: I see in the first the genius who creates and who divines; in the other, the mind that clarifies and enlivens. To Etienne belongs the power of the idea; to Isidore the lucidity that enables it to prevail. To one, boldness and sometimes temerity of views; to the other, prudence and wisdom of conception. To one, war and great blows of the sword; to the other, the peaceful and courteous battle, full of deference at the same time as firmness when he combats Cuvier, and for that reason, more sure of his triumph.16

Where Etienne was foolhardy, Isidore was cautious; where Etienne's writings were obscure and diffuse, Isidore's were remarkable for their clarity and systematization; and where Etienne was described as impetuous and emotional, Isidore was seen as calm and practical. Well received by all his contemporaries, Isidore Geoffroy did much to moderate the image of his father and of the so-

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called ecole philosophique whose leadership he inherited. Although he was said to have carried high the banner of the school of philosophical anatomy,17 he found no significant adversaries to engage in combat. His father's biographer and most ardent advocate, Isidore Geoffroy was nonetheless sufficiently influenced by the Cuvierian opposition to back away from several positions taken by the elder Geoffroy. Although he frequently used the phrase "unity of composition," he was less insistent than his father that it ought to apply to all animals. Indeed, he was at pains to distinguish between his father's method, the "theory of analogues," which he believed to be a true method applicable within embranchements, and his theory, "unity of composition," which was a goal toward which science should strive, but as yet unproven. In his hands unity of composition was no longer a demonstrated verity, soi pour soi was restricted to embryology and teratology, and the power of the environment to modify animal structure was reduced to producing a limited variability. And although he took the lead in defending synthesis in biology, Isidore Geoffroy always insisted that generalizations ought to be preceded by detailed factual work.18 In his laudatory and apologetic biography of his father written in 1847, Isidore Geoffroy tried, insofar as it was possible, to attribute to Etienne his own more moderate ideas. Thus he ascribed to his father his own restricted concept of transformism, despite the evidence that the elder Geoffroy had subscribed to a widespread mutability. In order to combat the critics who compared his father to the Naturphilosophen, Isidore Geoffroy purposely built up Geoffroy's credentials as a descriptive zoologist before entering into a discussion of his general ideas. His philosophical writings of the 1830s were conveniently ignored. Geoffroy thus emerged as a far more empirical and systematic thinker than in fact he was. As Isidore Geoffroy saw it, the clash in 1830 was inevitable, given the initial disagreement between Cuvier and Geoffroy dating from 1803 on the value of taxonomy. Cuvier's belief that taxonomy was the essence of science led logically to final causes, preformation, and immutability of species, while Geoffroy was led to reject final causes and to accept epigenesis and the variability of species. However, the differences between Cuvier and Geoffroy were not limited to ideas such as these, but extended as far as basic methodological principles. Isidore Geoffroy interpreted the Cuvier-Geoffroy debate as primarily a confrontation of two great schools of scientific methodology: the "school of facts" and the "school of ideas." The schools were by no means complementary. While Cuvier's "school of facts" was depicted as an unnecessarily restrictive force in science, Geoffroy's "school of ideas" was presented as advocating the true scientific method. The school of Cuvier, according to Isidore Geoffroy, could be characterized by "its almost exclusive research of observable facts," while that of Geoffroy was marked "by its constant tendency toward generalization and abstraction": The first, the ecole positive, as it calls itself, or the ecole timide, as others have named it, accords pre-eminence to facts over ideas. By the agency of its illustrious

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leader, it has even declared on several occasions that it rejects all theories as necessarily imbued with error, or accepts them only provisionally because they can lead to new facts. The second, the ecole systematique according to some, and the ecole progressive or philosophique according to others, subordinates, on the contrary, observation to abstraction, and so to say, places facts in the service of ideas. The latter are the true goal of science and the former only the means for achieving it.19

Like Goethe, who had offered a similar interpretation of the debate in his articles of 1830 and 1832, Isidore Geoffroy held that science ought to consist equally of facts and ideas. Whereas Cuvier and his school had sought to restrict science to particular facts alone, the Naturphilosophen, portrayed as the antithesis of Cuvier, had attempted to dispense with particular facts altogether. Only his father and his father's school had found the happy medium by discovering facts first and generalizing afterwards: "The facts first; after them, and by means of them, ideas."20 Sharing none of his father's tendency to rebel against the scientific establishment, Isidore Geoffroy deliberately avoided dealing with the religious and philosophical ramifications of the debate. Although he agreed with his father in rejecting final causes in science as a metaphysical attempt to divine the intentions of the Creator,21 he refused all further discussion of religion in his writings. Nor did he make mention of institutional or governmental politics. As far as Isidore Geoffroy was concerned, the conflict between his father and Cuvier was strictly a question of scientific ideas and scientific methodology. Thus, it was Isidore Geoffroy's role to moderate the controversy and to disassociate Geoffroy from his radical defenders. In his interpretation of the debate, Isidore Geoffroy placed his father in a positive light, for few scientists in the 1840s would dispute the necessity of theorizing in science. From the biography, which was widely read, the elder Geoffroy's system emerged not as the aberrant product of genius but as the cautiously and logically deduced consequence of an extensive interrogation of facts. By ignoring or silently revising his father's more radical hypotheses, Isidore Geoffroy was able to render the remaining parts of his father's doctrine acceptable to scientific contemporaries. Pierre Flourens, permanent secretary of the Academie des Sciences from 1833 to 1867, became the chief heir and apologist for Cuvier. Professor of human anatomy at the Museum in 1832, he traded the chair in 1838 for the recently created chair of comparative physiology. At the College de France he taught the course of natural history of organic bodies for many years before becoming titulary in 1855. Although not strictly a zoologist, Flourens became the major public spokesman for zoology in France in the three decades following the debate. Generally regarded as a disciple of Cuvier, Flourens did not altogether abandon the influence of his first mentor, Geoffroy. His doctrine was in fact a compromise between Cuvier and Geoffroy, as can be seen from his interpretation of the debate as a matter of one plan versus four. In his eloges of Cuvier and Geoffroy read before the Academie, Flourens was above all a diplomat. His eulogy of Cuvier, full of the highest praise for Cuvier's paleontological research and for his discovery of "the true method in

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natural history," made no mention whatever of the debate.22 In contrast, in his eloge of Geoffroy, read nearly twenty years afterward in 1852, he went out of his way to magnify the importance of the confrontation. "The world was divided," he declared, the "austere" thinkers siding with Cuvier and the bold minds supporting Geoffroy. In the eloge of Geoffroy, Flourens was careful not to take sides, but from his earlier writings his sympathies were well known. In some places, he agreed with Isidore Geoffroy that the debate concerned facts and ideas in science, and there he praised Geoffroy for his boldness of vision. But when he considered the conflict substantively rather than methodologically, he characterized it as a question of whether there were four distinct embranchements or only a single plan of animal structure.23 By choosing to describe the debate in this way, Flourens could readily argue that Cuvier was the victor. According to Flourens's reconstruction in the eloge of Geoffroy, the "main glory" of Cuvier was to have reformed the classification of the entire animal kingdom. He had justly established four plans, four essentially distinct types. "This beautiful order, exquisite fruit of the most perfect application of the [natural] method," was menaced by the increasing popularity of the ideas of Geoffroy, "who sought only a single plan, a single type."24 As most naturalists had adopted the four embranchements, Flourens could imply, without explicitly stating, that Cuvier had indeed prevailed. Flourens's interpretation of the debate in fact misrepresented Cuvier's position and revealed the impact that Geoffroy had made on French zoology. Flourens ascribed to Cuvier his own view that there was a unity of plan within each of the four great divisions of the animal kingdom. Unlike his benefactor, Flourens had no doubt of either the originality or the value of philosophical anatomy. While he noted that references to unity of plan in the animal kingdom could be found in the works of Buffon, Vicq d'Azyr, and Camper, he freely credited Geoffroy with interpreting the idea in an entirely original way and thus creating a new science. "That which distinguishes M. Geoffroy as a zoologist," wrote Flourens, "is the perception, as just as it was immediate, of the analogies of beings, what he himself so well called the "sentiment of affinities."25 Flourens acknowledged Geoffroy to have been the first to have brought comparison to bear on "the primitive and constituent elements of organs," and the first to study the fetal stage to discover homologies. "The glory of M. Geoffroy," Flourens confidently predicted, "will be to have founded the profound science of the intimate nature of beings: philosophical anatomy."26 For Flourens as for other leaders of French zoology, the triumph of four plans of nature over one was of vital importance, for the four embranchements served as the chief bulwark against the infiltration of evolutionary ideas. Homologies, unity of composition, teratology, arrests of development, and the principle of attraction of similar parts—Flourens praised them all when properly limited, but he thoroughly rejected any hint of transmutation of species. In an otherwise sympathetic review of the work of Buffon, published in 1844, for example, he took a harsh view of Buffon's speculations on evolution. Buffon had been led astray, according to Flourens, because he had not sufficiently studied the invertebrates to recognize that there were four distinct plans.27 In

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the eloge of Geoffroy, Flourens tried to minimize Geoffroy's commitment to evolution by disassociating evolution from the rest of his work. Evolutionary ideas were "only accessories" tacked on "toward the last years of his life" to "his principal ideas on the laws of animal organization": These views [on the mutability of species] where the real is not sufficiently separated from the ideal, are not peculiar to M. Geoffroy. They are divorced from that grand and beautiful collection of new and fundamental laws which constitutes his real doctrine and to which his name will always remain attached.28

The animal mummies that Geoffroy had brought back from Egypt early in his career were to Flourens the decisive proof that transformation of species had not occurred. Flourens did not follow Cuvier in insisting on the primacy of function or on the subordination of homologies to the principle of conditions of existence. Although Flourens defended final causes, he saw no need to link final causes exclusively to teleological explanation. For Flourens, the debate did not concern teleology versus morphology; he took for granted that both were necessary in comparative anatomy. Nor did it directly concern evolution. Even after Darwin, in 1864, he reiterated the point: "In other terms, is there a single plan, or are there four? That is, once again, the entire question, the entire discussion."29 With Cuvier's death in 1832, the scepter of zoology passed for a short time to his former protege and recent antagonist, Henri de Blainville. Blainville was named to Cuvier's chair of comparative anatomy at the Museum, which he held along with his professorship at the Faculte des Sciences until his death in 1850. Considered by many a misanthrope, a worthy successor to Moliere's Alceste,30 Blainville had little good to say about either opponent in the debate of 1830. He believed he owed nothing to them. In some respects Blainville was right. His own doctrine of the "animal series" was in one sense peculiar to himself and to a small number of disciples. Where Cuvier had found unbridgeable gaps, Blainville had sought transitional forms. Fossils as well as living animals found their place in the unique series of animal forms, which, like its predecessor, the chain of being, attested to God's wisdom in the Creation. In place of a succession of catastrophes followed by new creations, Blainville went back to the traditional eighteenth-century belief that there had been only a single creation. God had created all the members of the animal series at a single point in time, but since the beginning, many forms had gradually become extinct, leaving gaps in the once perfect scale.31 Despite the idiosyncrasies of Blainville's personality and scientific theories, he attracted a number of followers, all obscure names today, but some of whom held respectable posts in Paris and the provinces. Even Flourens was sympathetic to Blainville's animal series and to the possibility of a unique creation.32 Blainville won converts because the linear series and single Creation fulfilled a real need for those who, like Blainville himself, found neither successive creations nor evolution palatable. Anomalies in their day, Blainville's theories could not easily be supported by the data of comparative anatomy or

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paleontology. Blainville himself was soon overshadowed by a new and far more personable leader of French zoology, Henri Milne Edwards. In other respects, Blainville's doctrine, however unusual, owed significant debts to both Cuvier and Geoffroy. Like Cuvier, Blainville attached great importance to comparative anatomy, paleontology, and the natural system of classification. He was an ardent defender of "final causes" in science (and going further than Cuvier, one of the few French scientists to link explicitly science and religion). While his early version of the series tried to eliminate the four embranchements altogether by placing vertebrates, articulates, mollusks, and radiates at different levels in his system and by postulating such intermediary groups as the "articulo-mollusks," his final version of the series which he proposed in 1839 was in many respects closer to Cuvier. The four embranchements were in effect reinstated as four of five primary divisions of the animal kingdom. The five "Types"—Osteozoa (vertebrates), Entomozoa (articulates), Malacozoa (mollusks), Actinozoa (radiates), and Amorpha (no regular form)— could be ordered, but they did not necessarily shade into each other. In other words, Blainville admitted the possibility of a gap between the lowest form in one embranchement and the highest in the next. Blainville's final version of the animal series was, in fact, a hierarchical series of groups, as was found in most other classifications. The difference was that at each level, including the species, the groups could be linearly ordered.33 Blainville also appreciated the novelty and importance of philosophical anatomy, while at the same time rejecting unity of composition. Homologies, he believed, were the very essence of comparative anatomy. He admonished Cuvier for having limited comparative anatomy to a description of the parts and having ignored philosophical anatomy. Geoffroy's memoirs of 1807, Blainville wrote, "restore true comparative anatomy, that which Vicq d'Azyr had, so to say, created, which consists not in purely and simply describing the pieces of the skeleton of an animal, [or] in limiting oneself to comparing them to the more or less analogous [pieces] in animals of the same class, but in extending the comparison to the type itself as a whole, and above all, to the most elevated species." The search for homologies, he added, "was only a new denomination given to comparative anatomy, which considered scientifically, as it ought to be, could be nothing else."34 From the outset of his career, as we have seen, Blainville had been an enthusiastic promoter of philosophical anatomy, but in the end, he rejected in explicit terms Geoffroy's key generalization of philosophical anatomy, unity of composition. In 1840 Blainville wrote, "Just because there was a plan in the creation of each being, in each group of beings, and in the totality of creation, it is not necessary to conclude that there is a unity of plan, of composition, and of disposition in the organization of animals."35 The same organs did not reappear in differing degrees of development throughout the animal series. As one ascended the scale, new organs were added and other organs eliminated. Even the connections or arrangements of parts shifted when one passed from one type to another. According to Blainville, Geoffroy misunderstood the real significance of philosophical anatomy because he did not appreciate the animal

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series. In fact, by tracing organs through all their manifestations and revealing hidden affinities, homologies served as an indispensable aid in constructing the series. For Blainville, as for other French naturalists, teleology and morphology were not in contradiction. Despite his commitment to final causes, Blainville saw no need to insist that every structure served a useful function. Some structures—vestigial organs, in particular—existed solely to attest to the series. As the eighteenth-century chain of being had served to manifest God's design, the nineteenth-century animal series offered to Blainville an even more convincing proof of the existence of an all-wise Creator than any example of functional adaptation. Blainville took up the subject of the debate only in response to widespread criticism of the published lectures of his course on the history of the science of organization, taught at the Faculte des Sciences in 1839. Blainville's history of science was organized about the great men, each of whom personified an era. When he came to the modern era, instead of choosing the obvious leader of the life sciences, namely Cuvier, Blainville singled out Lamarck to represent the epitome of early nineteenth-century French science. Blainville, an antievolutionist, thus became one of the few French scientists to pay tribute to Lamarck in the pre-Darwinian era. A true savant devoted solely to science and not to patronage politics, Lamarck, according to Blainville, had been alone in understanding the significance of the animal series. As a deliberate slight to Cuvier, Blainville discussed Cuvier's work only as an appendix to that of Lamarck. With a perverse effrontery, he sought to minimize Cuvier's originality by ascribing the renaissance of comparative anatomy to Vicq d'Azyr and the creation of the science of paleontology to Pallas. Geoffroy was eliminated from the history altogether. Instead, Blainville turned to Lorenz Oken to personify the principles of philosophical anatomy. For his cavalier dismissal of Cuvier, Blainville received an almost universal reproach.36 Only then did he determine to justify his singular judgments by writing an epilogue to his history on Cuvier and his rival Geoffroy. Drafted some time in the late 1840s, but not published until 1890, Cuvier et Geoffroy Saint-Hilaire was a running diatribe against Cuvier. The book was intended, Blainville said, to show "that the works of M. G. Cuvier, however numerous they are, especially as they appeared in the enormous list of them that his panegyrists have given [us], are far from having the import and consequently the favorable influence that those of M. de Lamarck have had, to whose memory only a single eloge has been devoted, and what an eloge!"37 Blainville proceeded to dissect Cuvier's writings one by one, pointing out with obvious pleasure every minute anatomical error. An excellent anatomist, he had the advantage over Cuvier that he did his own dissections instead of assigning them to subordinates. Thus, he had little difficulty making legitimate corrections to his predecessor's work. Blainville found many reasons, both scientific and political, to attack Cuvier. He chastised him for accepting administrative positions in addition to his scientific posts and for politicizing science. He criticized him, too, for becoming

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mired down in classifications and descriptions and losing sight of any broader issues in science. In his estimation, Cuvier had lacked a true scientific doctrine; he was an "eclectic," a derisive term that Blainville associated with "individualistic Protestantism."38 Flourens, one of the many to label Blainville a misanthrope, accused him of being animated by preconceived views and a spirit of contradiction.39 Blainville had no higher opinion of Geoffroy than he did of Cuvier. In his book on Cuvier and Geoffroy, he indicated at every opportunity how Geoffroy had slighted him and failed to give him credit for his ideas. In return, Blainville described Geoffroy as a poor anatomist who had been led by his unrestrained imagination into the pursuit of "anatomical eccentricities."40 In his commentary on the debate, Blainville had few good words to say for either participant and did not think any of the issues were discussed in a competent manner. According to Blainville's characterization of events, both men were disputing an anatomical question when neither of them had a sufficient understanding of anatomy. Cuvier had been all but forced to descend into the polemical arena by Geoffroy's "more or less eccentric propositions" regarding the mollusks. The confrontation itself was dismissed as of little account: "one will see two zoologists who have never really studied in a practical way any of the points of the science of organization debating the details of positive anatomy and easily convincing themselves of each other's evident errors."41 Isidore Geoffroy, Flourens, and Blainville approached the debate from very different perspectives and as a result came to very different conclusions concerning the significance of the controversy. Isidore Geoffroy emphasized the contrast of methodology, Flourens, Geoffroy's assault upon the four embranchements, and Blainville, the personalities and ineptitude of the protagonists. Yet the doctrines of all three were shaped by the combined legacy of Cuvier and Geoffroy. All believed in some version of the four (Blainville adding a fifth) separate embranchements: vertebrates, mollusks, articulates, and radiates. All renounced (or did not insist upon) unity of composition in the animal kingdom as a whole. All pursued the goal of perfecting the natural system of classification. At the same time all incorporated philosophical anatomy as an integral part of their work. For all three, there existed in nature a plan or plans of animal structure that transcended similarities of function. Philosophical Anatomy Triumphant: Milne Edwards and "Physiological Zoology"

Of all French naturalists to respond creatively to the challenge posed by the controversy between Cuvier and Geoffroy, the most significant for the future of French zoology was Henri Milne Edwards. Blainville's reign as France's chief zoologist was relatively brief; within a decade, power had passed to a new naturalist and a new school—Milne Edwards and the so-called school of "physiological zoology."42 By midcentury, Milne Edwards had become the undisputed leader of French zoology. Professor at the Faculte des Sciences

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from 1844 until his death in 1885, he was the first French naturalist to train systematically several generations of disciples. For a long time just another public teaching forum, the Faculte des Sciences after the 1840s was becoming the focal point of zoological education. Because of his pivotal leadership in the Faculte, first as a professor and after 1849 also as dean, Milne Edwards shaped the direction of French zoology in a more definitive way than had Isidore Geoffroy, Flourens, or Blainville.43 More than any other naturalist he was responsible for the introduction of philosophical anatomy into the mainstream of French zoology. Profoundly influenced by the Cuvier-Geoffroy debate at a critical time in his career, Milne Edwards succeeded in combining teleological functionalism and morphology in a new and rich synthesis that was adopted by many of the leading French zoologists in the decades before Darwin. Milne Edwards's career was fostered primarily by his alliance with the Brongniarts, but also with aid from Geoffroy. Born in Bruges in 1800, a younger child in a large family of a Jamaican planter, Milne Edwards became a French citizen when Belgium became part of the Napoleonic Empire. In Paris he took a medical degree with little intention of practicing medicine, and he would doubtless have remained a wealthy dilettante and scientific amateur had not unforeseen financial circumstances in 1825 compelled him to earn a living. At the beginning of his scientific career he formed an intimate friendship and scientific collaboration with Audouin and soon became a protege of the Brongniart clan. When separate volumes of the Annales des sciences naturelles were instituted for botany and zoology in 1834, Milne Edwards joined Audouin as coeditor of the zoological series. Through the late 1820s and 1830s he supported himself by editing, writing textbooks, and teaching natural history in one of the Parisian secondary schools. As was often the case in this period, he was able to enter the Academie before he received a titulary professorship, inheriting in 1837 Frederic Cuvier's seat in the section of anatomy and zoology. In 1841 he entered the Museum when Audouin's unexpected death at the age of 44 left open the chair of entomology (a chair which he traded for that of mammalogy and ornithology in 1861). Because of Milne Edwards's contributions to morphology, Geoffroy had encouraged his career, choosing him in 1838 as his suppleant at the Faculte des Sciences. Upon Geoffroy's death, with the backing of Dumas, dean of the Faculte, and a fellow protege of Brongniart, he was appointed titulary professor. In contrast to his predecessors, who also taught in both places concurrently, Milne Edwards's first allegiance was to the Faculte rather than to the Museum. Milne Edwards's school has been called the "school of physiological zoology" because it placed new emphasis on the living animal. Where Cuvier had limited himself to studying the anatomy of dead invertebrates preserved in alcohol, Milne Edwards and Audouin observed embryological development and functional processes in live arthropods. This methodological innovation was of immense value in improving the natural system of classification, for in many cases stages in the development of invertebrates had previously been mistaken for new species. Milne Edwards and Audouin's excursions to the coasts of Normandy and Brittany in 1826, 1828, and 1829 to study marine invertebrates in their natural habitat have been said to mark the beginning of

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a long tradition of marine biology in France.44 Their memoirs on circulation in Crustacea that grew out of these travels won them, with the aid of Cuvier and Dumeril's enthusiastic report, the Academie's prize of experimental physiology in 1828.45 In addition to contributing to embryology, taxonomy, and comparative physiology, Milne Edwards and Audouin also turned their observations of invertebrates on the coast of France to the profit of philosophical anatomy. When Milne Edwards was first attracted to philosophical anatomy in the late 1820s, the conflict between Cuvier and Geoffroy was already apparent to the scientific community, and Milne Edwards sought to avoid becoming embroiled in it. Thus even his earliest work incorporated a synthesis between philosophical anatomy as advocated by Geoffroy and the limitations imposed by Cuvier. In his papers on the philosophical anatomy of articulates, he limited his discussion to a single embranchement, ignored Geoffroy's comparison of articulates and vertebrates,46 and offered no physiological explanations for structural modifications. Instead of comparing two forms by assuming questionable metastases in the manner of Geoffroy, or comparing two extreme forms directly, he proceeded by slow and sure steps through a progression of intermediary forms. He referred to Geoffroy in his papers as little as possible. The dual influence of Cuvier and Geoffroy was especially evident in Audouin and Milne Edwards's ingenious memoir on the philosophical anatomy of Crustacea published in 1828 and based on the materials gathered on the coast. Comparing the nervous system in the various orders of Crustacea, the authors intended "to show that there exists unity of composition in this system, and that the anomalous and varied modifications presented by animals of this class can all be reduced to a common type."47 By referring to a series of intermediary forms, they demonstrated that one could form a bridge from Talitra, or the sand-flea, whose nervous system consisted of two chains of thirteen ganglia, each united by transverse commissures, to the crab Maia, which has only two central ganglia, one at the head and one at the thorax. To corroborate data of comparative anatomy, they appealed to embryology, citing Serres and applying in a limited manner the theory of arrests of development. They noted that in certain larvae the nervous system was initially formed of two distinct chains as in Talitra, and that afterwards the chains fused toward the extremities to form two ganglia as in Maia. Thus, they discerned a parallel between the ontogeny of Crustacea and the series of adult forms.48 The difference between their circumspect approach and that of Geoffroy is evident from Geoffroy's report to the Academie on the memoir, in which he welcomed Audouin and Milne Edwards as members of the "new school," yet chided the two young men for not linking Crustacea and fishes and for not daring to compare the nervous systems of the crab and the crayfish (which had a ganglionic chain) directly.49 In his writings on philosophical anatomy in the 1830s, undertaken on his own, Milne Edwards began to work out a more flexible and fruitful approach to philosophical anatomy based on a deeper knowledge of embryology and a clearer understanding of relative perfection in the animal kingdom. Milne Edwards's new insight into progression in the animal kingdom was provided

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by the principle that came to be associated with his name—that of the division of physiological labor. First suggested in 1827, this principle was treated at greater length in the first volume of Histoire naturelle des crustaces published in 1834. In the simplest animals, Milne Edwards noted, there was little differentiation in structure and every part of the animal performed an identical set of functions. The internal anatomy of such an organism could be likened to a workshop in which each worker was employed at the same set of tasks. If such an organism, a polyp for instance, were cut into pieces, each piece would be able to survive and maintain all the functions of life. As one ascended the scale of being, however, the parts of organisms became increasingly dissimilar. While at first the same organ felt, moved, respired, and absorbed nutrients, step by step each function acquired its own instrument. Finally the various functions came to be performed by distinct parts. Complexity in the scale was thus characterized by a division and a distribution of functions.50 The principle of division of physiological labor enabled Milne Edwards to account for variation of form among Crustacea without the circumlocutions resorted to by Geoffroy. In his Histoire naturelle des crustaces, Milne Edwards envisioned the external skeleton of Crustacea as composed in essence of a series of "homologous" segments, each one a repetition of the previous. He claimed that in principle all Crustacea were composed of twenty-one segments. As a point of comparison, he singled out the case where the twenty-one segments were most distinct. The crustacean selected—the squillfish—also represented the primitive, least modified form, and thus was low in the series of Crustacea. As the division of physiological labor progressed, functions became localized, and the segments and their associated appendages became increasingly modified. Likewise, in the embryo the members at first all had the same form, but soon became dissimilar. Differences continued to be augmented until the animal reached the perfect state.51 In a memoir of 1835 on the value of embryology for the classification of Crustacea, Milne Edwards likened the "normal or common type" of a group of Crustacea, the point of departure for diverging development, to an archetype, although he did not use the term. The common type was "an ideal and abstract form which represents all that which the different organisms have in common and the mean of the differences which distinguish them from one another."52 Adopting the terminology Audouin used for insects, Milne Edwards determined homologies between segments, mouthparts, and legs in the various orders of Crustacea, but he did not feel compelled to provide explanations for all differences in number of parts. The appendages of the first segment, for example, were present only in the highest Crustacea. Each of the twenty-one segments except the last was provided with a pair of members which could vary considerably in form and function from order to order. By means of exhibiting a series of intermediary forms, Milne Edwards showed that the crab (a member of the Decapoda), which appeared to be covered by one large cephalothoracic carapace, and the Edriophthalma (Isopoda and Amphipoda), which appeared to be divided into a number of thoracic rings, were in fact similar in structure. He maintained that the carapace of the crab was nothing

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more than a prolongation of a cephalic segment, as found in the Edriophthalma.53 Thus Milne Edwards discerned unity in diversity, but without identifying a fixed number of parts in every animal. Philosophical anatomy as conceived by Milne Edwards was not an alternative to description and classification, but rather a powerful aid to achieving Cuvier's goal of a natural system of classification. Homologies were especially useful in classification when embryological evidence was brought to bear.54 For example, when Milne Edwards came to classify Crustacea, he rejected Cuvier's subordination of characters and the establishment of successive divisions. Instead he included among the Crustacea all forms that could be linked by "degradation" or simplification to the type of the group as represented by crabs and crayfish. In adopting this method, he wrote, one is not inhibited by these differences in structure, and one includes in the group of Crustacea all animals whose general organization, although less complicated, can be linked to that of the types of the class, and whose conformation recalls the transitory states through which the most perfect beings of the series have passed during the duration of their embryonic development.55

The key to classification was the study of development. The closer the pattern of development of two animals, the closer they ought to appear in the natural system of classification. Thus the parasitic Lernaea, formerly classed with the zoophytes, was transferred by Milne Edwards to the Crustacea because in its early stages of development it resembled other crustacea.56 This significant modification of Cuvier's principles of taxonomy could be traced on the one hand to Geoffroy and philosophical anatomy, and on the other to Blainville's serial classification of animals. In his early writings Milne Edwards was sympathetic to Serres's recapitulation theory, but later abandoned it for a branching conception of nature which recalled that of von Baer. He claimed, however, to have discovered it independently.57 While the recapitulation theory stipulated that a higher animal in the course of development repeated the forms of animals in the progressive series, the branching conception of nature suggested that perfection occurred by a series of progressive differentiations. Like the German embryologist, Milne Edwards came to believe that the process of development was a process of increasing individuality. Within a given embranchement, embryos seemed at first to follow similar paths and then at a certain point to diverge. Because the characters of each embranchement appeared almost from the outset in the developing embryo, embryology corroborated for Milne Edwards the primordial nature of the four embranchements. The characters of the embranchement gave way successively to those of the class, order, genus, and species. Thus, in theory at least, one could construct a natural classification based upon embryology. While the branching conception still allowed for arrests of development with respect to particular tissues or organs, it ruled out the recapitulation by the human embryo of forms of animals lower in the scale: I admit with Geoffroy Saint-Hilaire that there often exists a great analogy between the final state of some parts of the body of certain lower animals and the embry-

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onic state of these same parts in other animals belonging to the same type, but in which the organization is more perfected, and, with this philosopher, I will gladly call this state of permanent inferiority arrest of development. But I will guard myself against admitting with some of his disciples that the embryo of man or of any mammal represents at its various stages of development the less perfect species of animate creation.58

Perhaps viewing recapitulation as a portent of transformism, Milne Edwards launched a vigorous campaign against Serres. Milne Edwards's synthesis of philosophical anatomy and Cuvierian functionalism achieved its mature form in two works of the 1850s summarizing his doctrine: the introduction to his Lecons taught at the Faculte des Sciences and his Introduction a la zoologie generale. Milne Edwards combined on the one hand a commitment to the four embranchements as the cornerstone of zoology, and a theoretical belief in the primacy of function over structure, with a search for morphological regularities in nature. The theoretical framework which he articulated drew upon both Cuvier and Geoffroy, but the impress of Geoffroy is especially evident. Milne Edwards's mature works, in contrast to his earlier works, were full of enthusiastic praise for Geoffroy: for introducing the concept of homology, for promoting the study of embryology, for discovering the science of teratology, for advocating epigenesis, and for supporting a more philosophical approach to zoology in place of mere description.59 In the 1820s and 1830s it had been impolitic to pay tribute to Geoffroy, but once Geoffroy left active science, he could safely be recognized as one of the founders of modern zoology. Indeed, Milne Edwards credited Geoffroy with playing a major role in inspiring the movement toward a more "physiological" zoology: In recalling here this new phase of the natural history of animals, I could not, without injustice, forget the name of Etienne Geoffroy Saint-Hilaire, who, attacking with ardor a multitude of fundamental questions for the philosophy of zoology, has instilled in the minds of men a great impulse to progress and has done more than any other to direct the attention of observers to an order of facts from which this science now draws the most precious of its new wealth.60

Unlike Cuvier, Milne Edwards had no excessive fear of theoretical discussions in zoology. To be sure, he had less reason than Cuvier to fear generalizations in zoology, for science since Cuvier had become more bureaucraticized and insulated from public scrutiny, and professional standards were increasingly taken for granted. Naturphilosophie had ceased to be a threat, and evolution had not yet become one. Sharing little of Cuvier's fear that hypotheses in science could be abused, Milne Edwards freely declared that zoology ought to be philosophical. A naturalist should not limit himself to collecting facts, but rather seek out hypotheses, for they were the lifeblood of science. By linking facts, hypotheses made them useful, and moreover, provided an impulse to new research.61 Milne Edwards's theoretical discussions, however, had little in common with the broad speculations of Geoffroy. Rarely did he discourse on subtle fluids, the nature of life, or the fundamental laws of the universe. On most of the current speculative theories, Milne Edwards remained prudently

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on the side of the conservatives. He rejected spontaneous generation, the chain of being, the recapitulation theory, unity of composition as applied to the entire animal kingdom, and transformism. He was an acknowledged vitalist and advocate of final causes.62 His bulwark against all such theories was Cuvier's four embranchements: There is neither unity of composition nor unity of plan in this vast creation. The vertebrate does not resemble by its principal traits of structure either the mollusk, the insect, or the zoophyte, and as the greatest naturalist of our day, Georges Cuvier, has shown, there exist four fundamental types in the animal kingdom, four zoological conceptions from which all animal species seem to derive.63

While retaining the four separate embranchements as the cornerstone of zoology, Milne Edwards came to disagree with Cuvier on a number of important issues, foremost among them the role of biological "laws." Cuvier had not acknowledged laws of organization beyond the "conditions of existence," because any other regularities would have detracted from God's freedom to create. Milne Edwards deliberately sought patterns in nature, but at the same time he was careful not to claim for them the status of absolute law. Rather he cautiously referred to them as "tendencies of Nature in the creation of animated beings." Within each of the four basic plans, he noted first a "diversity of productions," and second a "law of economy" by which nature tended to employ the same materials in the same arrangement while varying their form and function.64 In principle, Milne Edwards agreed with Cuvier that God's conception of the function of an organ was prior to the assignment of structure, but in practice he often gave primacy to structure. He paid lip service to traditional teleology in passages such as the following: No, the physiological properties of the Animal are not, in my opinion, a consequence of its structure, but the raison d'etre of its structure. Each one of these admirable machines, in originating from the hand of the Creator, seems to me to have been called upon in advance to exercise a series of determined acts, and to carry in itself the germ of the power that will enable it to act,65before it is provided with the instruments necessary for the exercise of this force.

In fact, Milne Edwards was more inclined to explain diversity by morphological laws than by attributing it solely to the free choice of the Creator. In contrast to Cuvier, Milne Edwards regarded inequality of perfection among animals as "one of the characteristics of the zoological Creation" and an important means of achieving diversity. As a thorough opponent of the chain of being, Cuvier had eschewed almost all discussion of variable perfection in the animal kingdom. Instead he insisted that every animal could be said to be equally perfect, since each was perfectly fashioned to fulfill its role in nature. Milne Edwards replied that not all roles had the same importance. While denying the possibility of arranging all beings on a graduated linear scale of perfection, he nevertheless maintained that some animals could legitimately be said to be higher than others. Between the simplest and the most complex animals there existed "a multitude of intermediaries, and it is with reason that,

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in comparing them physiologically, the naturalist calls some higher animals and others degraded or lower animals."66 It was especially through the division of labor, Milne Edwards claimed, that nature tended to perfect the organism. At first nature merely repeated existing organs to give an organism a greater capacity for action. Only after exhausting this means did she form special instruments to serve each function. Moreover, these specialized instruments were initially derived from a common fund of materials. Materials serving a given function in one animal might be "borrowed" to serve another function in another animal. Nature resorted to all possible modifications of existing materials, before she undertook to create a new element: It seems thus that before having recourse to new resources to vary her products, she tried in some sense to exhaust each of the means that she used to create variations, and as much as she shows herself prodigal of variety in the work of Creation, she also appears economical in the means by which she obtains this wealth of results.67

While he did offer a few examples of possible new creations—e.g., the respiratory chamber in the higher mollusks68—he did not believe with Cuvier that God had unlimited freedom to create them. God introduced new elements only as a last resort. The principle of division of physiological labor and the branching conception of nature, both of immense importance for the future of biology, incorporated in a new synthesis Cuvier, Geoffroy, and German embryology. In articulating this new framework for zoology and comparative anatomy, Milne Edwards, the dominant figure in French zoology and leader of a school of disciples, successfully adapted and made fruitful application of many of the key concepts found in the work of Geoffroy Saint-Hilaire and the philosophical anatomists. At the same time he modified and restricted those concepts to meet some of the major objections of Cuvier. Milne Edwards did not discuss the debate in his published writings, but if he had, he would no doubt have agreed with his disciple Armand de Quatrefages that neither Cuvier nor Geoffroy emerged the victor. Milne Edwards's students continued cautiously to apply the theory of analogues, the principle of connections, the theory of accidental causes of monstrosities, and the theory of arrests of development, often acknowledging their indebtedness to Geoffroy. Geoffroy's ideas, thus circumscribed and refined, became part of the everyday arsenal of the French zoologist, and his eccentricities were all but forgotten. With Milne Edwards, there emerged a tacit consensus among French zoologists that traditional teleology and philosophical anatomy were equally necessary to explain the diversity of animal structure. Teleology and Morphology: The Debate in Britain

In France the reconciliation of morphology and teleology was for the most part implicit; in Britain, however, where natural theology was so widely popularized by both clergymen and scientists, there was a clear awareness that the def-

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inition of purpose in nature would have to be altered to accommodate the new understanding of unity of plan in animal structure. The shift in thinking was a significant one, as several recent historians have argued, for it prepared the way for Darwin by giving greater reign to the rule of law in place of the free will of the Creator. Because the British had to make a conscious effort to integrate homologies into natural theology, they interpreted the debate in the light of their own concerns as a conflict of "morphology" versus "teleology." These were terms that were rarely, if ever, used in France at this time. French naturalists, whether they considered themselves followers of Cuvier or of Geoffroy, were not fully aware that they were formulating a compromise between the two rivals. Cuvier's belief in the sufficiency of functional explanation was not usually combated directly, but most often simply set aside and ignored. Many—among them Blainville, Flourens, and Milne Edwards—continued to advocate final causes in biology, but saw no grounds for conflict. Without giving sustained attention to the problem, they as a matter of course adopted the idea that morphological similarities represented another form of design in nature. They assumed that philosophical anatomy revealed a larger purpose by enabling the naturalist to grasp laws of organization, to uncover unity in diversity, and to discern more accurately the unique natural system of creation. In Britain the compromise was of necessity couched in more explicit terms, with a definite acknowledgment that the acceptance of philosophical anatomy required a major modification of the accepted notion of final causes in biology. The British drew on both the French and the Germans for their understanding of philosophical anatomy, but the term "philosophical anatomy" and the concept of homology derived primarily from France and the controversy between Cuvier and Geoffroy. Principes de philosophie zoologique was not translated into English, but philosophical anatomy and the debate were brought to the attention of British scientists in the 1830s by a number of individuals, among them Robert Knox, Martin Barry, William B. Carpenter, Richard Owen, and above all, the Reverend William Whewell. Barry, Carpenter, and Owen combined morphology, teleology, and German embryology in a frutiful synthesis, similar to that of Milne Edwards, which had an important role in the genesis of Darwin's theory of evolution by natural selection.69 The conscious nature of the transformation of thought is well illustrated by the case of the philosopher and physical scientist William Whewell, whose three-volume History of the Inductive Sciences (1837) was responsible for introducing the debate to a broad segment of the educated British public.70 At first, clinging to the older notion of design in nature as restricted to adaptation, Whewell, fellow and tutor of Trinity College, Cambridge, and professor of mineralogy, rejected Geoffroy altogether. In his lengthy account of the debate, Whewell was disturbed by Geoffrey's contention that there existed ideally but one animal. Not only did Geoffroy's position tend toward evolution, but Geoffroy had dared to deny the importance of final causes in the life sciences: According to this theory [Geoffroy's theory of analogues], the structure and functions of animals are to be studied by the guide of their analogy only; our attention is to be turned, not to the fitness of the organization for any end of life or action,

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but to its resemblance to other organizations by which it is gradually derived from the original type.71

To a minister and author of one of the Bridgewater Treatises, this was unconscionable. He argued that in the biological sciences, as opposed to the physical sciences, teleological reasoning was indispensable. In the face of this perceived threat to religion, Whewell undertook a passionate defense of the traditional Argument from Design. Geoffroy's doctrine of unity of composition, insofar as it ignored purpose in nature, appeared to Whewell "utterly erroneous." Cuvier's "conditions of existence," by contrast, was pronounced by him to be a major contribution to biological theory, for Cuvier had appreciated the need for final causes. Reaffirming the time-honored view of purpose in nature, Whewell insisted that "The study of comparative anatomy is the study of the adaptation of animal structures to their purposes."72 Under the influence of Richard Owen, Whewell was soon to change his mind. He came to recognize that homologies were far more widespread than he had hitherto believed and that there existed many structures in animals of no immediate use to their possessor. In later editions of History of the Inductive Sciences, he admitted that the recent study of morphology had modified the former antithesis between "Morphology" and "Teleology." In order to reconcile the notion of design in nature with homologies, Whewell had to argue that there were other purposes in nature besides immediate utility. Law, similarity, and symmetry also attested to the wisdom of the Creator: If the general Plan be discovered after the contrivance has been noticed, the discovery may at first seem to obscure our perception of Purpose: but it will soon be found that it merely transfers us to a higher point of view. The adaptation of the Means to the End remains, though the means are part of a more general scheme than we were aware of.73

It was typical of the English response to contrast "morphology" and "teleology." Whewell's shift of thought, Michael Ruse has pointed out, was symptomatic of the very real difficulties faced by those trying to integrate science and religion in the decades before the Origin. The substitution of laws of structure, geological succession, and geographic distribution all tended to place limits on the action of the Creator and remove Him ever further from the immediate design of each created being.74 Martin Barry, a physician and embryologist who studied in Paris and Germany, and the physiologist William B. Carpenter both called upon von Baer's embryology in the 1830s to resolve some of the problems raised by the debate. On the basis of von Baer's embryological laws, Barry argued in two articles in 1837 that "unity of structure in the animal kingdom" lay not in adult animals but in the history of development. This was a subject, he noted in passing, to which Geoffroy had paid insufficient attention. Barry excused himself for concentrating in his papers on structure rather than function, for, he argued, both were "simultaneously contemplated in the same design." All animals shared a common origin and the same manner of development, according to von Baer's law of progression from the "homogeneous" to the "heterogeneous" or from

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the generalized to the specialized. It was in fulfillment of this law that useless and rudimentary parts found their meaning. The most reliable means of classification were not to be found in the consideration of the adult animal, where "function tends to embarrass," but in embryology, where "structure presents itself alone." Implicit in Barry's articles was the notion of a common structure and pattern of development upon which was superimposed teleological adaptation.75 Carpenter probably learned of the debate in 1834 from taking the course of comparative anatomy taught at the University of London by Robert Edmund Grant, a frequent visitor to Paris and sympathetic to Geoffroy.76 In his Principles of General and Comparative Physiology (1839), Carpenter incorporated von Baer's embryology into a predominantly morphological approach to nature. He insisted that, although the same elementary parts did not exist in all animals, in a more restricted sense within each of the four major divisions of nature, there was indeed a "unity of composition": "We arrive at the important truth that, where any new function, or great modification of function, is to be performed, no entirely new structure is evolved for the purpose;—the end being always attained by a corresponding modification in some structure already present."77 Arguing against Whewell's distinction between the need in the biological and in the physical sciences for teleological explanations, Carpenter also pointed to other ends in nature besides immediate utility. In fact, he claimed that the "philosophic Physiologist" must "disregard for a time, as in physical philosophy, the immediate purposes of the adaptations which he witnesses; and must consider these adaptations as themselves but the results or ends of the general laws for which he should search."78 Far from destroying design, the combination of unity and variety thus revealed by nature offered a stronger proof than ever of the existence of a wise Creator.79 "The Philosophical Anatomist, therefore, does not regard the object or function of a particular structure as a sufficient account of its existence; but, in attaining the laws of its formation independently of any assumption of an end, he really exhibits the primary Design in a much higher character, than in deducing it from any limited results of its operation [i.e., particular cases of adaptation]."80 Of all naturalists of any nationality, the man who gave the most deliberate thought to reconciling Cuvier and Geoffroy was the British comparative anatomist Richard Owen. As a young man, Owen was largely responsible for the introduction of Cuvierian comparative anatomy into Britain, and was thus sometimes called the "British Cuvier." His work is central to an understanding of the intellectual legacy of the Cuvier-Geoffroy debate. It was Owen who substituted the modern "homology" for Geoffroy's "analogy," clarified terminology, and publicized the concept. Trained in the British tradition of natural theology but also well versed in French science, Owen was especially able to articulate the contradictions between the two French rivals. Finally, it was largely through Owen that the new understanding of morphology reached Darwin and entered into his evolutionary synthesis.81 Like Milne Edwards and Louis Agassiz, the other two of the triumvirate of leaders of Cuvierian comparative anatomy in the 1840s and 1850s, Owen

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encountered and was deeply influenced by the Cuvier-Geoffroy controversy at a formative stage in his career. He first met Cuvier in 1830 when, in the wake of the July Revolution in France, Cuvier visited England. Owen, employed as assistant to the conservator of the Hunterian collection of anatomy at the Royal College of Surgeons, was still unknown in scientific circles in 1830. But because he could speak French fluently, he was entrusted with the task of escorting the prominent visitor through the institution. As a result of this opportune meeting, Owen received an invitation from the baron to come to Paris and study the rich collections at the Museum. The following year, the young man arrived in the French capital and was soon immersed in the issues raised by the debate.82 In 1865, in the "General Conclusions" of a three-volume work on the comparative anatomy of vertebrates, Owen, then superintendent of the natural history departments of the British Museum, looked back on the lasting impressions made by his visit to the Jardin des Plantes in 1831. In particular, he listed five issues brought forth by the Cuvier-Geoffroy debate: Unity of Plan or Final Purpose, as a governing condition of organic development? Series of species, uninterrupted or broken by intervals? Extinction, cataclysmal or regulated? Development, by epigenesis or evolution [preformation]? Primary life, by miracle or secondary law?

Upon his return home, he recalled, "I was guided in all my work with the hope or endeavour to gain inductive ground for conclusions on these great questions."83 Writing in 1865, six years after the publication of On the Origin of Species, Owen naturally emphasized the evolutionary debate. He wished to prove that, long before Darwin, he had come to the conclusion that species must be produced not by miraculous creation but by an unspecified "natural law or secondary cause."84 In 1831, however, the chief issue confronting Owen was the first named, rephrased by Owen more succinctly as "Homology or Teleology." Owen only slowly recognized the need for a compromise between Cuvier and Geoffroy. At the time the debate was raging in Paris, Owen thought Geoffroy's arguments in favor of homology were faulty and his examples inaccurate. "The logic, and, as it seemed, the facts, were on the side of teleology." But later in the decade, Owen's increasing knowledge of comparative anatomy brought about by his effort to classify the Hunterian collection "enforced a reconsideration of Cuvier's conclusions." He came, he said, to the realization that the concept of an "ideal type" was necessary "to make intelligible the 'unity in variety' pervading any group of organisms."85 Owen discussed the question of homology versus teleology in his courses on comparative anatomy given in 1837 and afterwards as Hunterian Professor at the Royal College of Surgeons, and in several publications in the 1840s, which included his Hunterian lectures on invertebrates (1843) and fishes (1844 and 1846) and, especially, two works entirely devoted to philosophical anatomy, On the Archetype and Homologies of the Vertebrate Skeleton (1848) and On the Nature of Limbs (1849).86

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In 1843 Owen offered for the first time in print a refinement of Geoffrey's terminology by substituting the term "homology" for Geoffroy's "analogy," and distinguishing between morphological similarities (homologies) and merely functional similarities (which he called analogies). He defined a homologue in nonevolutionary terms as "the same organ in different animals under every variety of form and function" and an analogue as "a part or organ in one animal which has the same function as another part or organ in a different animal." Analogies were thus evidence of convergence of functions due to adaptation to similar environments, not of essential similarities.87 In later publications, he subdivided homologies into three kinds of relationships: "special homology," corresponding to Geoffroy's "analogy"; "serial homology," or a comparison of parts within a single animal (also called "homotypes"); and "general homology," in which parts in a particular animal were compared to corresponding elements of an ideal type, or "Archetype."88 Owen realized that the acceptance of homologies entailed the abandonment of strict functionalism. To justify this step, Owen turned the familiar machine analogy of the natural theologians upside down.89 Authors of works on natural theology typically argued from human contrivance to God's design. In William Paley's oft-repeated example from his classic Natural Theology (1802), a watch, by its intricate mechanism, implied the existence of a watchmaker. How much more, the argument went, did the complex adaptations of animals imply the existence of an intelligent and all-foreseeing Creator. Owen, however, argued that in the case of machines made by man for the purpose of locomotion, the mechanisms were fundamentally different from those in animals. Machines were indeed all adapted expressly for their ends. Man did not place unnecessary shackles on his creative powers by devising instruments according to a common plan. The plan instead varied directly with the function: "There is no community of plan or structure between the boat and the balloon, between Stephenson's locomotive engine and Brunei's tunnelling machinery: a very remote analogy, if any, can be traced between the instruments devised by man to travel in the air and on the sea, through the earth or along its surface." Nor, Owen continued, should any greater conformity be expected in living things if final causes "were the sole governing principle of organization." "The teleologist would rather expect to find the same direct and purposive adaptation of the limb to its office as in the machine."90 Instead, one discovered that the limbs of vertebrates, whether employed for running, swimming, digging, or flying, were all constructed on a common plan. Many homologies could not be accounted for solely on the basis of functional requirements. One might argue, for example, that the bones of the human cranium were separate in infants and later fused together because parturition was thereby facilitated. But why then were the same bones separated in a bird which picked its way out of a shell, or in a marsupial which was only a fraction of an inch long when born? For what reason did the coracoid process of the scapula (the process that gives attachment to the biceps) develop from a separate center of ossification rather than by continuous ossification of the scapula? "These and a hundred such facts," wrote Owen, "force upon the contemplative anatomist the inadequacy of the teleological hypothesis to account

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for the acknowledged concordances expressed in this work by the term 'special homology.'"91 Such similarities of structure could not be explained, as Cuvier wished, by reference to similarity of functions. One could only explain the separate center of ossification of the coracoid process in human anatomy by observing that the process corresponded to a distinct bone, the coracoid, in monotremes, birds, and reptiles. But just as teleology did not suffice to explain the diversity of animal forms, neither did homology. Owen held Geoffrey's theory of unity of composition to be invalid even when restricted to the vertebrates. By no amount of juggling could all vertebrate skeletons be reduced to the same number of bones (or centers of ossification) arranged according to a single plan. The number of parts and even the number of centers of ossification could vary because of adaptive requirements. The femur, for example, was ossified from four centers in mammals, but from only one center in birds and reptiles. The explanation was teleological: as young mammals were ready to run at a early age, the shock to the leg would be diminished if the bones were ossified gradually from four separate points. Thus, Owen argued that one had to distinguish between separate points of the mammalian skeleton that were homologous to perfectly distinct bones in the lower vertebrates, and separate points of ossification which exist for a final cause.92 Owen contributed enormously to the clarification and extension of homologies of the vertebrate skeleton, yet he differed in significant ways from Geoffroy. Altogether, Owen's was a far more flexible and sophisticated treatment of philosophical anatomy than Geoffrey's. In his three-volume comparative anatomy of vertebrates, he painstakingly illustrated numbered homologous skeletal pieces throughout the vertebral series.93 Whenever possible, he corroborated his determinations of bones by embryological evidence. But Owen did not extend homologies beyond the vertebrates, for he believed with Cuvier and von Baer in the existence of four distinct types or embranchements. He rejected unity of composition in the animal kingdom and even within the confines of a single type. And he also firmly rejected another of the cornerstones of transcendental anatomy, the recapitulation theory. Instead, Owen, like Milne Edwards, came to adopt a branching conception of nature based on von Baer's embryology.94 This branching conception of nature was reflected in Owen's concept of the Archetype, perhaps his most significant departure from Geoffroy. The Archetype, Owen's ideal vertebrate and point of comparison, was the most primitive and unmodified form of vertebrate. It corresponded not only to a form low in the animal scale but also to a primitive and unmodified stage of embryological development. In structure, it resembled most closely the skeleton of a lungfish—essentially a series of vertebrae, the first four constituting the cranium, and scarcely developed limbs. In the lowest vertebrates, Owen claimed, the skeleton was not much removed from that of the Archetype. But the higher one went in the scale, the more the vertebrae were modified and specialized to perform different functions. Owen diagrammed the ideal vertebra, which consisted of a body or centrum, a neural arch above, a hemal arch

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below, and "diverging appendages." Single bones of the ideal plan could be "teleologically compounded" in order to serve special functions. Thus, the arms and legs were, according to Owen, teleological modifications of diverging appendages which have reached their highest development in man. Owen interpreted the arm as the diverging appendage of the fourth or occipital vertebra (one of the cranial vertebrae) of the Archetype, and the leg as the diverging appendage of another vertebra whose position in the series was less constant. He went so far as to speculate that on some other planet more than four diverging appendages could have been modified into additional limbs.95 The Archetype provided Owen with a clever solution to Geoffroy's problem of identifying the homologue in mammals of the operculum in fishes. According to Owen, the operculum was also a modified diverging appendage, found only in fishes, of the tympano-mandibular arch, or, in terms of the Archetype, a modified appendage of the hemal arch of the frontal vertebra, one of the vertebrae of the skull. This "solution," it can be seen, fell in the middle of the positions taken by Cuvier and Geoffroy. The operculum was not created de novo for fishes, but neither did there exist exact homologues of the opercular bones in the skeletons of the higher vertebrates.96 To explain the diversity of animal structure, Owen called upon the creative interaction of teleological and morphological forces in nature. In order to account for the underlying constancy of plan, he posited a "law of vegetative or irrelative repetition" resulting from a "polarizing force" (a concept derived from German science) similar to the force accounting for repetition of similar crystals. To account for modifications of the plan, he assumed the existence of an "adaptive or special organizing force." Both forces ultimately testified to the "superior design, intelligence and foresight" of the Creator. In the "adaptive force" could be found the familiar providential adaptation of means to ends, while the "polarizing force" enabled one to detect the Platonic Idea or ideal Exemplar in the Creation and pointed to the existence of an "Antecedent Mind" or Deity. Homologies therefore represented a higher form of teleology:97 The recognition of an ideal Exemplar for the Vertebrated animals proves that the knowledge of such a being as Man must have existed before Man appeared. For the Divine mind which planned the Archetype also foreknew all its modifications.98

Owen's synthesis of teleology and morphology was part of a wider development in British biology in the pre-Darwinian era. Ospovat has argued that the repudiation of strictly teleological explanation and the substitution of transcendental laws and forces by Owen and others in early nineteenth-century Britain marked a fundamental shift in the nature of biological explanation. He suggests that dividing pre-Darwinian naturalists into teleologists and nonteleologists is in some respects a more useful and revealing classification than grouping them according to the traditional dichotomy of creationists and evolutionists. The two groups not only disagreed in their explanation of animal structure, but also held divergent views on the relation of the history of life and the history of the earth. The teleologists simply assumed that changes in

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the earth's environment had necessitated the introduction of new species adapted to the new conditions. Those who rejected the sufficiency of teleological explanation could also reject the close parallel between the succession of fossils and the history of the earth, and instead attempt to discover laws or patterns of organic succession that were independent of geological changes. Owen, for example, believed that organic succession mirrored stages of embryological development. In the two decades before 1859, Ospovat argues, the writings of the nonteleologists represented the most stimulating and progressive current in biological thought, while the works of the teleologists by that time had little new to offer.99 It was from the works of the former group that Darwin borrowed theoretical concepts which he translated into evolutionary terms. In so doing, Darwin came to modify his initial conception of evolution by natural selection.

The Debate and Evolution in Britain and in France

Despite the innovative syntheses of such naturalists as Milne Edwards and Owen, homologies and adaptations were not satisfactorily reconciled until the advent of Darwin's theory of evolution in 1859. On the basis of the creationist hypothesis, at best an uneasy compromise could be reached, involving obscure tendencies of nature or mysterious polarizing and adaptive forces. Only on the assumption of descent with modification did the complex combination of unity of plan and adaptive variety fall into place. Darwin became familiar with the debate early in his career and quickly appreciated the importance of homologies for demonstrating evolution. Not long after beginning his first species notebook in 1837, Darwin read Principes de philosophie zoologique. He recognized the difficulties of linking mollusks and vertebrates by homologies, but pointed to "those beautiful passages from one to other organ" revealed by Milne Edwards's work on the mouthparts of Crustacea. Darwin saw even greater difficulties in Cuvier's position: "Cuvier has said each animal made for itself does not agree with old and modern types being constant." In other words, if each animal were really created for itself alone, there would be no reason for the animals of successive creations to share a common plan. Though Darwin thought Geoffroy's unity of composition was not workable, he was immediately sympathetic to the underlying assumption that nature worked by universal laws: "H[ilaire]. says grand idea God giving laws and on them leaving all to follow consequences."100 In 1837 Darwin also read and annotated Whewell's account of the debate in his History of the Inductive Sciences. To Whewell's defense of the utilitarian argument from design, Darwin responded in the margin by calling attention to mammae in males.101 From his earliest thinking about evolution, Darwin could call upon the new morphology with its appeal to law to counter the old reliance on God's free choice in assigning means to ends. If Ospovat is correct, Darwin only slowly divested himself of teleological

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reasoning. Between 1844 and 1859, Darwin substantially modified his theory. In the "Sketch" of 1842 and the "Essay" of 1844, he still thought the laws of evolution led to an overall harmony in nature. Although Darwin rejected teleological explanation of the school of Paley, he still retained a belief in perfect adaptation within the limits posed by unity of plan. Long after becoming an evolutionist he continued to believe in a teleology of a higher order. He thought of his 1840s theory of evolution as carrying on the goal of Owen's theories, that of reconciling teleology and morphology.102 Even in the Origin, there was a reminder of this. Darwin wrote: It is generally acknowledged that all organic beings have been formed on two great laws—Unity of Type, and the Conditions of Existence. . . . On my theory, unity of type is explained by unity of descent. The expression of conditions of existence, so often insisted on by the illustrious Cuvier, is fully embraced by the principle of natural selection.103

Ospovat sees the work of von Baer, Milne Edwards, and Owen, the latter two very much influenced by the debate, as of great importance to the development of Darwin's final theory of evolution. Darwin quickly adopted Owen's terminology. For Darwin, ancestors were archetypes, and evolution had to explain the transformation of a homogeneous unspecialized organism into a variety of specialized descendants. His 1840s theory assumed that transmutation was an intermittent process, occurring only in response to a change of external conditions. Once animals were readapted, evolution ceased. Only in the 1850s, through working out the consequences of the branching conception of nature, and especially the parallel between embryonic divergence and affinities in adults, did Darwin finally abandon the last remnant of teleological thinking. His effort at rethinking resulted in the principle of divergence, whereby the offspring of each species try to occupy as many diverse niches in the economy of nature as possible. In this more ecological understanding of nature, Darwin posited ample variation, a continuous struggle for existence, and organisms that were only relatively adapted to their surroundings.104 In the Origin Darwin reserved the evidence from morphology for the chapter just preceding the recapitulation and conclusion of the book. The subject of "Morphology," Darwin avowed, was "the most interesting department of natural history, and may be said to be its very soul." He marshaled example upon example in vertebrates, in articulates, and in plants, of structures employed for the most divergent of functions, but composed of homologous elements. Geoffroy was cited as an authority on the "high importance of relative connexion in homologous organs." Such evidence of constancy of plan, Darwin argued, citing Owen, could not be accounted for "by utility or by the doctrine of final causes." Darwin found equally inexplicable by the theory of special creation the many examples in nature of serial homologies. Nor could the creationist hypothesis explain why embryological characters were the most valuable characters in classification, why embryos of widely different animals of the same class were often similar, or why rudimentary organs were so common.105

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All of these morphological relationships could be readily explained by the theory of descent with modification. Modification by natural selection would alter the size and form of the skeletal pieces, but would have little tendency to alter the relative connection of the parts. "If we suppose that the ancient progenitor, the archetype as it may be called, of all mammals, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the whole class."106 In the case of serial homologies, Darwin argued that parts that were many times repeated were especially liable to vary, and would provide a fund of materials which could be adapted to the most varied of purposes while still retaining traces of their original similarity.107 To account for the embryological parallels, Darwin hypothesized that natural selection might affect the adult form of the animal while making little change in the protected embryonic form. If that were so, embryos of widely differing members of the same class might appear similar. The embryo would reveal to the naturalist the structure of the ancient and least modified progenitor of the class. Darwin attributed rudimentary organs to disuse and the tendency of nature to economize on the materials she uses. Thus, all the data of philosophical anatomy could be reinterpreted in terms of Darwin's theory of evolution. Philosophical anatomy, with its vision of an ideal unity of structure, did not disappear in Britain and America after Darwin. While belief in an archetype may have been difficult to reconcile with belief in natural selection and the struggle for existence, most naturalists preferred other theories of evolution that allowed for some measure of design in nature. Such theories were quite compatible with a belief in ideal forms. Thus, despite their conversion to evolution, many naturalists continued to interpret homologies and symmetries in nature not only as evidence of evolution but also as evidence of the Creator's plan. Owen himself adopted a teleological form of evolution and continued to publish his drawing and description of the Archetype. Jeffries Wyman, an American student of Owen, wrote within the framework of the Archetype and the polarizing and adaptive forces long after becoming an evolutionist.108 The term "philosophical anatomy" was replaced by "morphology," and morphological studies, often in a teleological context, continued unabated after Darwin. In fact, the late nineteenth century was the heyday of morphology. Although after Darwin homologies implied descent from a common ancestor, the method of establishing homologies was unchanged. In practice, naturalists formulated homologies in the same manner as before evolution, by tracing connections in adults and especially in embryos.109 In England the Cuvier-Geoffroy debate contributed to the development and relatively rapid acceptance of Darwin's theory; in France, where evolution met with widespread resistance, it played a different role as a historical precedent of a current controversy. The passions aroused by the Cuvier-Geoffroy debate had all but disappeared from French zoology when they were suddenly reawakened by the intrusion of Darwinian evolution. Reinterpreted as a battle over evolution, the debate became one of the focal points of the scientific and reli-

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gious controversy sparked by Darwin. Both sides on the issue turned to the history of French science to lend support to their claims. Opponents of evolution argued that the debate had long ago settled the issue. Proponents later in the century reinstated Lamarck to prominence, and tended to interpret the debate as the culmination of a longstanding war between the true founder of evolution and the reactionary chef d'ecole, Cuvier. Historians have sometimes expressed surprise that the French were so slow to accept evolution, but from the perspective of the Cuvier-Geoffroy debate, one would be astonished had it been otherwise.110 To read the arguments brought forth against Darwin in France is to be overwhelmed by a feeling of deja vu. Once more the entire arsenal of Cuvierian tactics was brought to bear: the conspiracy of silence, the sarcasm and ridicule, the charges of lack of novelty, and the confounding of Darwin's theory with Lamarckian transformism, spontaneous generation, and a host of other discredited speculative theories of the past. Flourens, for example, accused Darwin of following an a priori system, an anathema in France, while he followed only the facts. And Quatrefages, in typical Cuvierian fashion, rhetorically separated the "two Darwins," Darwin the patient observer and Darwin the rash theorizer. In this climate, it was small wonder that no accredited mainstream French scientist would translate Darwin and that the task was instead undertaken by a female republican propagandist, Clemence Royer, who added an inflammatory anti-clerical introduction.111 Nor was it any surprise that Darwin failed several times to be elected a corresponding member of the section of anatomy and zoology of the Academie, and that when he finally gained admission to the Academie in 1878 it was in the section of botany. In this way, the academicians could leave it ambiguous which of the "two Darwins" they were honoring. It was not long after 1859 that the interpretation of the Cuvier-Geoffroy debate became enmeshed in the controversy generated by the Origin. In 1864 Flourens wrote a series of five articles on the debate in the widely read Journal des savants. An unsuspecting reader might have been confused by Flourens's sudden hostility toward Geoffroy. Why did Flourens choose to rehash the events of the confrontation between Cuvier and Geoffroy at this late date and pronounce Cuvier the victor? Only in the last paragraph of the first article did Flourens's motive for writing become clear. Whereas in his eloge of Geoffroy Flourens also interpreted the debate as facts versus ideas, here he did not. The sole issue raised by the conflict was one plan of nature or four. Though Cuvier had defeated once and for all Geoffroy's attempt to demonstrate unity of composition in the animal kingdom, the issue had been recently raised again: The human mind always finds a certain pleasure in exercising itself against good sense. We all know, for example, and from sure science, that species do not change . . . we all know that, and yet a man arrives . . . a M. Darwin, who tells us that species do change . . . And already I see a certain public, at first astounded, then stupefied, and then not at all vexed to be furnished a new occasion to exercise itself against good sense, and not think like everyone else. . . . a man comes who repeats with assurance that all animals are composed alike, that there is unity of composition and unity of plan.112

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Thus Flourens sought to assimilate Darwin to Geoffroy and portray him as another misguided advocate of unity of composition. Unlike later writers, Flourens avoided discussion of Geoffroy's own theory of evolution. It was not necessary for Flourens's argument to suggest that Geoffroy was a transformist and that the debate concerned evolution. Rather, Flourens based his primary analogy between Geoffroy and Darwin not on the similarity of their conclusions but rather on methodological grounds. The two men had committed similar methodological sins by failing to see the limits of their respective theories. "M. Geoffroy deceives himself as M. Darwin has deceived himself," wrote Flourens. Darwin saw variability of species but failed to recognize the limits of variability, and so was led to "the transformation of the species." Similarly, Geoffroy had discovered homologies, but not seeing the limits of homologies, concluded that there existed unity of composition: These two incomplete minds have failed to turn everything topsy-turvy, and for the same cause—because they both lacked the superior and firm sense which sees the limits of phenomena, limits pre-estahlished and fixed, supreme guardians of the eternal distinction of species and organs.113

Like his mentor Cuvier, Flourens countered theories he opposed by insisting on limiting the applicability of generalizations to so-called observed phenomena. Under the circumstances, Milne Edwards and his disciple Armand de Quatrefages gave Darwin a much fairer hearing than did Flourens. At least they took him seriously, and though they disagreed with him, they supported his election as a corresponding member of the Academie. In Quatrefages's history of the French precursors of Darwin, first published in 1870, Geoffroy appeared as a transformist, chief among those who believed in a sudden transmutation of species, and the Cuvier-Geoffroy debate was reinterpreted in terms of the evolutionary debate. In fact, according to Quatrefages, the debate before the Academie began not with the memoir of Meyranx and Laurencet in February 1830, but rather with Roulin's memoir of 1828 on variability in domestic animals returned to the wild.114 It was this paper that induced Geoffroy to write his 1828 paper on fossil and living crocodiles. But for all his concern to be just, Quatrefages's conclusion might equally well have been written by Cuvier: Here is where we stand with respect to living organisms. We study them already made. We have not yet been able to penetrate into the workshop from whence they came; thus we can say nothing concerning the process of formation. . . . But if we truly wish to prepare for the future, let us learn to repress our ardor and our impatience. Let us use with gratitude the store of positive knowledge amassed by our predecessors. Let us increase it with our own labors, and let us guard ourselves against sacrificing it to hypotheses under the pretext of progress.115

By the 1880s, when transformism finally made headway among those in authority in French science, the Cuvier-Geoffroy debate was reinterpreted once more, this time as a central battle in the power struggle for the acceptance of evolution. Edmond Perrier, a professor and later director of the Museum and an early propagandist for evolution in France, devoted a chapter of his 1884

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review of French zoological philosophy before Darwin to the debate.116 Yet Geoffroy, instead of emerging as the ultimate victor, receded into the background as the debate came to be seen primarily as a confrontation not between Cuvier and Geoffroy, but between Cuvier and Lamarck. Lamarck was resurrected as "la fondateur de la doctrine d'evolution" a dedicated searcher for truth whose brilliant intuition had been stifled by the powerful Cuvier.117 In 1908 Lamarck's statue was placed at the entrance to the Museum, where it now stands across the expanse of formal gardens from the old statue of Buffon, which Geoffroy had so ardently defended in the 1830s. Geoffroy himself was largely forgotten. Then, in the early twentieth century, with evolution (as opposed to natural selection) no longer seriously contested among scientists, historians and biologists reread the transcript of the debate to discover that evolution was at most a secondary issue. Once again, the significance of the debate was revised in terms which reflected the tension in contemporary biology between morphology and functionalism. This new interpretation was put forward with particular emphasis by the twentieth-century British zoologist Edward Stuart Russell. A Lamarckian and opponent of materialistic and mechanistic explanations in biology, Russell advocated a return to teleological reasoning within the context of evolution. For Russell, what united Lamarck and Cuvier was more important than what divided them; both believed in the purposiveness of nature. He saw the debate as contrasting a common-sense functionalist view of nature with an abstract transcendental view. Although he recognized the importance of homologies, he castigated Geoffroy for his misguided effort to create a "pure morphology." A pure morphology was not possible, as Geoffroy himself unwittingly proved by allowing functional considerations to infiltrate into his work. Cuvier and Geoffroy emerged in Russell's classic work Form and Function (1916) as the greatest champions in history of two divergent strands of thought reaching back to Plato and Aristotle. Cuvier won the debate not by convincing his audience that species were fixed, but by demonstrating the necessity for teleological explanation in biology.118 More recent historians, heirs of the twentieth-century synthesis of Darwinian evolution, Mendelian genetics, and population dynamics, no longer feel the need to decide between Cuvier and Geoffroy. While generally following Russell and interpreting the debate as a contest between a teleological and a morphological approach to nature, they have portrayed the two views as more nearly complementary, although they have continued to believe that Cuvier had the upper hand in the argument. From the perspective of twentieth-century biology, neither Cuvier nor Geoffroy ultimately triumphed, for both are equally necessary to our modern understanding of biology. While agreeing in the main with Russell and later historians, we, taking as our point of reference the early nineteenth-century French context rather than the post-1859 British context, have given greater emphasis than has previously been given to Geoffroy and his contributions to the development of morphology. Using French rather than British terms, we may characterize the main intellectual issue of the debate as the "conditions of existence" versus philo-

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sophical anatomy. But the two rivals did not play equal but complementary roles. As contemporaries were well aware, the debate had much more to do with Geoffroy than with Cuvier; in this contest, Geoffroy was the challenger and Cuvier the defender. The debate provided an opportunity to test the originality and the scope of the theories and methods of philosophical anatomy. And despite Cuvier's skill in logical argument, Geoffroy did in the end succeed in convincing most of his contemporaries of the novelty and importance of homologies in comparative anatomy. Although Geoffroy's dream of uniting all animal life under a single principle—unity of organic composition—was quickly abandoned after 1830, the insights and discoveries of philosophical anatomy became a valued part of pre-Darwinian French and British zoology, and were afterwards incorporated into modern evolutionary biology. In its heyday in the early nineteenth century, philosophical anatomy, despite its occasional excesses, proved extraordinarily fruitful for anatomy and zoology. The search for transcendental patterns in nature stimulated extensive research and attracted new students to these fields. In invertebrate zoology, the concept of homology played an essential role in unraveling the classes, orders, genera, and species of the mollusks, articulates, and radiates. In vertebrate zoology, philosophical anatomy led for the first time to a minute study of individual bones, processes of bones, connecting elements, and centers of ossification. A uniform terminology was adopted which was extended from man as far as practicable through the vertebrate series. Philosophical anatomy encouraged the study of teratology because it suggested that malformations could give insight into relationships among normal animals. It gave special impetus to the study of embryology, because it was through careful observation of the stages of development that affinities could best be demonstrated. Finally, the realization that there existed structural affinities that could not be explained on the basis of functional similarity led to an undermining of traditional teleology, and so prepared the way for the Darwinian Revolution. The Cuvier-Geoffroy debate, like all celebrated scientific controversies, involved much more than the truth or falsity of particular scientific theories. It depended upon, and reinforced, broader religious, political, and social controversies, and it was used to feed the flames of opposition to professional science and its elitist system of institutions. It concerned personalities, style, and politics as well as science. One cannot say once and for all what the debate "really" meant in 1830 and afterwards, for it meant and it means something different to each of its participants and interpreters. Its richness is perhaps its greatest legacy. For Cuvier it offered a public forum to insist that morphological resemblances were subordinate to the "conditions of existence" and to rail against materialistic generalizations in science. For Geoffroy it was an opportunity to vindicate unity of composition and an entire movement of synthetic philosophy. For Flourens it was a question of one plan of animal structure versus four entirely separate ones, and for Isidore Geoffroy a confrontation between a school of facts and a school of ideas. For Raspail it was above all another example of dirty scientific politics, and for Quinet it foreshadowed the discovery of a grand unity in society. For Quatrefages it was a battle over trans-

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formism that Cuvier had long ago won. For Perrier it was a battle over transformism in which Lamarck ultimately triumphed. And for Russell it was the epitome of an ancient and still ongoing conflict between functionalism and morphology. It is both misleading and useless to judge such claims as either just or mistaken. In the end, a confrontation between two great men becomes a symbol, which like all symbols is inexhaustible. Each age can reinterpret it in terms of its own problems and find new meaning in it.

APPENDIX A

Chairs at the Museum d'Histoire Naturelle Related to Zoology (1793-1870)

Quadrupeds, Cetaceans, Reptiles, Birds, and Fishes

1793-1794 Etienne Geoffroy Saint-Hilaire (1772-1844) Chair split into the two following chairs in 1794: Mammals and Birds

1794-1841 Etienne Geoffroy Saint-Hilaire (1772-1844) 1841-1861 Isidore Geoffroy Saint-Hilaire (1805-1861) 1862-1876 Henri Milne Edwards (1800-1885) Reptiles and Fishes

1794-1825 Bernard de Lacepede (1756-1825) 1825-1857 Constant Dumeril (1774-1860) 1857-1870 Auguste Dumeril (1812-1870) Insects, Worms, Microscopic Animals, etc.

1793-1829 Jean-Baptiste Lamarck (1744-1829) Chair split into the two following chairs in 1830: Crustaceans, Arachnids, and Insects

1830-1833 Pierre-Andre Latreille (1762-1833) 1833-1841 Jean-Victor Audouin (1797-1841) 1841-1861 Henri Milne Edwards (1800-1885) (switched to chair of mammalogy) 1862-1895 Emile Blanchard (1819-1900) Mollusks, Worms, and Zoophytes

1830-1832 Henri Ducrotay de Blainville (1777-1850) (switched to chair of comparative anatomy) 1832-1865 Achille Valenciennes (1794-1865) 238

APPENDIX

239

1865-1869 Henri de Lacaze-Duthiers (1821-1901) (left chair for a chair of zoology at the Faculte des Sciences) 1869-1875 Gerard Paul Deshayes (1795-1875) Anatomy of Animals

1793-1802 A.-L.-F. Mertrud (d. 1802) Name of chair changed in 1802 to:

Comparative Anatomy

1802-1832 1832-1850 1850-1855 1855-1868 1868-1879

Georges Cuvier (1769-1832) Henri Ducrotay de Blainville (1777-1850) Georges Duvernoy (1777-1855) E. R. A. Serres (1786-1868) Paul Gervais (1816-1879)

Anatomy of Man

1793-1832 Antoine Portal (1742-1832) 1832-1838 Pierre Flourens (1794-1867) Name of chair changed in 1838 to:

Anatomy and Natural History of Man

1839-1855 E. R. A. Serres (1786-1868) (switched to chair of comparative anatomy) Name of chair changed in 1855 to:

Anthropology

1855-1892 Armand de Quatrefages (1810-1892)

Comparative Physiology

1837-1838 Frederic Cuvier (1773-1838) 1838-1867 Pierre Flourens (1794-1867) 1868 Paul Bert (1833-1886) (as suppleant) In December 1868, this chair was transferred to the Faculte des Sciences and Paul Bert made titulary. At the same time, Claude Bernard's chair of physiology at the Faculte des Sciences was transferred to the Museum.

240

APPENDIX

General Physiology

1868-1878 Claude Bernard (1813-1878) Paleontology

1853-1857 Alcide d'Orbigny (1802-1857) 1861-1868 Adolphe d'Archiac (1802-1868) 1869-1871 Edouard Lartet (1801-1871)

Compiled from various sources including Museum National d'Histoire Naturelle, Centenaire de la fondation du Museum d'Histoire Naturelle, lO juin 1793-10 juin 1893 (Paris, 1893), pp. iii-vi; and a list compiled for the Museum project at the Institut d'histoire et de sociopolitique des sciences, Universite de Montreal.

APPENDIX B

Chairs at the College de France Related to Zoology (1778-1870) Natural History

1778-1800 L.-J.-M. Daubenton (1716-1800) 1800-1832 Georges Cuvier (1769-1832) 1832-1837 Leonce Elie de Beaumont (1798-1874) In 1837 this chair was split into a chair of Natural History of Inorganic Bodies, given to Elie de Beaumont who held it until 1874, and a chair of the Natural History of Organized Bodies. Natural History of Organized Bodies

1837-1855 Georges Duvernoy (1777-1855) 1855-1867 Pierre Flourens (1794-1867) 1869-1903 Jules Marey (1830-1904) Comparative Embryology

1844-1873 Victor Coste (1807-1873) Anatomy

1773-1832 Antoine Portal (1742-1832) Medicine (chair created in 1769)

1796-1804 1804-1822 1822-1826 1827-1830 1830-1855 1855-1878

Jean-Nicholas Corvisart (1755-1821) Noel Halle (1754-1822) Theophile Laennec (1781-1826) Joseph Recamier (1774-1852) Francois Magendie (1783-1855) Claude Bernard (1813-1878)

Compiled from Le College de France (1530-1930) (Paris, 1932). 241

APPENDIX C

Chairs at the Faculte des Sciences in Zoology (1808-1870)

There were two chairs of zoology, a titulary chair and an adjunct. The senior man always became the titulary professor. i.

1808-1844 Etienne Geoffroy Saint-Hilaire (1772-1844) 1844-1885 Henri Milne Edwards (1800-1885) II.

1809 1812-1850 1850-1861 1861-1865 1865-1869 1869-1901

Georges Duvernoy (1777-1855) Henri Ducrotay de Blainville (1777-1850) Isidore Geoffroy Saint-Hilaire (1805-1861) Louis Pierre Gratiolet (1815-1865) Paul Gervais (1816-1879) Henri de Lacaze-Duthiers (1821-1901)

Compiled from the annual government Almanach and from individual biographies. 242

APPENDIX D

Zoologists at the Academie des Sciences (1795-1870)

Members of the Section of Anatomy and Zoology

The roman numerals correspond to the six seats of the section. [ ] Age at entrance to the Academie des Sciences of the ancien regime; ( ) age at entrance to the Institut de France.

i. 1795 1800 1814 1833 1862

L.-J.-M. Daubenton [27] (79) Antoine Olivier (44) Pierre-Andre Latreille (51) Isidore Geoffroy Saint-Hilaire (27) Emile Blanchard (42)

ii. 1795 Bernard de Lacepede (38) 1825 Henri de Blainville (48) 1851 Victor Coste (43) III.

1795 Jacques Tenon [35] (71) 1816 Constant Dumeril (42) 1860 Francois-Achille Longet (49) IV.

1795 1807 1844 1866

Auguste Broussonet [24] (34) Etienne Geoffroy Saint-Hilaire (35) Achille Valenciennes (50) Charles Robin (44) 243

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V.

1795 1803 1826 1838

Georges Cuvier (26) Philippe Pinel (57) Frederic Cuvier (53) Henri Milne Edwards (38)

VI.

1795 L.-C.-M. Richard (41) 1821 Jules-Cesar de Savigny (44) 1852 Armand de Quatrefages (42) Zoologists who entered the Academie des Sciences in other sections:

1795 1806 1828 1828 1831 1838 1847

Jean-Baptiste Lamarck, Botany [34] (51) Louis Bosc, Rural Economy (47) Pierre Flourens, Rural Economy (47) E. R. A. Serres, Medicine and Surgery (41) Henri Dutrochet, Rural Economy (54) Victor Audouin, Rural Economy (40) Georges Duvernoy, Free Academician (69)

Permanent Secretaries of the Academie des Sciences (1803-1870): Sciences mathematiques

1803 1822 ] 830 1853

J.-B.-J. Delambre Joseph Fourier Francois Arago Leonce Elie de Beaumont

Sciences physiques

1803 1832 1833 1868

Georges Cuvier Pierre-Louis Dulong Pierre Flourens Jean-Baptiste Dumas

Compiled from Le Comte de Franqueville, Le Premier Siecle de I'Institut de France (Paris, 1895), and Academie des Sciences, Index biographique des membres et correspondants de l'Academie des Sciences de 1666 d 1939 (Paris, 1939).

Notes

Chapter 1

1. Johann Peter Eckermann, Gesprache mit Goethe in der letzten Jahre seines Lebens, 2 vols. (Leipzig: Eugen Diederichs, 1902) [original edition, 3 vols., 1836-1848], 2: 473-474; idem, Conversations of Goethe with Eckermann and Soret, trans. John Oxenford (London, 1874), pp. 479480. My translation is largely based on Oxenford's. Soret's original notes on which Eckermann's account was based are in Frederic Soret, Conversations avec Goethe, ed. A. Robinet de Clery (Paris: Editions Montaigne, 1932), pp. 121-122. 2. Examples of historical works on biological controversies include John Farley and Gerald L. Geison, "Science, Politics and Spontaneous Generation in Nineteenth-Century France: The Pasteur-Pouchet Debate," Bulletin of the History of Medicine 48 (1974): 161-198; Shirley Roe, Matter, Life, and Generation: Eighteenth-Century Embryology and the Haller-Wolff Debate (Cambridge: Cambridge University Press, 1981); Philip J. Pauly, "The Loeb-Jennings Debate and the Science of Animal Behavior," Journal of the History of the Behavioral Sciences 17 (1981): 505-515; William B. Provine, The Origins of Theoretical Population Genetics (Chicago: University of Chicago Press, 1971) (an intellectual account of the biometry-Mendelian controversy); and Donald Mackenzie and Barry Barnes, "Scientific Judgement: The Biometry-Mendelian Controversy," in Barry Barnes and Steven Shapin, eds., Natural Order: Historical Studies of Scientific Culture (Beverly Hills: Sage, 1979) (a sociological account of this controversy). Since this manuscript was completed, two important works on scientific controversies have appeared: Martin J.S. Rudwick, The Great Devonian Controversy: The Shaping of Scientific Knowledge Among Gentlemanly Specialists (Chicago: University of Chicago Press, 1985); and James Secord, Controversy in Victorian Geology: The Cambrian-Silurian Dispute (Princeton: Princeton University Press, 1986). For a review of the literature on scientific controversy from a sociological perspective, see Steven Shapin, "History of Science and its Sociological Reconstructions," History of Science 20 (1982): 157-211. Most of the authors on scientific controversies, including the present author, have benefited greatly from Thomas S. Kuhn, The Structure of Scientific Revolutions, 2nd ed. (Chicago: University of Chicago Press, 1970). I have found helpful some of the writings of my former colleague at the University of Ottawa, Andrew Lugg. See, for example, "Disagreement in Science," Zeitschrift fur allgemeine Wissenschaftstheorie 9 (l978): 276-292. 3. See the following of Bourdier's writings: "Etienne Geoffroy Saint-Hilaire," in Charles Coulston Gillispie, ed., Dictionary of Scientific Biography, 16 vols. (New York: Charles Scribner's Sons, 1970-1980) (hereafter, DSB), 5: 355-358; "Georges Cuvier," ibid., 3: 521-528; "Geoffroy SaintHilaire Versus Cuvier: The Campaign for Paleontological Evolution (1825-1838)," in Cecil J. Schneer, ed., Toward a History of Geology (Cambridge, Mass.: M.I.T. Press, 1969), pp. 36-61; and "Trois siecles d'hypotheses sur 1'origine et la transformation des etres vivants (1550-1859)," Revue d'histoire des sciences 13 (1960): 1-44. The evolutionary component of the debate is also emphasized by Pietro Corsi, Oltre il mito: Lamarck e le scienze naturali del suo tempo (Bologna: il Mulino, 1983). 245

246

NOTES

4. E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology (London: John Murray, 1916), reprint ed. (Chicago: University of Chicago Press, 1982), pp. v, 31-44, 52-78, quotations on pp. 65, 78. A similar interpretation of the debate as a conflict between a morphological and a teleological or physiological view of animal organization is taken by William Coleman, Georges Cuvier, Zoologist: A Study in the History of Evolution Theory (Cambridge, Mass.: Harvard University Press, 1964), pp. 43, 152-153; Jean Piveteau, "Les discussions entre Cuvier et Geoffroy Saint-Hilaire sur 1'unite de composition du regne animal," Revue d'histoire des sciences 3 (1950): 343-363; and Francois Jacob, The Logic of Life: A History of Heredity, trans. Betty E. Spillman (New York: Pantheon, 1982), pp. 100-111. Theophile Cahn, La vie et I'oeuvre d'Etienne Geoffroy Saint-Hilaire (Paris: Presses Universitaires de France, 1962) evidently also believes that the debate concerned evolution, for he begins his chapter on the debate with an account of Geoffroy's evolutionary doctrine. In the remainder of the chapter, he seems to imply that the main issue was whether or not one could discover a system of laws of organization analogous to laws in the physical sciences. 5. See, for example, Edward Lurie, Louis Agassiz: A Life in Science (Chicago: University of Chicago Press, 1960), pp. 58-63. Rehbock (note 7) also tends to conflate Geoffroy and German Naturphilosophie. This intepretation of Geoffroy was especially prevalent in J.H.F. Kohlbrugge, Historische-kritische Studien uber Goethe als Naturforscher (Wurzberg: Kurt Kabitzsch, 1913), pp. 6163, who treated Geoffroy as an offshoot of fantastical Naturphilosophie challenging the modern positivistic science of Cuvier. Geoffroy was defended against this charge by Wilhelm Lubosch, "Der Akademiestreit zwischen Geoffroy St.-Hilaire und Cuvier im Jahre 1830 und seine leitenden Gedanken," Biologisches Zentralblatt 38 (1918): 357-384, 397-456. Lubosch emphasized Geoffroy's role as the founder of the concept of homology. 6. Russell, Form and Function, p. 65. According to Russell, Cuvier was able to subject Geoffroy's arguments to "destructive, and indeed final criticism" (ibid., p. 74). See also Piveteau, "Les discussions entre Cuvier et Geoffroy," p. 362. 7. Recent works that have appreciated the importance of morphology in the pre-Darwinian era include Dov Ospovat, "Perfect Adaptation and Teleological Explanation: Approaches to the Problem of the History of Life in the Mid-nineteenth Century," in Studies in History of Biology, vol. 2, ed. William Coleman and Camille Limoges (Baltimore: Johns Hopkins University Press, 1978), pp. 33-56; idem, The Development of Darwin's Theory: Natural History, Natural Theology, and Natural Selection, 1838-1859 (Cambridge: Cambridge University Press, 1981); Michael Ruse, The Darwinian Revolution: Science Red in Tooth and Claw (Chicago: University of Chicago Press, 1979); and Philip F. Rehbock, The Philosophical Naturalists: Themes in Early Nineteenth-Century British Biology (Madison: University of Wisconsin Press, 1983). 8. This is a major theme of Ospovat, Darwin's Theory. 9. Richard W. Burkhardt, Jr., "Lamarck, Evolution, and the Politics of Science," Journal of the History of Biology 3 (1970): 275-298; idem, The Spirit of System: Lamarck and Evolutionary Biology (Cambridge, Mass.: Harvard University Press, 1977), pp. 186-218. Chapter 2

1. Quoted in Isidore Geoffroy Saint-Hilaire, Vie, travaux et doctrine scientifique d'Etienne Geoffroy Saint-Hilaire (Paris, 1847); reprint ed. (Brussels: Culture et Civilisation, 1968), p. 25. 2. See, for example, J.P.F. Deleuze, Histoire et description du Museum Royal d'Histoire Naturelle, 2 vols. (Paris, 1823), esp. 1: 145; and Henri Daudin, Cuvier et Lamarck: les classes zoologiques et I'idee de serie animal, 1790-1830, 2 vols. in 1 (Paris: Felix Alcan, 1926), chapter 2. 3. Daudin, Cuvier et Lamarck, "Avant-propos." 4. On the popularity of natural history in the eighteenth century, see Daniel Mornet, Les sciences de la nature en France, au XVIII e siecle (Paris: Armand Colin, 1911), esp. pp. 1-2; and Yves Laissus, "Les cabinets d'histoire naturelle," in Rene Taton, ed., Enseignement et diffusion des sciences en France au XVII e siecle (Paris: Hermann, 1964), pp. 659-712. 5. Paul Lawrence Farber, "The Development of Taxonomy and the History of Ornithology," Isis 68 (1977): 550-566.

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6. On the variety of approaches to the life sciences in the eighteenth century, see Jacques Roger, "The Living World," in G.S. Rousseau and Roy Porter, eds., The Ferment of Knowledge: Studies in the Historiography of Eighteenth-Century Science (Cambridge: Cambridge University Press, 1980): 255-283. On the biological doctrines of Buffon and the philosophes, see Jacques Roger, Les sciences de la vie dans la pensee francaise du XV111e siecle: la generation des animaux de Descartes a I'Encyclopedie (Paris: Armand Colin, 1963). For an overview of personalities, institutions, and ideas in late eighteenth-century French science, see Charles Coulston Gillispie, Science and Polity in France at the End of the Old Regime (Princeton: Princeton University Press, 1980). 7. Paul Lawrence Farber, "Buffon and the Concept of Species," Journal of the History of Biology 5 (1972): 259-284; Phillip R. Sloan, "Buffon, German Biology, and the Historical Interpretation of Biological Species," British Journal for the History of Science 12 (1979): 109-153; Phillip R. Sloan and John Lyon, "Introduction" in Lyon and Sloan, eds., From Natural History to the History of Nature: Readings from Buffon and His Critics (Notre Dame, Ind.: University of Notre Dame Press, 1981), pp. 1-32; and Roger, "Living World," p. 278. 8. Toby A. Appel, "B.-G.-E. de la Ville-sur-1llon, comte de Lacepede," DSB, 1: 546-548. 9. The Encyclopedie methodique, edited by C.J. Pancoucke, 193 vols. (Paris, 1782-1830) was to be the sequel to the grand Encyclopedie of Diderot and D'Alembert. The work was interrupted by the Revolution and completed mainly during the Restoration. Daubenton, the editor of the zoology volumes, wrote the sections on mammals, reptiles, and fishes, Olivier began the volumes on "insects," and Bruguiere those on "worms." Encyclopedie methodique. Histoire naturelle des animaux, ed. L.-J.-M. Daubenton, 10 vols. (Paris, 1782-1825); Encyclopedie methodique. Histoire naturelle des vers, 2 vols. (1789-1830). Vicq d'Azyr contributed a volume on comparative anatomy: Encyclopedie methodique. Systeme anatomique des quadrupedes, vol. 2 (Paris, 1792). 10. On the French Linnaeans, see Frans A. Stafleu, Linnaeus and the Linnaeans: The Spreading of Their Ideas in Systematic Botany, 1735-1789 (Utrecht: International Association for Plant Taxonomy, 1971), pp. 267-291. See also J. Lamoureux, "Antoine Gouan," Nouvelle biographie generale, 46 vols. (Paris, 1852-1877) (hereafter Nouvelle biographie generate), 21: columns 360-363; Jean Motte, "P.-A.-M. Broussonet," DSB, 2: 509-511; Georges Cuvier, "Eloge historique de PierreMarie-Auguste Broussonet, lu le 4 Janvier 1808," in idem, Recueil des eloges historiques lus dans les seances publiques de I'Institut Royal de France, 3 vols. (Paris, 1819-1827), 1: 311-342; Jean Theodorides, "Jean-Guillaume Bruguiere (1749-1798) et Guillaume-Antoine Olivier (1756-1814), medecins, naturalistes et voyageurs," Congres National des Societes Savantes. Section des Sciences, Montpellier, 1961, Comptes rendus (Paris: Gauthier-Villars, 1962), pp. 173-183; Georges Cuvier, "Extrait d'une notice biographique sur Bruguieres, lue a la Societe philomatique dans sa seance generale du 30 Janvier 1799," in idem, Recueil des eloges historiques, 2: 425-442; idem, "Eloge historique de Guillaume-Antoine Olivier, lu le 8 Janvier 1816," in ibid., 2: 235-265. 11. On the natural system of classification in botany, see Stafleu, Linnaeus and the Linnaeans, pp. 321-332; and Henri Daudin, De Linne a Jussieu; methodes de la classification et idee de serie en botanique et en zoologie (1740-1790) (Paris: Felix Alcan, 1926). 12. Aubin-Louis Millin, "Discours sur 1'origine & les progres de 1'histoire naturelle en France," Actes de la Societe d'Histoire Naturelle de Paris 1 (1792): i-xvi, quotation on p. xii. On the Societe Linneene and the Societe d'Histoire Naturelle, see Daudin, Cuvier et Lamarck, pp. 6-11; Roger Hahn, The Anatomy of a Scientific Institution: The Paris Academy of Sciences, 1666-1803 (Berkeley: University of California Press, 1971), pp. 112-114, 179-182; Gillispie, Science and Polity, pp. 191-192. On Millin, see [Du Mersan] "Aubin-Louis Millin," Nouvelle biographie generale, 35: columns 537-541. 13. On the Jardin du Roi in the eighteenth century, see Yves Laissus, "Le Jardin du Roi," in Taton, ed., Enseignement et diffusion des sciences, pp. 287-341; and Gillispie, Science and Polity, pp. 143-184. 14. On Daubenton, see Georges Cuvier, "Eloge historique de Daubenton, lu le 5 avril 1800," in Recueil des eloges, 1: 37-80; and Camille Limoges, "L.-J.-M. Daubenton," DSB, 15: 111-114. The volumes on which Daubenton collaborated were: Georges-Louis Leclerc, comte de Buffon, Histoire naturelle generate et particuliere, avec la description du cabinet du roi. Quadrupedes, vols. 4-15 (Paris, 1753-1767). 15. Most of the following discussion of the transformation of the Jardin into the Museum is

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based upon E.T. Hamy, Les derniers jours du Jardin du Roi et la fondation du Museum d'Histoire Naturelle (Paris, 1893). This work, which reprints the major documents pertaining to the foundation of the Museum, also appears in Museum National d'Histoire Naturelle, Centenaire de la fondation du Museum d'Histoire Naturelle, lO juin 1793-10 juin 1893 (Paris, 1893), pp. 1-162. On the personnel of the Jardin, see Laissus, "Le Jardin du Roi." 16. Hamy, Derniers jours, p. 37n. 17. Ibid., pp. 97-98, 100. 18. Ibid., 106. 19. Ibid., pp. 65-66, 66n. For other accounts, see Deleuze, Histoire et description du Museum, 1: 72ff; and Paul Antoine Gratacap Cap, Le Museum d'Histoire Naturelle: histoire de la fondation et des developpements successifs de I'etablissement (Paris, 1854), p. 89ff. 20. See Charles Coulston Gillispie, "The Encyclopedie and the Jacobin Philosophy of Science: A Study in Ideas and Consequences," in Marshall Clagett, ed., Critical Problems in the History of Science (Madison: University of Wisconsin Press, 1959), pp. 255-289, esp. pp. 265-266; L. Pearce Williams, "The Politics of Science in the French Revolution," in ibid., pp. 291-308. 21. The Museum's budget rose considerably through the first part of the century. On the Museum's financial resources in the nineteenth century, see Camille Limoges, "The Development of the Museum d'Histoire Naturelle of Paris, c. 1800-1914," in Robert Fox and George Weisz, eds., The Organization of Science and Technology in France, 1808-1914 (Cambridge: Cambridge University Press, 1980), pp. 211-240, esp. pp. 217-219. 22. In the following discussion of Geoffroy's early career, I have generally followed Isidore Geoffroy Saint-Hilaire's biography of his father, Vie, travaux et doctrine scientifique d'Etienne Geoffroy Saint-Hilaire (note 1). This work, although obviously biased, is closest to the events and probably the most reliable for basic information. Theophile Cahn, La vie et I'oeuvre d'Etienne Geoffroy Saint-Hilaire (Paris: Presses Universitaires de France, 1962), the only other full-length biography, has some helpful additional material, but has no footnotes, an inadequate bibliography, and is insufficiently critical of its sources. Franck Bourdier, "Etienne Geoffroy Saint-Hilaire," DSB, 5: 355-358, is informative and provocative, as are all the writings of Bourdier, but also tendentious and at times inaccurate. Bourdier has on many occasions pointed the way to little-known but excellent sources on French science. Pierre Flourens, "Eloge historique d'Etienne Geoffroy SaintHilaire," Memoires de I'Academie des Sciences, 23 (1853): i-lxxi, also in idem, Recueil des eloges historiques lus dans les seances publiques de I'Academie des Sciences, 3 vols. (Paris, 1856-1862), 1: 229-284, is useful more for Flourens's opinions than for its account of Geoffroy's life. For a list of Geoffroy's works, see I. Geoffroy, op. cit, pp. 421-471; and Royal Society of London, Catalogue of Scientific Papers (1800-1863), 2: 824-832. Collections of Geoffroy's manuscripts may be found in the Bibliotheque Centrale du Museum National d'Histoire Naturelle, Archives de I'Academie des Sciences de 1'Institut de France, Archives Nationales de France, Wellcome Institute (London), American Philosophical Society (Philadelphia), and the Special Collections Branch, Smithsonian Institution Libraries, (Washington, D.C.). 23. Cahn, Geoffroy Saint-Hilaire, pp. 19-20. On Hairy, see R. Hooykaas, "Rene-Just Hauy," DSB, 6: 178-183. 24. Bourdier, "Geoffroy Saint-Hilaire," p. 355, claims without documentation that Geoffroy embraced revolutionary ideas and frequented radical committees and clubs. 25.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 11-19, quotation on p. 19. 26. Ibid., pp. 20-22. Correspondence regarding Lacepede's resignation and the appointment of Geoffroy is located in Archives Nationales de France, F17 1227, dossier 1, piece 3. Lacepede's letter of resignation is dated 4 March 1793 and the nomination of Geoffroy is dated 7 March. I am grateful to Toby Gelfand for researching this material for me. 27. Louis de Launay, Une grande famille de savants: les Brongniart (Paris: G. Rapilly et fils, 1940), pp. 49-59. Quotations, presumably taken from letters of Brongniart to his family, are on pp. 58, 59. 28. Cahn, Geoffroy Saint-Hilaire, p. 21. On Lakanal and Geoffroy, see Owsei Temkin and C. Lilian Temkin, "The Relationship between Geoffroy Saint-Hilaire and Lakanal," Bulletin of the History of Medicine 17 (1945): 305-314. 29.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 34-37, 39n.

NOTES

249

30. Ibid., pp. 41-42. 31. Cuvier, "Eloge historique de Daubenton," pp. 41-43; Johann Wolfgang von Goethe, "Derniers pages de Goethe expliquant a 1'Allemagne les sujets de philosophie naturelle controversees au sein de 1'Academie des Sciences de Paris," Revue encydopedique, 53 (1832): 563-573 and 54 (1832): 54-68, esp. 53: 564-568. For a recent analysis of the differences in style and doctrine of the two men, see Paul Lawrence Farber, "Buffon and Daubenton: Divergent Traditions Within the Histoire naturelle" Isis 66 (1975): 63-74. 32. Georges-Louis Leclerc, comte de Buffon, Histoire naturelle generale et particuliere, avec la description du cabinet du roi, vol. 1 (Paris, 1749), "Premier discours. De la maniere d'etudier & de traiter 1'Histoire Naturelle," pp. 3-62. Buffon's "Initial Discourse" has been recently translated into English by John Lyon in Lyon and Sloan, From Natural History to the History of Nature, pp. 89-128. 33. On Buffon's later philosophy of classification, see Farber, "Buffon and the Concept of Species"; Sloan, "Buffon, German Biology and the Historical Interpretation of Biological Species"; and Sloan and Lyon, "Introduction," in Lyon and Sloan, From Natural History to the History of Nature. 34. Georges-Louis Leclerc, comte de Buffon, Histoire naturelle generale et particuliere, avec la description du cabinet du roi, vol. 4 (Paris, 1753), "L'asne," pp. 377-403, quotations on pp. 379, 380, 381. 35. Cuvier, "Eloge historique de Daubenton," p. 58. 36. L.-J.-M. Daubenton, "Introduction a la histoire naturelle," in Encyclopedie methodique. Histoire naturelle des animaux, ed. L.-J.-M. Daubenton, vol. 1 (Paris, 1782), pp. i-xv; idem, "Sur la definition et les limites de 1'histoire naturelle," in Seances des ecoles normales, recueillies par des stenographes, et revues par les professeurs, new ed., 10 vols. (Paris, 1800-1801), 1: 95-112. On Daubenton's definition of the scope of natural history, see also Richard W. Burkhardt, Jr., The Spirit of System: Lamarck and Evolutionary Biology (Cambridge, Mass.: Harvard University Press, 1977), pp. 19-22. 37. Daubenton, "Introduction a la histoire naturelle," p. iv; idem, "Sur la nomenclature methodique de 1'histoire naturelle," in Seances des ecoles normales, 1: 425-444, quotation on p. 433, quoted and translated by Burkhardt, Spirit of System, p. 125. 38. Daubenton, "Sur la definition et les limites de 1'histoire naturelle," quotation on p. 112. 39. Daubenton, "Sur les voyages et les theories des naturalistes," in Seances des ecoles normales, 2: 171-188; idem, "Sur les rapports que 1'on a recherches entre les corps bruts et les corps organises," in ibid., 4: 3-13; and idem, "Sur la redaction de 1'histoire naturelle," in ibid., 1: 288-302, quotations on pp. 288, 290. 40. Etienne Geoffroy Saint-Hilaire, "Memoire sur un nouveau genre de quadrupedes de 1'ordre Glires," 12pp., Geoffroy Saint-Hilaire Papers, American Philosophical Society. The American Philosophical Society also holds two later manuscript versions of this memoir. On Daubenton's advice to Geoffroy, see I. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 43, 43n, 131, 131n. 41. Etienne Geoffroy Saint-Hilaire, "Extrait d'un memoire sur un nouveau genre de quadrupedes, de 1'ordre des Rongeurs (Glires L.)," La decade philosophique 4 (1795): 193-206. Daubentoniidae is now a family, although it still contains only a single species. 42. Etienne Geoffroy Saint-Hilaire, "Memoire sur les rapports naturels des Makis Lemur, L., et description d'une espece nouvelle de mammifere," Magasin encydopedique [2e annee] 1 (1796): 20-50, quotations on p. 20. 43. Ibid., p. 44. 44. For accounts of Cuvier's education and early career, see Dorinda Outram, Georges Cuvier: Vocation, Science and Authority in Post-Revolutionary France (Manchester: Manchester University Press, 1984); Georges Louis Duvernoy, Notice historique sur les ouwages et la vie de M. le Bon Cuvier (Paris, 1833); John Vienot, Le Napoleon de la intelligence: Georges Cuvier, 1769-1832 (Paris: Fischbacher, 1932); and especially Georges Cuvier, "Memoires pour celui qui fera mon eloge, ecrits au crayon dans ma voiture pendant mes courses de Paris en 1822 et 1823," Bibliotheque de 1'Institut de France, Fonds Flourens, Ms 2598 (3) (henceforth, Cuvier, Manuscript autobiography). This autobiography was partially published (omitting some of Cuvier's more critical comments about colleagues) as Georges Cuvier, "Memoires pour servir a celui qui fera mon

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eloge . . ." in Flourens, Recueil des eloges, 1: 169-193. Dates have been taken mainly from this latter source. I am grateful to William Coleman for making available to me his typed transcript of the manuscript version. The best accounts of Cuvier's scientific thought are William Coleman, Georges Cuvier, Zoologist: A Study in the History of Evolution Theory (Cambridge, Mass.: Harvard University Press, 1964); and Daudin, Cuvier et Lamarck. For additional biographical material on Cuvier, see below, Chapter 3, and the thorough bibliography of works about Cuvier by Dorinda Outram, "Scientific Biography and the Case of Georges Cuvier: With a Critical Bibliography," History of Science 14 (1976): 101-137. 45. Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 172. 46. Ibid., p. 174. This was probably an introduction to the works of Kielmeyer's later teacher, Johann Friedrich Blumenbach. On Blumenbach and Kielmeyer, see Timothy Lenoir, "Kant, Blumenbach, and Vital Materialism in German Biology," Isis 71 (1980): 77-108. 47. Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 173; Christoph Heinrich Pfaff, "Notice biographique sur Georges Cuvier," in Lettres de Georges Cuvier a C.M. Pfaff, 1788-1792, sur I'histoire naturelle, la politique et la litterature, trans. Louis Marchant (Paris, 1858), pp. 11-44, 293-296, esp. pp. 17-18. 48. On Cuvier's aversion to hypotheses, see Pfaff, Lettres de Georges Cuvier, pp. 72-73,134,160, 215,270. 49. Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 180n. 50. Ibid., pp. 180-181; Cuvier, Manuscript autobiography, pp. 23-24, 26; "Principaux discours prononces sur le tombe de M. Cuvier, le 16 mai 1832," Annales des sciences naturelles 26 (1832): 394-415, quotation on p. 404. For a variant, see Flourens, "Eloge historique d'Etienne Geoffroy Saint-Hilaire," p. viii. Outram suggests that the biographical tradition with respect to Cuvier tries to cover up the political nature of Cuvier's success in Paris and emphasizes instead the generous assistance of others in response to talent. She shows that Cuvier waged a deliberate campaign to become known. Outram, Georges Cuvier, chapters 3 and 8. 51. Cuvier, Manuscript autobiography, pp. 24-25, 27; Cuvier, "Memoires pour celui qui fera mon eloge," pp. 180-181. 52.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 65n; The Fables of La Fontaine, trans. Marianne Moore (New York: Viking, 1964), pp. 40-41, quotation on p. 41. 53. Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 181. For variants, see Cahn, Geoffroy Saint-Hilaire, p. 25 (who cites the passage as coming from a letter to a friend); and Flourens, "Eloge historique d'Etienne Geoffroy Saint-Hilaire," p. ix. For Cuvier's unkind comments on Hauy, see Cuvier, Manuscript autobiography, pp. 26-27. 54.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 60-70. 55. For other accounts of Geoffroy's early friendship and collaboration with Cuvier, see Flourens, "Eloge historique d'Etienne Geoffroy Saint-Hilaire," pp. viii-ix; and Cahn, Etienne Geoffroy Saint-Hilaire, pp. 24-28. On Geoffroy's nostalgia, see, for example, his eulogy of Cuvier in "Principaux discours prononces sur le tombe de M. Cuvier," pp. 402-408. 56. On the need to reform Linnaean natural history, see Cuvier, "Memoires pour servir a celui qui fera mon eloge," pp. 70-72. 57. Etienne Geoffroy Saint-Hilaire and Georges Cuvier, "Memoire sur une nouvelle division des Mammiferes et sur les principes qui doivent servir de base dans cette sorte de travail," Magasin encydopedique [ler annee] 2 (1795): 164-190; idem, "Lettre du citoyen Geoffroy, . . . et du citoyen Cuvier aux redacteurs du Magasin encydopedique sur le rhinoceros bicorne," Magasin encydopedique [l er annee] 1 (1795): 326-328; idem, "Histoire naturelle des orang-outangs," Magasin encydopedique [ler annee] 3 (1795): 451-463; Georges Cuvier and Etienne Geoffroy Saint-Hilaire, "Memoire sur les rapports naturels du Tarsier (Didelphis macrotarsus Gm.)," Magasin encydopedique [ler annee] 3 (1795): 147-154; idem, "Memoire sur les especes d'elephans," Bulletin de la Societe Philomatique 1 (1795): 90. 58. Pfaff, Lettres de Georges Cuvier, pp. 178-179. 59. Ibid., p. 214; Georges Cuvier, "Memoire sur les cloportes terrestres," Journal d'histoire naturelle 2 (1792): 18-31, quotation on pp. 18-19. Franck Bourdier, "Georges Cuvier," DSB, 3: 521528, claims that for a time Cuvier was converted to a belief in the chain of being. It seems to me that Bourdier mistakes discussions of graded nuances between families of animals for belief in a linear chain.

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60. Pfaff, Lettres de Georges Cuvier, pp. 134-136, 246-247. On the introduction of comparative anatomy into taxonomy, see Daudin, Cuvier et Lamarck and Coleman, Georges Cuvier, Zoologist, pp. 74-106. Michel Foucault, The Order of Things: An Archaeology of the Human Sciences (New York: Vintage, c.1970, 1973), pp. 263-279, explores the nature of the radical shift from classical natural history as represented by Linnaeus and the new functional doctrine based on comparative anatomy of Cuvier. 61. Geoffroy and Cuvier, "Memoire sur une nouvelle division des Mammiferes." Daudin, Cuvier et Lamarck, argues that in the end empirical techniques of classification were more important in overhauling the classification of the animal kingdom than the rational principles developed by Cuvier. 62. Georges Cuvier, "Memoire sur la structure interne et externe, et sur les affnites des animaux auxquels on a donne le nom de Vers," La decade philosophique 5 (1795): 385-396. 63. Cuvier and Geoffroy, "Memoire sur les rapports naturels du Tarsier," p. 154. 64. Geoffroy and Cuvier, "Histoire naturelle des orang-outangs," p. 452. 65. For the text of the letter, see Henry Fairfield Osborn, Cope, Master Naturalist (Princeton: Princeton University Press, 1931), pp. 9-10, quotation on p. 9. A slightly variant text of Geoffroy's letter was printed in Science, n.s. 19 (1904): 798-800. Although Lamarck signed the letter on behalf of the Museum administration, it was written by Geoffroy and it was to Geoffroy that Peale addressed his reply. The letter is therefore not an early indication of Lamarck's commitment to evolution, as has been suggested. Peale's reply, dated 30 April 1797, is in the American Philosophical Society. 66. See Bourdier, "Georges Cuvier," p. 523; idem, "Etienne Geoffroy Saint-Hilaire," p. 356. 67. On the history of the Museum, see Deleuze, Histoire et description du Museum; P. Bernard, L. Couailhac, P. Gervais, and E.M.M. Lamaout, Le Jardin des Plantes (Paris, 1842); and Cap, Le Museum d'Histoire Naturelle. For a modern analysis of the Museum's "Golden Age" in the first part of the century and subsequent decline in the period 1840-1870, see Limoges, "The Development of the Museum d'Histoire Naturelle." 68. On the lack of community at the Museum, see Outram, Georges Cuvier, chapter 8. 69. On the cabinet of the Stadholder, see Florence F.J.M. Pieters, "Notes on the Menagerie and Zoological Cabinet of Stadholder William V of Holland, Directed by Aernout Vosmaer," Journal of the Society for the Bibliography of Natural History 9 (1980): 539-563. 70. Deleuze, Histoire et description du Museum, 1: 67-157, esp. pp. 103-104. On the Cabinet, see Franck Bourdier, "Le Cabinet d'Histoire Naturelle du Museum, 1635-1935," Sciences, no. 18 (March-April 1962): 35-50. 71. Ibid., 2: 652. The following description of the Museum's resources is based on Deleuze. Deleuze presents an extended comparison of the facilities of the Museum in 1789 and 1822 (ibid., 1: 185-192). The entire second volume is devoted to a description of the contents of the Cabinet including the galleries of zoology (ibid., 2: 409-652) and the galleries of comparative anatomy (ibid., 2: 653-672, esp. pp. 670-672). On the library, see ibid., 2: 693-702, on the menagerie, see ibid., 1: 111-112, 2: 673-692), and on the journal, see ibid., 1: 99-100. 72. For other descriptions of the collection of comparative anatomy, see Georges Cuvier, "Notice sur I'etablissement de la collection d'anatomie comparee du Museum," Annales du Museum d'Histoire Naturelle 2 (1803): 409-414; and Achille Valenciennes, "Catalogue des preparations anatomiques laissees dans le Cabinet d'anatomie comparee du Museum d'histoire naturelle, par G. Cuvier, faisant suite a la notice inseree dans le tome II des Annales du Museum," Nouvelles annales du Museum d'Histoire Naturelle 2 (1833): 417-508. 73. Etienne Geoffroy Saint-Hilaire and Frederic Cuvier, Histoire naturelle des mammiferes, avec des figures originales, coloriees, dessinees d'apres des animaux vivans, publiee sous I'autorite de l'Administration du Museum d'Histoire Naturelle, 4 vols. (Paris, 1824-1842). 74. The Museum's journals from 1802 to 1874 are as follows: Annales, 20 vols. (1802-1813); Memoires, 20 vols. (1815-1832); Nouvelles annales, 4 vols. (1832-1835); Archives, 10 vols. (18391861); and Nouvelles archives, 10 vols. (1865-1874). 75. See Richard W. Burkhardt, Jr., "Lamarck, Evolution and the Politics of Science," Journal of the History of Biology 3 (1970): 275-298; idem, The Spirit of System, pp. 13-45 (on Lamarck as "naturalist-philosopher"), 186-202 (on the reception of his evolutionary theory); and Pietro Corsi, Oltre il mito: Lamarck e le scienze naturali del suo tempo (Bologna: il Mulino, 1983).

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76. Henri Marie Ducrotay de Blainville, Histoire des sciences de l'organisation et de leurs progres comme base de la philosophie, ed. F.-L.-M. Maupied, 3 vols. (Paris, 1858), 3: 358. 77. Bernard de Lacepede, Histoire naturelle des poissons, 5 vols. (Paris, 1798-1803), esp. "Discours sur la duree des especes," 2: xxiii-lxiv; Appel, "Lacepede." 78. Cuvier, Manuscript autobiography, p. 30. See also Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 182. 79. Cuvier, "Memoires pour servir a celui qui fera mon eloge," p. 184. One-third of the original sixty resident members of the First Class of the Institut were chosen by the Directory, which selected the most prestigious scientists of the day, among them Daubenton and Lacepede, named by an arrete of 20 November 1795. The nominated members were to select the remaining twothirds. Former members of the Academie des Sciences were generally given seats, even if some shuffling of sections had to be made to accommodate them. Thus the elderly anatomist and surgeon, Jacques-Rene Tenon, and Broussonet were elected to the section on 9 December. The last two members, Cuvier and Richard, were initially chosen on 13 December. 80. Ibid., p. 184. 81. Cuvier wrote, "That which gave me most favor among the savants, was that I was almost the only one who then envisioned natural history under a rational and philosophical point of view who utilized anatomy in zoology." Ibid., p. 182. 82. This argument was emphasized by Blainville, who wished to minimize Cuvier's scientific talent. See Henri Marie Ducrotay de Blainville, Cuvier et Geoffroy Saint-Hilaire: biographies scientifiques (Paris, 1890), p. 31. Chapter 3

1. Etienne Geoffroy Saint-Hilaire, Cours de I'histoire naturelle des mammiferes (Paris, 1829), "Discours preliminaire," p. 20; "Principaux discours prononces sur la tombe de M. Cuvier, le 16 mai 1832," Annales des sciences naturelles 26 (1832): 394-415, quotations on pp. 402, 403. 2. Georges Louis Duvernoy, Notice historique sur les ouwages et la vie de M. le Ber Cuvier (Paris, 1833), p. 1; Pierre Flourens, "Eloge historique de Georges Cuvier," in idem, Analyse raisonnee des travaux de Georges Cuvier, precedee de son eloge historique (Paris, 1841), pp. 1-72, quotation on p. 1. 3. William Coleman, Georges Cuvier, Zoologist: A Study in the History of Evolution Theory (Cambridge, Mass.: Harvard University Press, 1964); Henri Daudin, Cuvier et Lamarck: les classes zoologiques et I'idee de serie animale, 1790-1830, 2 vols. in 1 (Paris: Felix Alcan, 1926). 4. The writings of Dorinda Outram, especially her recent book, analyze in depth Cuvier's role as a public scientist and as a cultivator and dispenser of patronage. See Dorinda Outram, Georges Cuvier: Vocation, Science and Authority in Post-Revolutionary France (Manchester: Manchester University Press, 1984); idem, ed., The Letters of Georges Cuvier: A Summary Calendar of Manuscript and Printed Materials Preserved in Europe, the United States of America, and Australasia (Chalfont St. Giles: British Society for the History of Science, 1980), "Introduction," pp. 1-11; idem, "The Language of Natural Power: The 'Eloges' of Georges Cuvier and the Public Language of Nineteenth Century Science," History of Science 16 (1978): 153-178; and idem, "Politics and Vocation: French Science, 1793-1830," British Journal for the History of Science 13 (1980): 2743. See also my dissertation, "The Cuvier-Geoffroy Debate and the Structure of Nineteenth Century French Zoology," Ph.D. Dissertation, Princeton University, 1975, chapter 2. 5. By contrast, none of Geoffroy's book-length works achieved a contemporary second edition or translation. Jean Chandler Smith, Smithsonian Institution Libraries, is currently compiling a bibliography of all of Cuvier's writings, including translations. 6. The revolutionary character of each of these works and their importance in directing future research are recognized by Coleman, Georges Cuvier, Zoologist, pp. 2, 60. 7. Georges Cuvier, Lecons d'anatomie comparee, 5 vols. (Paris, 1800-1805). At the time of his death, Cuvier was working on a second edition of Lecons, published 1835-1846. 8. Ibid., 1: 47. 9. Georges Cuvier, Recherches sur les ossemens fossiles de quadrupedes, ou I'on retablit les car-

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acteres de plusieurs especes d'animaux que les revolutions du globe paroissent avoir detruites, 4 vols. (Paris, 1812); reprint ed. (Brussels: Culture et Civilisation, 1969). A second edition appeared in 1821-1824, a third in 1825, and a fourth in 1834-1836. 10. Georges Cuvier, "Memoire sur les especes d'elephans vivantes et fossiles," Memoires de I'Jnstitut National des Sciences et Arts 2 (1798): 1-22. On vertebrate paleontology before Cuvier, see John C. Greene, The Death of Adam: Evolution and Its Impact on Western Thought (Ames, Iowa: Iowa State University Press, 1959), chapter 4. 11. Cuvier, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 66-67. (Each memoir is separately paginated.) 12. See, for example, Honore de Balzac, The Magic Skin [Le peau de chagrin] trans. Katherine Prescott Wormeley (Boston, 1894), pp. 25-26. 13. Cuvier, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 58. 14. Ibid., 1: 4, 61. 15. Martin Rudwick, The Meaning of Fossils: Episodes in the History of Paleontology, 2nd ed. (New York: Science History Publications, 1976), chapter 3, stresses the massive evidence in Cuvier's time for catastrophes and insists that while Cuvier's conclusions were catastrophic, his geological method was actualistic. That is, he inferred past events by analogy with observable processes. On Cuvier's paleontology, see also Coleman, Georges Cuvier, Zoologist, chapter 5; and Peter J. Bowler, Fossils and Progress: Paleontology and the Idea of Progressive Evolution in the Nineteenth Century (New York: Science History Publications, 1976), pp. 16-22. 16. Cuvier, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 81-82. 17. Georges Cuvier, Le regne animal distribue d'apres son organisation, pour servir de base a I'histoire naturelle des animaux et d'introduction a I'anatomie comparee, 4 vols. (Paris, 1817); 2nd ed., 5 vols. (Paris, 1829-1830). 18. Etienne Geoffroy Saint-Hilaire and Georges Cuvier, "Memoire sur une nouvelle division des Mammiferes et sur les principes qui doivent servir de base dans cette sorte de travail," Magasin encyclopedique [ler annee] 2 (1795): 164-190; Georges Cuvier, "Memoire sur la structure interne et externe, et sur les affinites des animaux auxquels on a donne le nom de Vers," La decade philosophique 5 (1795): 385-396. 19. Georges Cuvier, Tableau elementaire de I'histoire naturelle des animaux (Paris, 1798), reprint ed. (Brussels: Culture et Civilisation, 1969); idem, Lecons d'anatomie comparee, 1: 60-88 and tables. 20. Georges Cuvier, "Sur un nouveau rapprochement a etablir entre les classes qui composent le regne animal," Annales du Museum d'Histoire Naturelle 19 (1812): 73-84. 21. Ibid., p. 76. The same statement appears in Cuvier, Le regne animal, 1: 57. 22. For an example of a disciple attributing philosophical anatomy to Cuvier, see Georges Louis Duvernoy, "Anatomie," in Dictionnaire universel d'histoire naturelle, ed. Charles Dessalines d'Orbigny, 13 vols. (Paris, 1842-1861), 1: 438-454, esp. section on "Anatomie philosophique, transcendante et speculative," pp. 445-449. On Blainville, see Toby A. Appel, "Henri de Blainville and the Animal Series: A Nineteenth-Century Chain of Being," Journal of the History of Biology 13 (1980): 291-319. On the many meanings of "type," see Paul Lawrence Farber, "The TypeConcept in Zoology during the First Half of the Nineteenth Century," Journal of the History of Biology 9 (1976): 93-119. For an example of a more modern interpretion of Cuvier as a believer in abstract types, see Edward Lurie, Louis Agassiz: A Life in Science (Chicago: University of Chicago Press, 1960), pp. 60-62. 23. Cuvier, "Sur un nouveau rapprochement," p. 79. 24. Flourens, "Eloge historique de Georges Cuvier," p. 49. 25. Cuvier, Lecons d'anatomie comparee, 1: xix. 26. Cuvier, Le regne animal, 1: 12. 27. On teleology in Cuvier's biological thought, see Coleman, Georges Cuvier, Zoologist, chapter 2; and E.S. Russell, Form and Function: A Contribution to the Study of Animal Morphology (London: John Murray, 1916); reprint ed. (Chicago: University of Chicago Press, 1982), chapter 3. 28. Cuvier, Le regne animal, 1: 6. 29. Cuvier, Lecons d'anatomie comparee, 1: 47, 55-56; idem, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 58-63, quotation on p. 61.

254

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30. Cuvier, Lecons d'anatomie comparee, 1: 47-53. 31. See Charles Coulston Gillispie, "Etienne Bonnot, abbe de Condillac," DSB, 3: 380-383. 32. Cuvier, Le regne animal, 1:10. 33. Cuvier, Lecons d'anatomie comparee, 1: v-vi. 34. Georges Cuvier, Rapport historique sur les progres des sciences naturelles depuis 1789, et sur lew etat actuel (Paris, 1810), p. 9. 35. Comte's heroes of positivism, celebrated in his calendar of his "religion of humanity," included Stahl, Broussais, Lamarck, and Gall, men whom Cuvier and his disciples would have branded as speculators with insufficient respect for "the positive facts." Comte chose his heroes for the very reason Cuvier rejected them: they all had a materialist tinge to their writings. For his critique of the history of biology, see Auguste Comte, Cows de philosophie positive, 2nd ed., 6 vols. (Paris, 1864), vol. 3. See also idem, The Catechism of Positive Religion, trans. Richard Congreve (Clifton, N.J.: Augustus M. Kelly, 1973), Table D. 36. This point was noted by William Coleman, "Les organismes marins et 1'anatomie comparee dite experimentale: 1'oeuvre de Georges Cuvier," Vie et milieu, Supplement no. 19 (1965): 225238, esp. pp. 236-237. 37. Cuvier, Rapport historique, pp. 11-12. 38. Georges Cuvier, Recueil des eloges historiques lus dans les seances publiques de I'lnstitut Royal de France, 3 vols. (Paris, 1819-1827), 1: vi. For a complete bibliography of Cuvier's eloges, see Outram, "The Language of Natural Power." Outram sees in the eloges and in French science generally a contradiction between the ideology of the purity of scientific inquiry and the highly personal and political nature of the patronage system. 39. Cuvier, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 25-35, quotation on p. 28. See also Richard W. Burkhardt, Jr., "Lamarck, Evolution, and the Politics of Science," Journal of the History of Biology 3 (1970): 275-298, esp. pp. 294-295. Another early example of Cuvier's ridiculing of Lamarck was noted by a foreign visitor, Joseph Marzari Pencati, who attended Cuvier's course at the Athenee in 1804. See Danielle Plan, Un genevois d'autrefois: HenriAlbert Gosse, 1753-1816 (Paris, 1909), pp. Ixxxii-lxxxiv. 40. Georges Cuvier, "Eloge historique de Daubenton, lu le 5 avril 1800," in Recueil des eloges, 1: 37-80, quotations on pp. 42-43, 56. 41. Georges Cuvier, "Eloges historiques de Charles Bonnet et H.B. de Saussure, lus le 3 Janvier 1810," in Recueil des eloges, 1: 383-409, quotations on pp. 395, 396, 401. 42. Cuvier, Rapport historique, p. 255; idem, Lecons d'anatomie comparee, 1: 6-8. 43. Cuvier, Lecons d'anatomie comparee, 1: 1-6, 19; idem, Tableau elementaire, pp. 5-6; idem, Rapport historique, pp. 224-225; idem, Le regne animal, 1: 13-16. See also Xavier Bichat, Recherches physiologiques sur la vie et la mort, 2nd ed. (Paris, 1802), pp. 1 -2, 78-84. 44. See Chapter 2, note 59. 45. Daudin, Cuvier et Lamarck, 2: chapter 10; Jean-Baptiste Lamarck, Philosophie zoologique, ou, Exposition des considerations relatives a I'histoire naturelle des animaux, . . . (Paris, 1809). On Lamarck's theory of evolution, see Richard W. Burkhardt, Jr., The Spirit of System: Lamarck and Evolutionary Biology (Cambridge, Mass.: Harvard University Press, 1977). 46. Cuvier, Lecons d'anatomie comparee, 1: 59-60, quotation on p. 60. See also Cuvier's critique of the chain of being in "Eloges historiques . . . Bonnet," p. 400. 47. Cuvier, "Sur un nouveau rapprochement," pp. 80-81. 48. Cuvier, Le regne animal, 1: xx-xxi. 49. Cuvier, "Memoire sur les especes d'elephans," p. 12. 50. See Coleman, Georges Cuvier, Zoologist, p. 1. 51. Burkhardt, "Lamarck, Evolution and the Politics of Science," esp. pp. 283-284; idem, The Spirit of System, chapter 7. 52. Cuvier's arguments against evolution are examined by Coleman, Georges Cuvier, Zoologist, chapters 6 and 7. Cuvier's most extensive critique of evolution appeared in Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 73-81. 53. Cuvier is a prime example of what has been termed a "public scientist," that is, a scientist who justified the scientific enterprise to political and social authorities. See Frank M. Turner, "Public Science in Britain, 1880-1919," Isis 71 (1980): 589-608. By extending the analogy with

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the situation Turner describes in Victorian Britain, one might interpret Cuvier's pronouncements as an appropriate professionalizing strategy for the political and social context of early nineteenthcentury France. See Frank M. Turner, "The Victorian Conflict between Science and Religion," Isis 69 (1978): 356-376. 54. On Cuvier as an educator, see Outram, Georges Cuvier, chapter 4; John Vienot, Le Napoleon de I'intelligence: Georges Cuvier, 1769-1832 (Paris: Fischbacher, 1932), chapter 5; and Georgette Legee, "La participation de Georges et de Frederic Cuvier a 1'organisation de 1'instruction publique (1802-1838), Histoire et nature 4 (1974): 47-72. 55. On Cuvier as an administrator, see Outram, Georges Cuvier, chapters 4 and 5; Vienot, Le Napoleon de I'intelligence, chapter 5 (pp. 113-143) and pp. 231-234; Henry Puget, "Cuvier au Conseil d'Etat," Revue politique et parlementaire 152 (1932): 300-319; and Baron E. Pasquier, Eloge de M. le Baron Cuvier (Paris, 1832), 55pp., also published in Paris; ou, Le livre des cent-etun 9 (1832): 329-390. 56. When Lyell visited Paris in 1823, he reported that Alexander von Humboldt, speaking of Cuvier "with great vivacity and force," informed him, "No, Cuvier gives no lectures, and the reason I regret to say is, that he is still a Politician." Lyell to his father, 8 July 1823, in Mary Katherine Lyell, Life, Letters and Journals of Sir Charles Lyell, Bart., 2 vols. (London, 1881), 1: 126. 57. Coleman points to Cuvier's conservatism in politics and concern for order. Outram argues that, judged by the standards of his time, Cuvier, although he had little concern for social justice, was not, in fact, a conservative. Coleman, Georges Cuvier, Zoologist, pp. 11, 14, 17; Outram, Georges Cuvier, pp. 95, 105 58. Christoph Heinrich Pfaff, Lettres de Georges Cuvier a C.M. Pfaff, 1788-1792, sur I'histoire naturelle, la politique et la litterature, trans. Louis Marchant (Paris, 1858), pp. 115, 204. Franck Bourdier, "Georges Cuvier," DSB, 3: 521-528, claims (p. 522), without citing evidence, that Cuvier only feigned sympathy for the Revolution because he feared that his letters would be opened by the French police. This seems far-fetched. 59. Pfaff, Lettres de Georges Cuvier, pp. 271, 286, quotation on p. 281. 60. For example, in his eloge of Daubenton (note 40, p. 68), he wrote of 1793 as a time when "by a reversal of ideas that will long be memorable in history, the most ignorant portion of the people had to pronounce on the fate of the most instructed and the most generous." Cuvier devoted most of his eloge of Olivier to detailing the evils of political chaos in the Orient during the period of Olivier's voyage, which Cuvier regarded as hardly worse than the Terror that Olivier had escaped. Georges Cuvier, "Eloge historique de Guillaume-Antoine Olivier, lu le 8 Janvier 1816," in idem, Recueil des eloges, 2: 235-265. 61. Pfaff, Lettres de Georges Cuvier, p. 118. 62. On Mesmer, see Robert Darnton, Mesmerism and the End of the Enlightenment in France (Cambridge, Mass.: Harvard University Press, 1968). On Gall, see Owsei Temkin, "Gall and the Phrenological Movement," Bulletin of the History of Medicine 21 (1947): 275-321. On Pouchet, see John Farley and Gerald L. Geison, "Science, Politics and Spontaneous Generation in Nineteenth-Century France: The Pasteur-Pouchet Debate," Bulletin of the History of Medicine 48 (1974): 161-198. On the reception of Darwin, see below, chapter 8. 63. Quoted by Vienot, Le Napoleon de I'intelligence, p. 137. 64. For example, Humboldt remarked to Lyell in 1823: "You observe that his soirees are mostly attended by English; the truth is, the French savans have in general cut him; his continual changing over to each new party that came into power at length disgusted almost all, and you know that it has been long a charge against men of science, that they were pliant tools in the hands of princes and ministers, and might be turned which way they pleased. That such a man as Cuvier should have given a sanction to such an accusation was felt by all as a deep wound to the whole body." M.K. Lyell, Life, Letters, 1: 126-127. 65. Pasquier, "Eloge," p. 40. See also Puget, "Cuvier au Conseil d'Etat," p. 314; Vienot, Le Napoleon de I'intelligence, chapter 5; and Coleman, Georges Cuvier, Zoologist, pp. 10-11. 66. On Cuvier's religious views, see Coleman, Georges Cuvier, Zoologist, pp. 15-16; Vienot, Le Napoleon de I'intelligence, chapter 6; and Outram, Georges Cuvier, chapter 7. 67. Georges Cuvier, "Eloge historique de Joseph Priestley, lu le 24 juin 1805," in Cuvier, Recueil des eloges, 1: 189-234, quotations on pp. 191, 192, 222, 234.

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68. See, for example, Reijer Hooykaas, Natural Law and Divine Miracle: The Principle of Uniformity in Geology, Biology and Theology (Leiden: E.J. Brill, 1963), pp. 197-198; and Michael Ruse, The Darwinian Revolution: Science Red in Tooth and Claw (Chicago: University of Chicago Press, 1979), p. 63. 69. Noe'1-Antoine Pluche, Le spectacle de la nature, 8 vols. (Paris, 1732-1750); Camille Limoges, "Noel-Antoine Pluche," DSB, 11: 42-44; Rene-Antoine Ferchault de Reaumur, Memoires pour servir a I'histoire naturelle des insectes, 6 vols. (Paris, 1734-1742). On British natural theology, see Charles Coulston Gillispie, Genesis and Geology: A Study in the Relations of Scientific Thought, Natural Theology, and Social Opinion in Great Britain, 1790-1850 (New York: Harper & Row, 1959). 70. This argument was suggested by a conversation with Gerald Geison. On the different form taken by French industrialization, see Rondo Cameron and Charles E. Freedeman, "French Economic Growth: A Radical Revision," Social Science History 1 (Winter 1983): 3-30. I thank Andrew Butrica for this reference. British editors tended to inject more overt references to religion into Cuvier's works. See, for example, Georges Cuvier, The Animal Kingdom, 15 vols., ed. Edward Griffith (London, 1827-1835), 1: xi. On Robert Jameson's editing of Cuvier, see Rudwick, The Meaning of Fossils, pp. 132-135. 71. Turner, in writing of post-Darwinian science in Britain, links emerging professionalism with demands for autonomy from religious authorities. The argument might be adapted to the somewhat different situation in early nineteenth-century France. See Turner, "The Victorian Conflict Between Science and Religion." 72. Roger Hahn, "Laplace and the Vanishing Role of God in the Physical Universe," in Harry Woolf, ed., The Analytic Spirit: Essays in the History of Science in Honor of Henry Guerlac (Ithaca, N.Y.: Cornell University Press, 1981), pp. 85-95. 73. Henri Marie Ducrotay de Blainville, Histoire des sciences de I'organisation et de lews progres comme base de la philosophie, ed. F.L.M. Maupied, 3 vols. (Paris, 1858). 74. Quoted by Pol Nicard, "Etude sur la vie et les travaux de M. de Blainville," in Henri Marie Ducrotay de Blainville, Osteographie, ou, Description iconographique comparee du squelette et du systeme dentaire des mammiferes recents et fossiles pour servir de base a la zoologie et a la geologie, 4 vols. (Paris, 1839-1864), 1: i-ccxxiii, esp. pp. cxiv-cxx, quotation on p. cxix. See also Appel, "Blainville and the Animal Series," pp. 313-314. 75. Hooykaas, Natural Law and Divine Miracle, pp. 197-198; Coleman, Georges Cuvier, Zoologist, pp. 176-182; Rudwick, The Meaning of Fossils, pp. 111-112, 133-135. 76. Cuvier, Recherches sur les ossemens fossiles, "Discours preliminaire," 1: 94-110. 77. M.K. Lyell, Life, Letters, 1: 276; Georges Cuvier, "Memoires pour servir a celui qui fera mon eloge, ecrits au crayon dans ma voiture pendant mes courses en 1822 et 1823," in Pierre Flourens, Recueil des eloges historiques, lus dans les seances publiques de I'Academie des Sciences, 3 vols. (Paris, 1856-1862), 1: 169-193, quotation on p. 190; Charles Leopold Laurillard, "Cuvier," in Biographie universelle, ancienne et moderne, 85 vols. (Paris, 1811-1862) (henceforth Biographie universelle), 61: 588-605, quotation on p. 600. On Sand and Balzac, see below, chapter 7. 78. Cuvier, Rapport historique, p. 387. In his first book, Tableau elementaire, Cuvier had written (p. xii): "Those who are peaceably occupied with the study of nature should be little tempted to hurl themselves into the stormy career of ambition. They should succumb with difficulty to brutal or cruel passions, the usual stumbling-blocks of those ardent minds who are incapable of mastering their actions. Pure as the objects of their researches, they should be aroused toward all that surrounds them by that same beneficence that they see shown by nature toward all her productions." 79. Cuvier, Le regne animal, 1: xix-xx. 80. On patronage in early nineteenth-century French science, see Maurice Crosland, The Society of Arcueil: A View of Science at the Time of Napoleon I (Cambridge, Mass: Harvard University Press, 1967); Outram, Georges Cuvier, idem, "Politics and Vocation"; Robert Fox, "The Rise and Fall of Laplacian Physics," in Historical Studies in the Physical Sciences, vol. 4., ed. Russell McCormmach (Princeton; Princeton University Press, 1975), pp. 89-136; idem, "Scientific Enterprise and the Patronage of Research in France, 1800-1870," Minerva 11 (1973): 442-473; and Eugene Frankel, "Corpuscular Optics and the Wave Theory of Light: The Science and Politics of a Revolution in Physics," Social Studies of Science 6 (1976): 141-184.

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81. On Cuvier's suppleants, see Henri Marie Ducrotay de Blainville, Cuvier et Geoffroy SaintHilaire; biographies scientifiques (Paris, 1890), pp. 243-244. On Lacepede, see Toby A. Appel, "B.-G.-E. de la Ville-sur-1llon, comte de Lacepede," DSB, 7: 546-548. 82. Maurice Crosland, "The Development of a Professional Career in Science in France," Minerva 13 (1975): 38-57, quotation on p. 54; Paul Antoine Gratacap Cap, Le Museum d'Histoire Naturelle: histoire de la fondation et des developpements successifs de I'etablissement (Paris, 1854), p. 7; E.T. Hamy, Les derniers jours du Jardin du Roi et la fondation du Museum d'Histoire Naturelle (Paris, 1893), pp. 146-150. 83. On the College de France, see Jean Torlais, "Le College Royal," in Rene Taton, ed., Enseignement et diffusion des sciences en France au XVIII e siecle (Paris: Hermann, 1964), pp. 261286; College de France, Le College de France (1530-1930): livre jubilaire compose a I'occasion de son quatrieme centenaire (Paris: Presses Universitaires de France, 1932), especially the article by Andre Mayer, "L'histoire naturelle et la physiologie au College de France," pp. 167-190. The College de France did become a center for experimental physiology. Francois Magendie, holder of the chair of medicine from 1830 to 1855, and Claude Bernard, holder of the chair from 1855 to 1878, set up laboratories and performed much of their celebrated research there. 84. The professors of histoire naturelle medicale were Claude Richard (1754-1821), primarily a botanist, Achille Richard (1794-1859), a physician and botanist, and Alfred Moquin-Tandon (1804-1863), who published in both botany and zoology. 85. A. Railliet and L. Moule, Histoire de I'Ecole d'Alfort (Paris: Asselin et Houzeau, 1908), pp. 60-77, 130-131, 136-137, 313-314. On Alfort in the eighteenth century, see Caroline C. Hannaway, "Veterinary Medicine and Rural Health Care in Pre-Revolutionary France," Bulletin of the History of Medicine 51 (1977): 431-447. 86. On the ecoles centrales and the lycees, see L. Pearce Williams, "Science, Education and the French Revolution," Isis 44 (1953): 311-330; Francois Joseph Picavet, Les ideologues: essai sur I'histoire des idees et des theories scientifiques, philosophiques, religieuses, etc. en France depuis 1789 (Paris, 1891), pp. 32-69; and F.V.A. Aulard, Napoleon I" et le monopole universitaire: origines et fonctionnement de I'Universite Imperiale (Paris: A. Colin, 1911). The secondary schools in existence in a given year, the chairs and the names of the professors can be found by consulting the Almanach national, Almanach imperial, or Almanach royal published annually by the French government. In the 1830s, Milne Edwards was teaching at the College Henri V in Paris before he obtained posts at the Faculte des Sciences and Museum. 87. On the Faculties, see Terry Shinn, "The French Science Faculty System, 1808-1914: Institutional Change and Research Potential in Mathematics and the Physical Sciences," in Historical Studies in the Physical Sciences, vol. 10, ed. Russell McCormmach and Lewis Pyenson (Baltimore: Johns Hopkins University Press, 1979), pp. 271-332. Older sources include Louis Liard, L'enseignement superieur en France, 1789-1893, 2 vols. (Paris, 1888-1894); and Aulard, Napoleon Ier et le monopole universitaire, esp. pp. 325-329. The faculties in existence in a given year, the occupants of the chairs, and the requirements for degrees can be found in the Almanach imperial or Almanach royal. 88. The Paris Faculte des Sciences was intentionally organized so that it would predominate over the provincial faculties. It was to consist of eight professors chosen from among the professors already teaching in Paris: two from the Ecole Polytechnique, two from the College de France, two from the Museum, and two professors of mathematics from the lycees. Thus set up, the Paris Faculte boasted some of the greatest names in French science: Biot, Poisson, Lacroix, Gay-Lussac, Thenard, Hauy, Desfontaines, and Geoffroy. See Aulard, Napoleon Ier et le monopole universitaire, p. 327; and Almanach imperial (1810). 89. An arrete of the council of the University of 23 May 1809 automatically granted the doctorat es sciences to members of the First Class of the Institut and to professors at the Ecole Polytechnique, Museum, and College de France. See Aulard, Napoleon Ier et le monopole universitaire, p. 174. 90. For an example of a zoologist who followed this pattern perfectly, see Jean-Victor Audouin, Journal d'un etudiant en medecine et en sciences a Paris sous la Restauration (1817-1818), ed. Jean Theodorides (Paris, 1959), also in Histoire de la medecine 9 (Nov. 1958): 4-63; 9 (Dec. 1958): 5-56; 10 (Jan. 1959): 5-48. Audouin became professor of entomology at the Museum in 1833.

258

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91. The history of the chair of zoology is discussed by Henri de Lacaze-Duthiers, "Dix-sept annees d'enseignement de la zoologie en Sorbonne," Revue scientifique, 3rd series, 37 (1886): 737748. On Milne Edwards, see below, chapter 8. 92. On the Ecole Normale, see Crosland, Society of Arcueil, pp. 218-219; and Craig Zwerling, "The Emergence of the Ecole Normale Superieure as a Centre of Scientific Education in the Nineteenth Century," in Robert Fox and George Weisz, eds., The Organization of Science and Technology in France, 1808-1914 (Cambridge: Cambridge University Press, 1980), pp. 31-60. 93. Pierre Lafitte, "Documents pour servir a 1'histoire de 1'Athenee," Revue occidentale, 2nd series, 10 (July-Dec. 1984): 261-274. 94. The question of whether early nineteenth-century French science was "professionalized" has been much debated. Some historians, among them Crosland, would readily grant the savants of postrevolutionary France the status of professional scientists because they held full-time remunerated positions obtained after having completed a period of training. For other historians, professional science is inconceivable unless it is centered in the university. The sociologist Ben-David has argued that French science in this period was not truly professional, because it did not combine teaching and research in a university setting and because it did not continue to expand and flourish after its moment of glory during the Napoleonic regime. The new institutions of higher education established in France between 1793 and 1800 represent to him not a novel departure but rather the culmination of eighteenth-century patterns of support. More recently, Outram has suggested that sociological theories of professionalization and institutionalization are of only limited use in understanding French science in this period. Although Outram seems to me to underestimate the institutional transformation of French science, she justly points to the highly personal nature of research programs and patronage, the limited number of posts, and the close linkage of science with political life, all factors that do not coincide with common definitions of professionalization. On this issue and the related question of whether and why French science declined after 1830, see Maurice Crosland, "The Development of a Professional Careeer in Science in France," also in idem, ed., The Emergence of Science in Western Europe (New York: Science History Publications, 1976), pp. 139-159; Joseph Ben-David, "The Rise and Decline of France as a Scientific Center," Minerva 8 (1970): 160-179, also in idem, The Scientist's Role in Society (Englewood Cliffs, N.J.: Prentice-Hall, 1971), pp. 88-107; Roger Hahn, "Scientific Careers in Eighteenth Century France," in Crosland, ed., Emergence of Science, pp. 127-138; Robert Fox, "Scientific Enterprise and the Patronage of Research in France"; Robert Gilpin, France in the Age of the Scientific State (Princeton: Princeton University Press, 1968); Harry W. Paul, "The Issue of Decline in Nineteenth-Century French Science," French Historical Studies 1 (1972): 416-450; Outram, Georges Cuvier, and idem, "Politics and Vocation." 95. On the Academie des Sciences in the eighteenth century, see Roger Hahn, The Anatomy of a Scientific Institution: The Paris Academy of Sciences, 1666-1803 (Berkeley: University of California Press, 1971); and James E. McClellan III, "The Academie Royale des Sciences, 1699-1793: A Statistical Portrait," Isis 72 (1981): 541-567. For an account of the functioning of the Institut during the Empire, see Crosland, Society of Arcueil, chapter 3. For the membership of the Academie, see Index biographique des membres et correspondants de I'Academie des Sciences de 1666 a 1939 (Paris: Gauthier-Villars, 1939) and later editions; and the tables according to section in Le Comte de Franqueville, Le premier siecle de I'lnstitut de France, 25 octobre 1795-25 octobre 1895 (Paris, 1895), pp. 444-447. Hahn claimed (p. 304) that since the nineteenth-century Academie was no longer a locus for original research, "it ceased to be a vital force for the scientific community"; it did little more than reward discoveries made elsewhere. Crosland, however, recognizes the central importance of the Academie's role in providing "a focus of ambition in clearly designated channels." Maurice Crosland, "The French Academy of Sciences in the Nineteenth Century," Minerva 16 (1978): 73-102, quotation on p. 101. 96. On the overlap of Museum and Academie membership, see Camille Limoges, "The Development of the Museum d'Histoire Naturelle of Paris, c. 1800-1914," in Fox and Weisz, eds., The Organization of Science and Technology in France, pp. 211-240, esp. p. 217. After 1802, if a vacancy occurred in any of the ecoles speciales, which included the Museum, Bonaparte as First Consul would be given a choice among three candidates, one presented by the institution in question, one by the inspectors general of public education, and one by the Academie des Sciences.

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During the early Restoration this system was continued in modified form; the king filled vacancies on the presentation of one candidate each by the Museum and Academie. See J.P.F. Deleuze, Histoire et description du Museum Royal d'Histoire Naturelle, 2 vols. (Paris, 1823), 1: 74, 74n. The election of candidates by both Museum and Academie, which continued even after the July Revolution, could lead to vicious politicking, as occurred in the cases of Lamarck's successor in 1830 and Cuvier's and Portal's successors in 1832. 97. For information on who read papers at meetings of the Academie, see Proces-verbaux des seances de I'Academie des Sciences tenues depuis la fondation de I'lnstitut jusqu'au mois d'aout 835 [1795-1835], 10 vols. (Hendaye: Academie des Sciences, 1910-1922) (hereafter, Proces-verbaux). On Cuvier's encouragement of Audouin, Magendie, and Flourens, see below, chapter 5, note 51. 98. See, for example, Bourdier, "Georges Cuvier," p. 527. 99. Nicard, "Blainville," pp. cxcviii-ccv; E.T. Hamy, ed., "Notes intimes sur Georges Cuvier redigees en 1836 par le Dr Quoy pour son ami J. Desjardins, de Maurice," Archives de medecine navale 86 (1906): 450-475, quotations on pp. 457, 462. 100. On Laurillard and Dumeril, see Pierre Flourens, "Eloge historique d'Andre-Marie-Constant Dumeril," Memoires de I'Academie des Sciences 35 (1866): i-xxii. On Duvernoy, see [E. Desmarest] "Duvernoy," Nouvelle biographie generale, 15: columns 554-556. On F. Cuvier, see Franck Bourdier, "Frederic Cuvier," DSB, 3: 520-521; and Pierre Flourens, "Eloge historique de F. Cuvier," Memoires de I'Academie des Sciences 18 (1842): i-xxviii. On Valenciennes, see Toby A. Appel, "Achille Valenciennes," DSB, 13: 554-555. 101. On Flourens, see below, chapters 4 and 5. 102. On Blainville, see William Coleman, "Henri Marie Ducrotay de Blainville," DSB, 2: 186188; Pierre Flourens, "Eloge de Marie-Henri Ducrotay de Blainville," Memoires de I'Academie des Sciences 27, Part 2 (1860): i-ix, and also in Recueil des eloges, 1: 285-341; and Appel, "Blainville and the Animal Series." The following account is especially derived from Nicard, "Blainville," also published separately (Paris, 1890). Nicard's biography includes the text of a letter by Blainville to Cuvier, written some time between 1830 and 1832 and probably never sent, which details the history of their stormy relationship (pp. cxcviii-ccv). 103. The methodical manner in which Cuvier organized his working space and time was nicely described by Lyell. See M.K. Lyell, Life, Letters, 1: 249-250. 104. Hamy, "Notes intimes sur Georges Cuvier," p. 470. Quoy wrote, "In the Museum, Cuvier was esteemed, a bit feared, but not loved" (p. 470). Chapter 4

1. See below, note 58. 2. On Belon, see E.W. Gudger, "Some Early and Late Illustrations of Comparative Anatomy," Annals of Medical History, n.s. 1 (1929): 334-355, esp. pp. 336-341; F.J. Cole, A History of Comparative Anatomy From Aristotle to the Eighteenth Century [1949] (New York: Dover, 1975), pp. 7-10; and M. Wong, "Pierre Belon," DSB, 1: 595-596. 3. On Goethe, see below, chapter 6. 4. Felix Vicq d'Azyr, "Memoire sur les rapports qui se trouvent entre les usages et la structure des quatre extremites dans I'homme et dans les quadrupedes," in Oeuvres de Vicq-d'Azyr, ed. Jacques Moreau de la Sarthe, 6 vols. (Paris, 1805), 4: 313-337. 5. On the term "homology," see below, chapter 5, note 4. Richard Owen, Lectures on the Comparative Anatomy and Physiology of the Invertebrate Animals, Delivered at the Royal College of Surgeons in 1843 (London, 1843), pp. 374, 379; idem, On the Nature of Limbs (London, 1849), p. 5. 6. Etienne Geoffroy Saint-Hilaire, Philosophie anatomique; des organes respiratoires sous le rapport de la determination et de I'identite de lews pieces osseuses (Paris, 1818), reprint ed. (Brussels: Culture et Civilisation, 1968), p. 9. In this work, Geoffroy uses the phrase "unity of composition" several times in passing. In later writings, he gives it more emphasis. 7. Isidore Geoffroy Saint-Hilaire, Vie, travaux et doctrine scientifique d'Etienne Geoffroy SaintHilaire (Paris, 1847), reprint ed. (Brussels: Culture et Civilisation, 1968), p. 74.

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8. Georges Cuvier, "Memoires pour servir a celui qui fera mon eloge," in Pierre Flourens, Recueil des eloges historiques lus dans les seances publiques de I'Academie des Sciences, 3 vols. (Paris, 1856-1862), 1: 169-193, quotation on p. 185. 9. On the Commission on Science and the Arts, see Maurice Crosland, The Society of Arcueil: A View of French Science at the Time of Napoleon I (Cambridge, Mass.: Harvard University Press, 1967), pp. 14-17; and J. Christopher Herold, Bonaparte in Egypt (New York: Harper & Row, 1962), pp. 164-176. 10. Etienne Geoffroy Saint-Hilaire, Lettres ecrites d'Egypte a Cuvier, Jussieu, Lacepede, Monge, Desgenettes, Redoute Jeune, Norry, etc., aux professeurs du Museum et a sa famille, ed. E.T. Hamy (Paris: Hachette, 1901), p. 31. Other sources for Geoffroy's voyage to Egypt include I. Geoffroy, Etienne Geoffroy Saint-Hilaire, chapter 3; and Jean-Eduoard Goby, "Etienne Geoffroy SaintHilaire en Egypte, 1798-1801," Cahiers d'histoire egyptienne, series 5, fascicule 2-3 (June 1953): 139-160. Further information on some of the manuscripts written by Geoffroy in Egypt can be found in Yves Laissus, "Catalogue des manuscrits d'Etienne Geoffroy Saint-Hilaire conserves au Museum," Revue d'histoire des sciences 25 (1972): 365-390; and idem, "Catalogue des manuscrits d'Etienne Geoffroy Saint-Hilaire conserves aux Archives de 1'Academic des Sciences de 1'Institut de France," Histoire et nature 1 (1973): 71-89. Accounts of the meetings of the Institut d'Egypte were published in La decade egyptienne, 3 vols. (Cairo, 1798-1800); Institut d'Egypte, Memoires sur I'Egypte, 4 vols. (Paris, 1798-1801); Courier de I'Egypte; and Magasin encyclopedique. 11. According to Isidore Geoffroy, the first year in Egypt was one of the happiest of his father's life. I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 81. 12. Herold, Bonaparte in Egypt, p. 169. 13. Geoffroy, Lettres ecrites d'Egypte, pp. 137-138. 14. Jean-Baptiste Lamarck, Georges Cuvier, and Bernard de Lacepede, "Rapport des professeurs du Museum, sur les collections d'histoire naturelle rapportes d'Egypte par E. Geoffroy," Annales du Museum d'Histoire Naturelle 1 (1802): 234-241, quotation on p. 241. Joseph Dombey (17421794) had explored Peru, Chile, and Brazil from 1777 to 1785, leaving most of his collections to the Jardin des Plantes. 15. Geoffroy discovered that the plover was the trochilus of the ancients, and he saw this bird fly unafraid into the mouth of a crocodile, as in the legend of Herodotus. See I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 89. 16. Geoffroy, Lettres ecrites d'Egypte, pp. 95-96. 17. Etienne Geoffroy Saint-Hilaire, "Observations sur 1'aile de 1'autruche," La decade egyptienne 1 (1798): 46-51, quotations on pp. 49, 50. This memoir also appeared in Institut d'Egypte, Memoires sur I'Egypte 1 (1798): 79-87. 18. Etienne Geoffroy Saint-Hilaire, "Memoire sur les prolongemens frontaux des animaux ruminans," Memoires de la Societe d'Histoire Naturelle (1799): 91-101, quotation on pp. 96-97. 19.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 95. 20. Geoffroy, Lettres ecrites d'Egypte, pp. 129-130. 21. Etienne Geoffroy Saint-Hilaire, "Exposition d'un plan d'experiences pour parvenir a la preuve de la co-existence des sexes dans les germes de tous les animaux," 20pp., Dossier Etienne Geoffroy Saint-Hilaire, Archives de 1'Academie des Sciences, Institut de France. Another version of this manuscript is located in F17 1099, Archives Nationales de France. 22. Ibid. Geoffroy quoted this conclusion in a letter to Cuvier dated 9 November 1800. See Geoffroy, Lettres ecrites d'Egypte, p. 193. 23. Etienne Geoffroy Saint-Hilaire, "Histoire de la formation de 1'oeuf, servant d'introduction au developpement des experiences annonces dans la derniere seance a 1'egard des oiseaux, experiences entreprises dans la vue d'arriver a des preuves directes de la coexistence des sexes dans les germes des etres vivans," 20pp., F17 1099, Archives Nationales de France. Another version of this memoir, titled "Histoire naturelle de 1'oeuf," 10pp., without the list of experiments, is located in Dossier Etienne Geoffroy Saint-Hilaire, Archives de 1'Academie des Sciences, Institut de France. 24. Geoffroy, "Exposition d'un plan d'experiences." 25. Geoffroy, "Histoire de la formation de 1'oeuf." 26. "Extrait des registres de 1'Institut du Kaire. Rapport sur une suite d'experiences proposees par le citoyen Geoffroy," signed by Joseph Fourier, 3pp., Ms 2712, IX, Bibliotheque Centrale du

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Museum National d'Histoire Naturelle. Other versions of this report appeared in Institut d'Egypte, Memoires sur I'Egypte 3 (1800): 385-387, and in Courier de I'Egypte, No. 111, 30 germinal, an IX [19 April 1801]. 27. Etienne Geoffroy Saint-Hilaire, "Experiences dans la vue de rechercher les causes qui determine, dans les plantes dioiques, 1'avortement d'un sexe de preference et a la exclusion de 1'autre," 5 diagrams, Dossier Etienne Geoffroy Saint-Hilaire, Archives de 1'Academic des Sciences, Institut de France. 28. Etienne Geoffroy Saint-Hilaire, "Considerations sur la fibre musculaire au moyen desquelles on explique sans recourir a la supposition d'un fluide nerveux, les causes mecaniques de la paralyse musculaire, du sommeil periodique, de 1'affaiblement, de 1'epuisement, de la reparation, et de la contraction naturelle des muscles," 23pp., Ms 2712, VI, Bibliotheque Centrale du Museum National d'Histoire Naturelle. 29. Etienne Geoffroy Saint-Hilaire, "Memoire dans lequel on compare les organes de la respiration des diverses classes d'animaux, on evalue la force de compression qui y condense Pair, on exprime la quantite d'oxigene qui s'y degage pendant la respiration et dans lequel on recherche les causes qui engourdissent les reptiles et certaines mammiferes," 37pp., Ms 2712, X, Bibliotheque Centrale du Museum National d'Histoire Naturelle. 30. In September 1801, Geoffroy wrote to Cuvier, "Is it because I have inundated you with my letters that I have merited your writing to me never . . . " Geoffroy, Lettres ecrites d'Egypte, p. 202. 31. Ibid., p. 195. 32. Etienne Geoffroy Saint-Hilaire, Etudes progressives d'un naturaliste pendant les annees 1834 et 1835 (Paris, 1835), pp. 149-151. On the importance of electric fish as a possible key to the nature of nervous action, see Roderick W. Home, "Electricity and the Nervous Fluid," Journal of the History of Biology 3 (1970): 235-251, esp. pp. 249-250. 33. Etienne Geoffroy Saint-Hilaire, "Du calorique considere comme principe vital et de ses lois de circulation dans les corps organises," 37pp., Ms 2722, III, Bibliotheque Centrale du Museum National d'Histoire Naturelle; idem, "Ma physique, ou systeme d'idees sur le fluide nerveux et subsidiairement sur les elemens reels des corps," 75pp., Ms 2722, I, Bibliotheque Centrale du Museum National d'Histoire Naturelle. See also Laissus, "Manuscrits de Geoffroy conserves au Museum," pp. 379-380. 34. Etienne Geoffroy Saint-Hilaire, Notions synthetiques, historiques et physiologiques de philosophie naturelle (Paris, 1838), pp. xxv-xxviii. 35. On the Laplacian program, see Robert Fox, "The Rise and Fall of Laplacian Physics," in Historical Studies in the Physical Sciences, vol. 4, ed. Russell McCormmach (Princeton: Princeton University Press, 1975), pp. 89-136. 36. Buffon in 1765 had taken up the program of the motion of the smallest particles of matter, arguing against the mathematician Alexis-Claude Clairaut that these particles still obeyed a Newtonian formula for attraction. See John Lyon and Phillip R. Sloan, eds., From Natural History to the History of Nature: Readings from Buffon and His Critics (Notre Dame, Ind.: University of Notre Dame Press, 1981), pp. 77-85. 37. Geoffroy, "Du calorique," p. 4. On Geoffroy's system, see Geoffroy, Lettres ecrites d'Egypte, pp. 217-220. 38. Geoffroy, Lettres ecrites d'Egypte, pp. 219-220. 39. Ibid., p. 205. 40. Ibid., p. 216. 41. Ibid., p. 220. 42.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 104-108. 43. Geoffroy, "Du calorique," p. 35. 44. Lamarck, Cuvier, and Lacepede, "Rapport," p. 235. The report was read by Lacepede. 45. Ibid., p. 236. 46. Jean-Baptiste Lamarck, Philosophie zoologique (Paris, 1809), pp. 69-71, quotation on p. 69. 47. Georges Cuvier, Recherches sur les ossemens fossiles de quadrupedes, 4 vols. (Paris, 1812), "Discours preliminaire," 1: 79-81. 48. Charles Lyell, Principles of Geology, 3 vols. (London, 1830-1833), 2: 28-31. 49. [Etienne Geoffropy Saint-Hilaire] Catalogue des mammiferes du Museum National d'Histoire

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Naturelle [Paris, 1803]. This was an uncharacteristic book for Geoffroy: a straightforward listing of Museum specimens according to order, genus, and species. One novelty of the classification was the creation of separate orders for marsupials ("Pedimanes") and monotremes (the name was Geoffroy's). I consulted a photostatic copy of this work in the Division of Mammalogy, National Museum of Natural History, Smithsonian Institution. See also I. Geoffroy, Etienne Geoffroy SaintHilaire, pp. 115-119. 50.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 116-117. 51. Etienne Geoffroy Saint-Hilaire, "Histoire naturelle et description anatomique d'un nouveau genre de poisson du Nil, nomme Polyptere," Annales du Museum d'Histoire Naturelle 1 (1802): 57-68; idem, "Sur les branchies du Silurus anguillaris," Bulletin de la Societe Philomatique (1802): 105; idem, "Observations anatomiques sur le Crocodile du Nil," Annales du Museum d'Histoire Naturelle 1 (1803): 37-52. 52. Etienne Geoffroy Saint-Hilaire, "Memoire sur 1'anatomie comparee des organes electriques de la raie torpille, du gymnote engourdissant, et du silure trembleur," Annales du Museum d'Histoire Naturelle 1 (1802): 392-407, quotation on p. 394. 53. Proces-verbaux, 1: 24, 27-28, 45, 67, 68. 54. Ibid., 1:395. 55. Ibid., 2: 101, 109. 56. Henri Deherain, "Lettres a G. Cuvier sur 1'organisation de 1'Institut en 1'an XI," Journal des savants (1916): 368-376, esp. pp. 372-373. 57. Proces-verbaux, 2: 643. Apparently there was an argument at the Institut on whether human anatomists belonged in the section of anatomy and zoology (ibid., 2: 640). 58. Georges Cuvier, Lecons d'anatomie comparee, 5 vols. (Paris, 1800-1805), 1: 331-337 (on the pectoral fin), 1: 248-250 (on the furcula), 2: 15-32 (on bones of the skull), and 4: 378-379 (on the operculum); Etienne Geoffroy Saint-Hilaire, "Premier memoire sur les poissons, ou 1'on compare les pieces osseuses de leurs nageoires pectorales avec les os de 1'extremite anterieur des autres animaux a vertebres," Annales du Museum d'Histoire Naturelle 9 (1807): 357-372, esp. pp. 360361. 59. France. Commission des Sciences et Arts d'Egypte, Description de I'Egypte; ou, Recueil des observations et des recherches qui ont ete faites en Egypte pendant I'expedition de I'armeefrancaise. Histoire naturelle, 2 vols. (Paris, 1809-1813). The volumes, although dated 1809 and 1813, actually appeared in 1826. Geoffroy left much of the description of new species to his son and to Victor Audouin. 60. Etienne Geoffroy Saint-Hilaire, Principes de philosophie zoologique, discutes en mars 1830, au sein de I'Academie Royale des Sciences (Paris, 1830), pp. 127-128. See also the article on the Tetraodon physa in Description de I'Egypte, 1: 19-37. 61. For a comparison of Geoffroy's doctrine with that of Aristotle, see Georgette Legee, "Les lois de 1'organisation d'Aristote a Geoffroy Saint-Hilaire," Histoire et nature 1 (1973): 3-25. 62. Geoffroy, "Premier memoire sur les poissons," p. 358. 63. Ibid., p. 370. 64. Ibid., p. 370. 65. Etienne Geoffroy Saint-Hilaire, "Second memoire sur les poissons: considerations sur 1'os furculaire, une des pieces de la nageoire pectorale," Annales du Museum d'Histoire Naturelle 9 (1807): 413-427. 66. Etienne Geoffroy Saint-Hilaire, "Troisieme memoire sur les poissons, ou lon traite de leur sternum sous le point de vue de sa determination et de ses formes generates, " Annales du Museum d'Histoire Naturelle 10 (1807): 87-104, quotations on pp. 88, 89, 90. 67. Geoffroy, "Premier memoire sur les poissons," p. 364. 68. Etienne Geoffroy Saint-Hilaire, "Considerations sur les pieces de la tete osseuse des animaux vertebres, et particulierement sur celles du crane des oiseaux," Annales du Museum d'Histoire Naturelle 10 (1807): 342-365; idem, "Determination des pieces qui composent le crane des crocodiles," Annales du Museum d'Histoire Naturelle 10 (1807): 249-264. 69. Geoffroy, "Considerations sur les pieces de la tete osseuse," p. 344. 70. Ibid., p. 344. 71. Ibid., pp. 342-343.

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72. Ibid., p. 345. 73.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 131-133. 74. J.H.F. Kohlbrugge, Historische-kritische Studien uber Goethe ah Naturforscher (Wurzburg: Curt Kabitzsch, 1913), pp. 61-113. 75. On Naturphilosophie, see below, chapter 5. 76. Franck Bourdier, "Geoffroy Saint-Hilaire versus Cuvier: The Campaign for Paleontological Evolution (1825-1838)," in Cecil J. Schneer, ed., Toward a History of Geology (Cambridge, Mass.: M.I.T. Press, 1969), pp. 36-61, esp. p. 37; F. Dagognet, "Valentin Hauy, Etienne Geoffroy SaintHilaire, Augustin P. de Candolle: une conception d'ensemble mais aussi un ensemble de conceptions," Revue d'histoire des sciences 25 (1972): 327-336. 77. Proces-verbaux, 3: 523-525 [4 May 1807], 564-565 [24 August 1807], 575-576 [7 September 1807]. 78. Ibid., 3: 525, 575. 79. Ibid., 3: 575-576. 80. Ibid., 3: 523. 81. Ibid., 3: 565. 82. Ibid., 3: 576. 83. Ibid., 3: 579. 84. Ibid., 3: 585. 85. Pierre Flourens, "Eloge historique d'Etienne Geoffroy Saint-Hilaire," Memoires de I'Academie des Sciences 23 (1853): i-ixxi, esp. p. ix. See also Cuvier, Manuscript autobiography, p. 34. 86. On the mission to Portugal, see I. Geoffroy, Etienne Geoffroy Saint-Hilaire, chapter 6; and E.T. Hamy, "La mission de Geoffroy Saint-Hilaire en Espagne et en Portugal (1808): histoire et documents," Nouvelles Archives du Museum d'Histoire Naturelle, 4th series, 10 (1908): 1-58. 87.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 190-191. 88. "Obseques de M. Geoffroy Saint-Hilaire," Gazette medicale de Paris 12 (1844): 409-422, quotation on pp. 415-416. 89. Geoffroy, Philosophie anatomique, p. xxxix. 90. Jules-Cesar Savigny, Memoires sur les animaux sans vertebres (Paris, 1816), p. iii. On Savigny, see Mary P. Winsor, "Marie-Jules-Cesar Lelorgne de Savigny," DSB, 12: 130-131; E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology (London: John Murray, 1916), reprint ed. (Chicago: University of Chicago Press, 1982), pp. 83-85; and Paul Pallary, "Marie-Jules-Cesar Savigny: sa vie et son oeuvre," Memoires presentes a I'lnstitut d'Egypte 17 (1931), 20 (1932), and 23 (1934), esp. 23 (1934). This work notes that after their return from Egypt, Savigny and Geoffroy engaged in an unpleasant dispute over the division of labor in writing up the materials from the expedition. Apparently they were later reconciled. 91. Savigny, "Observations sur la bouche des papillons, des phalenes et des autres insectes lepidopteres, suivies de quelques considerations sur la bouche des dipteres, des hemipteres at des apteres suceurs," in idem, Memoires sur les animaux sans vertebres, pp. 1-37. See also the extract of this memoir in Bulletin de la Societe Philomatique (1814): 168-170; and the report to the AcademiE read by Lamarck in Proces-verbaux, 5: 408-411. 92. Savigny, "Observations generales sur la bouche des arachnides, des crustaces et des entomostraces," in idem, Memoires sur les animaux sans vertebres, pp. 39-118, quotation on p. 47. See also the report to the Academie by Lamarck, Cuvier, and Latreille in Proces-verbaux, 5: 521526. 93. Henri Marie Ducrotay de Blainville, "Prodrome d'une nouvelle distribution systematique du regne animal," Journal de physique 83 (1816): 244-267, esp. p. 265. This memoir was also published in Bulletin de la Societe Philomatique (1816): 105-124. 94. Henri Marie Ducrotay de Blainville, "Considerations generales sur le squelette des animaux vertebres et articules," Bulletin de la Societe Philomatique (1817): 109; idem, "Note sur les animaux articules," Journal de physique 89 (1819): 467-472; idem, "Sur la concordance des anneaux du corps des Entomozoaires hexapodes adultes," Bulletin de la Societe Philomatique (1820): 3337. 95. Henri Marie Ducrotay de Blainville, "Sur l'existence de veritable ongles a l'aile de quelques especes d'oiseaux," Bulletin de la Societe Philomatique (1819): 41; idem, "Note sur 1'existence des

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nerfs olfactifs dans le dauphin, et par analogie, dans les autres cetaces," Bulletin de la Societe Philomatique (1815): 193-195; idem, "Considerations sur les organes de la generation," Bulletin de la Societe Philomatique (1818): 155-162. On Blainville and philosophical anatomy, see Toby A. Appel, "Henri de Blainville and the Animal Series: A Nineteenth-Century Chain of Being," Journal of the History of Biology 13 (1980): 291-319, esp. pp. 302-307. 96. Henri Dutrochet, "Recherches sur la metamorphose du canal alimentaire chez les insectes," in idem, Memoires pour servir a I'histoire anatomique et physiologique des vegetaux et des animaux, 2 vols. (Paris, 1837), 2: 331-360, quotations on pp. 358-359. 97. Henri Dutrochet, "Observations sur la structure et la regeneration des plumes, avec des considerations generales sur la composition de la peau des animaux vertebres," in idem, Memoires 2: 361-384. 98. On Dutrochet's relationship to Geoffroy, see Joseph Schiller and Tetty Schiller, Henri Dutrochet (Henri du Trochet, 1776-1847): le materialisme mecaniste et la physiologie generale (Paris: Albert Blanchard, 1975). The Schillers include texts of some twenty letters from Dutrochet to Geoffroy. On Dutrochet, see also Vladislav Kruta, "Rene-Joachim-Henri Dutrochet," DSB, 4: 263-265. 99. Constant Dumeril, "Considerations generates sur 1'analogie qui existe entre tous les os et les muscles du tronc dans les animaux," Magasin encydopedique [annee 1808] 3: 111-143. For Geoffroy's comments, see his manuscript lecture notes dated 21 March 1821 in the Geoffroy SaintHilaire Papers, American Philosophical Society; and Etienne Geoffroy Saint-Hilaire, "Composition de la tete osseuse de 1'homme et des animaux," Annales des sciences naturelles 3 (1824): 173192, 246-299, esp. pp. 176-177. See also Pierre Flourens, "Eloge historique d'Andre-Marie Constant Dumeril," Memoires de l'Academic des Sciences 35 (1866): i-xxii, esp. p. xv. A commission was named at the Institut to report on Dumeril's memoir, but no report appears to have been made. But see Cuvier's commentary in Magasin encydopedique [annee 1809] 1: 163-164. Dumeril's early work also included memoirs which bridged Cuvier's four embranchements. In 1812 for the competition for adjoint professor at the Faculte des Sciences, he submitted a monograph on "the fishes which most resemble the invertebrates." There he argued that certain jawless fishes, including lampreys, which he named cyclostomes, differed from all other fishes. They occupied "the last rank" among vertebrates, he claimed, and moreover, he demonstrated by comparative anatomy that some species formed "a veritable natural transition in the scale of animal organization" to the invertebrates. See Constant Dumeril, Dissertation sur les poissons qui se rapprochent le plus des animaux sans vertebres (Paris, 1812), 40pp, esp. pp. 7-8. 100. Augustin-Pyramus de Candolle, Theorie elementaire de la botanique (Paris, 1813), esp. pp. 77-123. On Candolle and "philosophical botany," see Michel Guedes, "La theorie de la metamorphose en morphologie vegetate: A.-P. de Candolle et P.-J.-F. Turpin," Revue d'histoire des sciences 25 (1972): 253-270; Peter J. Stevens, "Hauy and A.-P. de Candolle: Crystallography in the Development of Botanical Systematics and Comparative Morphology, 1780-1840," Journal of the History of Biology 17 (1984): 49-82; and Alphonse de Candolle, Memoires et souvenirs de Augustin-Pyramus de Candolle (Geneva, 1862), pp. 214-216. 101. Georges Cuvier, "Sur la composition de la tete osseuse dans les animaux vertebres," Annales du Museum d'Histoire Naturelle 19 (1812): 123-128, quotations on p. 123. 102.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 198-201. 103. Geoffroy recounted the steps leading up to his discovery of the homologues of the operculum in Philosophie anatomique, pp. 15-30. 104. Henri Marie Ducrotay de Blainville, "Memoire sur 1'opercule des poissons," Bulletin de la Societe Philomatique (1817): 104-112. 105. Geoffroy, Principes de philosophie zoologique, p. 129. 106. Etienne Geoffroy Saint-Hilaire, "Du squelette des poissons ramene dans toutes ses parties a la charpente osseuse des autres animaux vertebres, et premierement de 1'opercule des poissons," Bulletin de la Societe Philomatique (1817): 125-127; idem, Philosophie anatomique, pp. 29-30. 107. Isidore Geoffroy, for example, always referred to the search for homologies as "philosophical anatomy." See I. Geoffroy, Etienne Geoffroy Saint-Hilaire, chapter 8. One of the reviewers, A.-J.-L. Jourdan, specifically faulted Geoffroy for the title, since the work did not present a philosophy but only the materials for one (see below, note 119, p. 212). 108. Geoffroy, Philosophie anatomique, pp. xv, xxxii. Isidore Geoffroy insisted that Geoffroy's

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method (theory of analogues) was distinct from his theory (unity of organic composition) and that supporters of the Cuvierian school had confounded them. It seems that Geoffroy himself confounded them. I. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 217-220. 109. Geoffroy, Philosophie anatomique, p. xviii. 110. Ibid., p. xxviii. 111. Ibid., p. xxxviii. 112. Ibid., p. xxx. This sentence was often repeated in Geoffroy's writings as well as in contemporary discussions of Geoffroy's work. 113. Ibid., pp. 4-11. For an explanation of Geoffroy's use of metastasis as well as a useful analysis of Philosophie anatomique, see Russell, Form and Function, chapter 5. 114. Ibid., p. 37. 115. Ibid., pp. 51-54. Geoffroy's particular homology is not now considered valid, though his belief that homologies ought to exist was justified. In the period before Darwin, a considerable number of naturalists took up the problem of the homologies of the opercular bones. For the resolution of this question, see below, chapter 8. 116. Ibid., p. xi. 117. Ibid., pp. 57-137. Geoffroy had previously named the bones of the sternum in his article on the tortoise in Description de I'Egypte. For a good discussion of Geoffroy's treatment of the sternum, see Russell, Form and Function, pp. 56-60. 118. Geoffroy, Philosophie anatomique, p. xxix. 119. Ibid., pp. 284-304, quotation on p. 289. Reviews of Philosophie anatomique include that of Etienne Pariset in Revue encydopedique 3 (1819): 32-40; that of A.-J.-L. Jourdan in Journal universel des sciences medicales 15 (1819): 67-88, 196-213; that of Frederic Cuvier in Journal complementaire des sciences medicales 3 (1819): 142-157; that of U. Coste in Bibliotheque medicale 65 (1819): 178-195; and Pierre Flourens, Analyse de la Philosophie anatomique, ou I'on considere plus particulierement l'influence qu'aura cet ouvrage sur I'etat actuel de la Physiologie et de I'Anatomie, 28pp. (Paris, 1819), also published as "Essai sur l'esprit et sur 1'influence de la Philosophie anatomique," Revue encydopedique 5 (1820): 217-232. Jourdan (op. cit, p. 211) commented, "Despite the care that he takes to excuse himself, this digression is entirely out of place . . ." Pariset (op. cit., p. 40) noted that Geoffroy had written as if convinced, but he failed to provide proof. F. Cuvier (op. cit., p. 155) refused to comment on the doctrine of sound because it would take him too far from his main subject. 120. Georges Cuvier, "Analyse des travaux de l'Academie Royale des Sciences, pendant 1'annee 1818: partie physique," Memoirs de I'Academic des Sciences 3(1818): clxxix-ccxxx, esp. pp. ccxiiiccxviii, quotation on p. ccxiv. 121. Georges Cuvier, "Analyse des travaux de 1'Academie Royale des Sciences pendant 1'annee 1817," Memoirs de I'Academic des Sciences 2 (1817): xciii-cixix, esp. pp. cxviii-cxxcii, quotation on p. cxx. 122. Cuvier, "Analyse . . . 1818," p. ccxvi. 123. Cuvier, "Analyse . . . 1817," pp. cxxiv-cxxv. 124. Ibid., p. cxxvii. 125. Pariset, Review of Philosophie anatomique, p. 33. 126. Ibid., p. 33. 127. Jourdan, Review of Philosophie anatomique, p. 209n. 128. Ibid., p. 212. 129. F. Cuvier, Review of Philosophie anatomique, p. 147. 130. Ibid., p. 157. 131. See note 119. Georgette Legee, "Cuvier (1769-1832), Geoffroy Saint-Hilaire (1772-1844) et Flourens (1794-1867)," Histoire et biologie 2 (1969): 10-34, esp. p. 11. Although most writers on Flourens have neglected the patronage and influence of Geoffroy, Legee is an exception. On Flourens's career, see below, chapter 5. 132. Flourens, Analyse de la Philosophie anatomique, pp. 5, 6. 133. Ibid., p. 11. 134. Ibid., p. 14. "It is perfectly certain that no one has ever proved anything by them [final causes], except their very inability to prove anything," Flourens wrote (p. 28).

266

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Chapter 5

1. Georges Cuvier and Achille Valenciennes, Histoire naturelle des poissons, vol. 1 (Paris, 1828), pp. 1, 551. 2. Etienne Geoffroy Saint-Hilaire, Cows de I'histoire naturelle des mammiferes (Paris, 1829), "Discours preliminaire," pp. 20-21. This work was also published as Lecons sur I'histoire naturelle des mammiferes (Paris, 1828). The lectures are separately paginated. 3. Lenoir distinguishes three traditions of Naturphilosophie: transcendental Naturphilosophie (the Gottingen school), speculative or romantic Naturphilosophie (Schelling, Oken, Goethe), and metaphysical Naturphilosophie (Hegel). See Timothy Lenoir, "The Gottingen School and the Development of Transcendental Naturphilosophie in the Romantic Era," in Studies in History of Biology, vol. 5, ed. William Coleman and Camille Limoges (Baltimore: Johns Hopkins University Press, 1981), pp. 111-205, esp. pp. 112-115. 4. Geoffroy and Serres ascribed the first use of the term "homology" to German anatomists, but they did not cite any specific source. Even Richard Owen, who gave the term its modern meaning, did not know where the word came from and could only refer to Geoffroy. See Geoffroy's lecture notes dated 10 March 1821, Geoffroy Saint-Hilaire Papers, American Philosophical Society, where Geoffroy compared the "theory of homologues" to the "theory of analogues"; Etienne Geoffroy Saint-Hilaire, "Composition de la tete osseuse de 1'homme et des animaux," Annales des sciences naturelles 3 (1824): 173-192, 245-299, esp. p. 174; and Richard Owen, On the Archetype and Homologies of the Vertebrate Skeleton (London, 1848), p. 5. 5. See Lorenz Oken, Elements of Physiophilosophy, trans. Alfred Tulk (London, 1847), pp. 363384. On German transcendental anatomy, see E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology (London: John Murray, 1916), reprint ed. (Chicago: University of Chicago Press, 1982), pp. 89-101. 6. On the philosophical foundations of German biology, see Timothy Lenoir, "Kant, Blumenbach, and Vital Materialism in German Biology," Isis 71 (1980): 77-108; idem, The Strategy of Life: Teleology and Mechanics in Nineteenth-Century German Biology (Dordrecht: D. Reidel, 1982), esp. pp. 1-61; idem, "The Gottingen School"; and L. Pearce Williams, "Kant, Naturphilosophie and Scientific Method," in Ronald N. Giere and Richard S. Westfall, eds., Foundations of Scientific Method: The Nineteenth Century (Bloomington, Ind.: University of Indiana Press, 1973), pp. 3-22. 7. For information on the translations of Geoffroy's memoirs into German, see Royal Society of London, Catalogue of Scientific Papers (1800-1863), 2: 824-832. 8. References to political events of the Restoration period are from Alfred Cobban, A History of Modern France. Volume 2:1799-1871 (Baltimore: Penguin Books, 1961); Frederick B. Artz, Reaction and Revolution: 1814-1832 (New York: Harper & Row, 1934) and Guillaume de Bertier de Sauvigny, The Bourbon Restoration, trans. Lynn M. Case (Philadelphia: University of Pennsylvania Press, 1966). 9. See, for example, Mary Katherine Lyell, Life, Letters and Journals of Sir Charles Lyell, Bart., 1 vols. (London, 1881), 1: 126-127, 136-137. Lyell, visiting Paris in 1823, wrote to his father, "Cuvier is himself notoriously considered an Ultra, and I fear with too much reason, one of the most mischievous Ultras in the present Conseil d'Etat." 10. On the vertebral theory of the skull, see Geoffroy, "Composition de la tete osseuse." 11. Geoffroy argued that the connections of the Weberian ossicles were with the vertebrae, and that therefore the bones were vertebral branches. Etienne Geoffroy Saint-Hilaire, "Sur une nouvelle determination de quelques pieces mobiles chez la carpe, ayant ete considerees comme les parties analogues des osselets de 1'oreille; et sur la necessite de conserver le nom de ces osselets aux pieces de 1'opercule," Memoires du Museum d'Histoire Naturelle 11 (1824): 143-160. The Weberian ossicles are now in fact believed to be derived from the adjacent vertebrae. It is thought that they convey vibrations from the swim bladder to the ear, a function analogous to that of the ear ossicles. 12. Proces-verbaux, 6: 521, 7: 4, 6, 7, 9, 11-12, 13. 13. Etienne Geoffroy Saint-Hilaire, "Memoires sur 1'organisation des insectes. Premier memoire sur un squelette chez les insectes, dont toutes les pieces identiques entre elles, dans les divers ordres

NOTES

267

du systeme entomologique, correspondent a chacun des os du squelette dans les classes superieures," Journal complementaire des sciences medicales 5 (1820): 340-351, quotation on p. 341, also published in Annales generales des sciences physiques 3 (1820): 165-180. On Geoffroy's research on the skeleton of insects, see Russell, Form and Function, pp. 60-64. 14. Geoffroy, "Premier memoire," pp. 344-345, quotation on p. 345; Etienne Geoffroy SaintHilaire, "Troisieme memoire: sur une colonne vertebrale et de ses cotes dans les insectes apiropodes," Journal complementaire des sciences medicales 6 (1820): 138-168, also published in Annales generales des sciences physiques 5 (1820): 96-132. 15. Etienne Geoffroy Saint-Hilaire, "Considerations generales sur la vertebre," Memoires du Museum d'Histoire Naturelle 9 (1822): 89-119. For an example of a post-Darwinian physiologist tackling the same problem, see Gerald L. Geison, "Walter Holbrook Gaskell," DSB, 5: 279-284. 16. Geoffroy, "Premier memoire," pp. 346-347; idem, "Troisieme memoire," p. 150. 17. Geoffroy, "Considerations generales sur la vertebre," opposite pp. 115, 119. 18. Ampere wrote a three-part anonymous article on homologies between articulates and vertebrates that was mistakenly attributed to Audouin as well as to Geoffroy in Royal Society of London, Catalogue of Scientific Papers (1800-1863), 1: 118, 2: 827. See [Andre-Marie Ampere] "Considerations philosophiques sur la determination du systeme solide et du systeme nerveux des animaux articules," Annales des sciences naturelles 2 (1824): 295-310, 3 (1824): 199-203,453-456; and Francois Arago, "Ampere," in Oeuvres de Francois Arago, vol. 2 (Paris, 1854), pp. 1-116, esp. pp. 70-75. J.F. Meckel was among those who publicly objected to Geoffroy's homologies. Geoffroy responded to Meckel in a polemical article, "Systeme intra-vertebrale des insectes," Annales de la medecine physiologique 3 (1823): 233-252. 19. Geoffroy, "Troisieme memoire," p. 166. 20. Etienne Geoffroy Saint-Hilaire, "Second memoire: sur quelques regies fondamentales en philosophie naturelle," Journal complementaire des sciences medicales 6 (1820): 31 -36, quotation on p. 34n, also published in Annales generales des sciences physiques 3 (1820): 263-268. See also Geoffroy, "Premier memoire," p. 350, where Geoffroy sees Cuvier as condemning his views. 21. Georges Cuvier, "Analyse des travaux de 1'Academie Royale des Sciences, pendant 1'annee 1820. Partie physique," Memoires de I'Academie des Sciences 4 (1819-1820): ccxxix-cccxvii, esp. pp. cclxii-cclxxiii. 22. Geoffroy, "Troisieme memoire," p. 168. 23. On Latreille's career, see Richard W. Burkhardt, Jr., "Pierre-Andre Latreille," DSB, 8: 4849; A.-J.-L. Jourdan, "Latreille," Biographie universelle, 70: 350-354; and "Latreille," Nouvelle biographie generale, 29, columns 850-854. 24. Lamarck's report to the Institut on Savigny's first memoir on insects noted that Latreille had foreshadowed some of the ideas. See Proces-verbaux, 5: 408-409. 25. Pierre-Andre Latreille, "De la formation des ailes des insectes, et de 1'organisation exterieure des ces animaux, comparee, en divers points avec celles des arachnides et des crustaces," pp. 121. This article was published with "Passage des animaux invertebres aux vertebres," pp. 22-40, in a 44 page pamphlet (Paris, n.d. [1820]) which was intended to be a supplement to his Memoires sur divers sujets d'histoire naturelle des insectes, de geographie ancienne et de chronologie (Paris, 1819). A copy of the pamphlet is bound with the book at the New York Public Library. The "Supplement," pp. 40-41, tried to moderate the claims of the articles by stating that there was still a "hiatus that is now impossible for us to fill" between the lowest fish and the highest crustacean (p. 41). 26. Latreille, "De la formation des ailes des insectes," pp. 3-4. 27. Latreille, "Passage des animaux invertebres." See also Russell, Form and Function, p. 86. 28. Cuvier, "Analyse . . . 1820," p. cclxxvi. 29. Ibid., p. cclxxiii. 30. Pierre-Andre Latreille, "De 1'organisation exterieure des cephalopodes, comparee avec celle de divers poissons," Memoires de la Societe d'Histoire Naturelle de Paris 1 (1823): 269-272, esp. p. 271. 31. "Latreille," Nouvelle biographie generale, 29: column 851. 32. Latreille to Cuvier, 30 August 1831, Fonds Cuvier, Ms. 3253.32, Bibliotheque de 1'Institut de France.

268

NOTES

33. On Audouin's career, see Jean Theodorides, "Jean Victor Audouin," DSB, 1:328-329; Henri Milne Edwards, Notice sur la vie et les travaux de Victor Audouin (Paris, 1850), 22pp.; and M. Duponchel, "Notice sur la vie et les travaux de Jean-Victor Audouin," Annales de la Societe Entomologique de France 11 (1842): 95-164. 34. Jean-Victor Audouin, Journal d'un etudiant en medecine et en sciences a Paris sous la Restauration (1817-1818), ed. Jean Theodorides (Paris, 1959), pp. 102-103. This work also appeared in Histoire de la medecine 9 (Nov. 1958): 4-63, 9 (Dec. 1958): 5-56, 10 (Jan. 1959): 5-48. 35. Etienne Geoffroy Saint-Hilaire, "Rapport lu a 1'Academie des Sciences dans sa seance du fevrier 1820, sur un memoire de M. Audouin concernant l'organisation des insectes," Journal complementaire des sciences medicales 6 (1820): 36-42; Report of Lacepede, Cuvier, Bosc and Dumeril, Proces-verbaux, 7: 32-33. 36. Geoffroy, "Rapport . . . sur un memoire de M. Audouin," p. 42; Proces-verbaux, 7: 11-12. 37. Proces-verbaux, 1: 33. 38. Jean-Victor Audouin, "Recherches anatomiques sur le thorax des animaux articules et celui des insectes hexapodes en particulier," Annales des sciences naturelles 1 (1824): 97-135, 416-432, quotation on p. 99. See also Russell, Form and Function, pp. 85-86. 39. Audouin, "Recherches anatomiques," p. 100 40. Ibid., p. 116. 41. Ibid., p. 110. 42. Ibid., pp. 105-106. 43. Audouin to Cuvier, 17 July 1820, Fonds Cuvier, Ms. 3242, Bibliotheque de 1'Institut de France. 44. Georges Cuvier, "Rapport sur une memoire de M. Audouin, intitule, 'Recherches anatomiques sur le thorax des animaux articules, et celui des insectes en particulier'," Annales generales des sciences physiques 7 (1821): 396-406, quotation on p. 397. Cuvier's report also appeared in Proces-verbaux, 7: 144-147. 45. Ibid., pp. 398, 399. 46. Audouin, "Recherches anatomiques," p. 104. 47. On this election, see Pol Nicard, "Etude sur la vie et les travaux de M. de Blainville," in Henri Marie Ducrotay de Blainville, Osteographie, ou, Description iconographique comparee du squelette et du systeme dentaire des mammiferes recents et fossiles pour servir de base a la zoologie et a la geologie, 4 vols. (Paris, 1839-1864), 1: i-ccxxiii, esp. pp. lxxxiv-lxxxix. 48. On Flourens, see Vladislav Kruta, "Marie-Jean-Pierre Flourens," DSB, 5: 44-45; Alfred Vulpian, "Eloge historique de M. Flourens," Memoires de 1'Academie des Sciences 44 (1888): cxlixclxxxiv; and J.M.D. Olmsted, "Pierre Flourens," in E.A. Underwood, ed., Science, Medicine and History, vol. 2 (London: Oxford University Press, 1953), pp. 290-302. All of these sources recognize Cuvier's role in promoting Flourens's career, but none mention Geoffroy's influence. One historian who does discuss Flourens's shift of allegiance is Georgette Legee, "Cuvier (1769-1832), Geoffroy Saint-Hilaire (1772-1844) et Flourens (1794-1867)," Histoire et biologie 2 (1969): 10-34. 49. Robert M. Young, "Franz Joseph Gall," DSB, 5: 250-256, esp. p. 250. For an example of Geoffroy's favorable response to Gall, see Etienne Geoffroy Saint-Hilaire, Philosophie anatomique, vol. 2., Des monstruosites humaines (Paris, 1822), reprint ed. (Brussels: Culture et Civilisation, 1968), pp. 17, 17n. 50. Geoffroy, Philosophie anatomique, 2: ix; Etienne Geoffroy Saint-Hilaire, "La Zoologie a-telle, dans 1'Academie des Sciences, une representation suffisant? La Physiologie n'y a-t-elle pas ete entierement oubliee?," Revue encyclopedique 13 (1822): 501-511. 51. For Cuvier's critique of Flourens's terminology, see Georges Cuvier, "Rapport sur le memoire de M. Flourens lu dans la seance du 22 juillet 1822," Proces-verbaux, 7: 349-354. For an appreciation of Cuvier's influence on the physiological doctrine of Flourens and also of Magendie, see Michael Gross, "The Lessened Locus of Feelings: A Transformation in French Physiology in the Early Nineteenth Century," Journal of the History of Biology 12 (1979): 231-271, esp. pp. 252258. 52. Henri Marie Ducrotay de Blainville, "Prodrome d'une nouvelle distribution systematique du regne animal," Journal de physique 83 (1816): 244-267. This memoir also appeared in Bulletin de la Societe Philomatique (1816):105-124.

NOTES

269

53. Henri Marie Ducrotay de Blainville, De I'organisation des animaux; ou, Principes d'anatomie comparee (Paris, 1822), Tables. 54. Ibid., pp. xvi, xxi, Iv-lvii, 16. 55. Ibid., pp. 4-5, quotation on p. 5. 56. On the significance of Laplace and the Laplacian program, see Maurice Crosland, The Society of Arcueil: A View of French Science at the Time of Napoleon I (Cambridge, Mass.: Harvard University Press, 1967); Robert Fox, "The Rise and Fall of Laplacian Physics," in Historical Studies in the Physical Sciences, vol. 4, ed. Russell McCormmach (Princeton: Princeton University Press, 1975), pp. 89-136; Eugene Frankel, "J.B. Biot and the Mathematization of Experimental Physics in Napoleonic France," in ibid., vol. 8, ed. Russell McCormmach and Lewis Pyenson (Baltimore: Johns Hopkins University Press, 1977), pp. 33-72; and idem, "Corpuscular Optics and the Wave Theory of Light: The Science and Politics of a Revolution in Physics," Social Studies of Science 6 (1976): 141-184. 57. Roger Hahn, "Dominique Francois Jean Arago," DSB, 1: 200-203. 58. Frankel, "Corpuscular Optics," pp. 171-173. 59. Flourens, not yet a member of the Academie, who wrote analyses of the work of the Academie for Annales generales des sciences physiques, noted in 1820 (3: 244): "According to the regulations, these meetings are supposed to be secret. The Academie has, however, judged it a propos to stray on this point from the letter of the law and to admit a certain number of young savants." On the debate over regulations in 1825, see Proces-verbaux, 8: 206, 210, 214. 60. Proces-verbaux (see chapter 3, note 97). The Comptes rendus began to appear in 1835. 61. Proces-verbaux, 8: 308. 62. On this election, see Nicard, "Etude sur Blainville," pp. lix, ccxxiii, quotation on p. lxix. 63. On Serres, see Elizabeth B. Gasking, "Antoine Etienne Reynaud Augustin Serres," DSB, 12: 315-316; and "Serres," Nouvelle biographie generale, 43: columns 797-799. On Geoiffoy's initial encounter with Serres, see Alfred Moquin-Tandon, Un naturaliste a Paris sous Louis-Philippe: journal de voyage inedit (1834), 2nd. ed. (Paris: Mercure de France, 1944), pp. 113-114. 64. Proces-verbaux, 6: 508, 509, 7: 21-22, 34, 130, 157, 165. His first paper on osteogenesis appears not to have been published as such. 65. Geoffroy reported on Serres's memoir on the laws of osteogenesis in 1819, on his research on the comparative anatomy of the brain in 1824, and on his work on animal monsters in 1825. Proces-verbaux, 6: 112-113, 127, 304-307; Journal complementaire des sciences medicales 3 (1819): 67-72; ibid. 19 (1824): 148-161; and Revue encydopedique 23 (1824): 324-340. 66. On Serres's efforts to obtain a seat in the Academie, see Proces-verbaux, 7: 210, 211, 8: 184, 186, 204, 206, 307, 308, 458, 459, 9: 95, 96. 67. The following discussion of Serres's doctrine is based upon E.R.A. Serres, Anatomie comparee du cerveau, dans les quatre classes des animaux vertebres, applique a la physiologie et a la pathologie du systeme nerveux, 2 vols. (Paris, 1824-1826); idem, "Recherches d'anatomie transcendante, sur les lois de 1'organogenie appliquees a l'anatomie pathologique," Annales des sciences naturelles 11 (1827): 47-70; idem, Recherches d'anatomie transcendante et pathologique. Theorie des formations et des deformations organiques, appliquee a I'anatomie de Ritta-Christina, et de la duplicite monstrueuse (Paris, 1832); and on Geoffroy's report on Serres's memoir, "Des lois de 1'osteogenie," in Proces-verbaux, 6: 415-417. 68. See, for example, Blainville, De I'organisation des animaux, pp. 4-5. 69. Serres, Anatomie comparee du cerveau, "Discours preliminaire," 1: Ixvii-lxviii. 70. Ibid., 1: lxviii. 71. See Stephen Jay Gould, Ontogeny and Phylogeny (Cambridge, Mass.: Harvard University Press, 1977), pp. 47-52; and Russell, Form and Function, pp. 79-83. 72. Serres, Anatomie comparee du cerveau, "Discours preliminaire," 1: liv-lv. 73. Ibid., 2: 24. See also E.R.A. Serres, "Explication du systeme nerveux des animaux invertebres," Annales des sciences naturelles 3 (1824): 377-380, esp. p. 380. Serres argued that the nervous system of invertebrates corresponded to the intervertebral or spinal ganglia of the nervous system of vertebrates. 74. Serres, Recherches d'anatomie transcendante, p. 9. 75. Ibid., "Dedicace."

270

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76. On Meckel, embryology, and teratology, see Gould, Ontogeny and Phylogeny, pp. 45-47; Guenter B. Risse, "Johann Friedrich Meckel," DSB, 9: 252-253; Owen E. Clark, "The Contributions of J.F. Meckel, the Younger, to the Science of Teratology," Journal of the History of Medicine 24 (1969): 310-322; and Lenoir, Strategy of Life, pp. 56-61. 77. Serres, "Recherches d'anatomie transcendante," pp. 49-51, quotation on p. 50; Serres, Anatomie comparee du cerveau, "Preface," 1: x-xi, "Discours preliminaire," 1: xviii-xxii. 78. Georges Cuvier, "Rapport fait a 1'Academie des Sciences en mars 1821," in Serres, Anatomie comparee du cerveau, "Preface," 1: xviii-xxxviii. 79. Moquin-Tandon, Un naturaliste a Paris, p. 114. Moquin-Tandon described Serres as "a bit cold" in his manner and not especially attractive in appearance, but nonetheless esteemed him for his frankness, his reliability, and his lack of the petty jealousies that characterized most of the savants of the time. Ibid., pp. 113, 117-118. 80. Ibid., pp. 114-115. 81. The American comparative anatomist Jeffries Wyman, who attended the courses of Serres and Isidore Geoffroy in 1841, commented, "Serres and St Hilaire give you but little else than the famous laws of 'arrest of development' and 'unity of composition.'" George E. Gifford, ed., "An American in Paris, 1841-1842: Four Letters from Jeffries Wyman," Journal of the History of Medicine 22 (1967): 274-285, quotation on p. 277. 82. See, for example, Etienne Geoffroy Saint-Hilaire, "Exposition d'un plan d'experiences pour parvenir a la preuve de la co-existence des sexes dans les germes de tous les animaux" (1800), Dossier Etienne Geoffroy Saint-Hilaire, Archives de 1'Academie des Sciences, Institut de France. 83. Isidore Geoffroy Saint-Hilaire, Vie, travaux et doctrine scientifique d'Etienne Geoffroy SaintHilaire (Paris, 1847), reprint ed. (Brussels: Culture et Civilisation, 1968), p. 254. On Geoffroy and teratology, see ibid., chapter 9. 84. Isidore Geoffroy Saint-Hilaire, Histoire generale et particuliere des anomalies de I'organisation chez I'homme et les animaux. . . ou, Traite de teratologie, 3 vols. (Paris, 1832-1836). 85. Blumenbach's biological philosophy, described by Lenoir as "vital materialism" or "teleomechanism," centered on the notion of a "formative force" regulating the development and form of each class of animals. This force was inseparable from matter, yet not reducible to it. Teratology was important to Blumbenbach's research program, because the study of malformations revealed variations in the operation of the force. See Lenoir, "Kant, Blumenbach and Vital Materialism," pp. 78-96; and Lenoir, Strategy of Life, pp. 17-35. Geoffroy adopted the term "formative force" (latinized as nisus formativus) to signify the course of normal development, but had little understanding of the philosophical subtleties of German vital materialism. 86. Etienne Geoffroy Saint-Hilaire, "Considerations d'ou sont deduites des regies pour Pobservation des monstres, et pour leur classification," Annales generales des sciences physiques 8 (1821): 74-87, quotation on p. 74. This paper was reprinted in Geoffroy, Philosophie anatomique, 2: 103123. 87. Ibid., pp. 75-76. 88. Etienne Geoffroy Saint-Hilaire, "Memoire sur plusieurs deformations du crane de l'homme, suivi d'un essai de classification des monstres acepahles," in idem, Philosophie anatomique, 2: 3101, also published in Memoires du Museum d'Histoire Naturelle 1 (1821): 85-162. 89.1. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 283. 90. Geoffroy, "Premier memoir," p. 350n. 91. Clark (note 76) regards Meckel as an adherent of epigenesis. This would contradict Geoffroy and Isidore Geoffroy's insistence on Meckel's preformationism. 92. Geoffroy, Philosophie anatomique, 2: 480. 93. Ibid., p. 489. 94. Geoffroy, Philosophie anatomique, 2: 495, 528. 95. Ibid., p. 15. 96. Etienne Geoffroy Saint-Hilaire, "Considerations generales sur la monstruosite, et description d'un genre nouveau observe dans 1'espece humaine, et nomme Aspalasome," Annales des sciences naturelles 4 (1825): 450-468. 97. Geoffroy, Philosophie anatomique, 2: 493. 98. Etienne Geoffroy Saint-Hilaire, Fragment sur la Nature, ou quelques idees generales du

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271

monde physique considerees d'ensemble et dans I'unite, 27pp. [Paris, 1829], esp. p. 23, extracted from Encyclopedie moderne [1st ed.], vol. 17 (Paris, 1829); idem, "Monstre," in Dictionnaire classique d'histoire naturelle, vol. 11 (Paris, 1827), pp. 108-151, esp. pp. 139-141; idem, Extrait d'un rapport fait a I'Academic des Sciences, le 19 octobre 1829 . . . Sur deux freres attaches ventre a ventre depuis leur naissance. . . , 14pp. (Paris, n.d. [1829]) (I used a copy of this report in the National Library of Medicine, Bethesda, Md.); Proces-verbaux, 8: 332-336, esp. p. 334. 99. On Geoffroy's role in the foundation of a French school of experimental teratology, see Jane M. Oppenheimer, "Some Historical Relationships between Teratology and Experimental Embryology," Bulletin of the History of Medicine 42 (1968): 149-159; idem, "Historical Introduction to Teleostean Development," Osiris 2 (1936): 124-148; Jean Rostand, "E. Geoffroy SaintHilaire et la teratogenese experimentale," Revue d'histoire des sciences 17 (1964): 41-50; Frederick B. Churchill, "Chabry, Roux, and the Experimental Method in Nineteenth-Century Embryology," in Giere and Westfall, eds., Foundations of Scientific Method, pp. 161-205, esp. pp. 181-186; and especially, J.L. Fischer, "Le concept experimental dans 1'oeuvre teratologique d'Etienne Geoffroy Saint-Hilaire," Revue d'histoire des sciences 25 (1972): 347-364, which quotes extensively from Geoffroy's manuscript notes on his experiments. 100. Etienne Geoffroy Saint-Hilaire, "Des differens etats de pesanteur des oeufs, au commencement et a la fin de 1'incubation," Journal complementaire des sciences medicales 1 (1820): 271278. 101. Quoted by Fischer, "Le concept experimental," p. 350. 102. Etienne Geoffroy Saint-Hilaire, "Sur les organes sexuels et sur les produits de generation des poules dont on a suspendu la ponte, en fermant l'oviductus," Memoires du Museum d'Histoire Naturelle 9 (1822): 1-24, quotation on p. 24. Geoffroy discussed his experiments through 1822 in Philosophie anatomique, 2: 509-513. 103. Etienne Geoffroy Saint-Hilaire, "Sur les deviations organiques provoquees et observees dans un etablissement d'incubations artificielles," Memoires du Museum d'Histoire Naturelle 13 (1825): 289-296, also published in Journal complementaire des sciences medicales 24 (1826): 256263; idem, "Monstre," p. 149; Fischer, "Le concept experimentale," pp. 359-360. At another incubation oven at Bourg-la-Reine, where the air was especially dry, Geoffroy noted that many of the chicks had digits curved backward. 104. Toby A. Appel, "Dominique-Auguste Lereboullet," DSB, 8: 253-255. 105. See, for example, Georges Cuvier, "Analyse des travaux de l'Academic Royale des Sciences, pendant l'annee 1826. Partie physique," Memoires de I'Academie des Sciences 9 (1830): xcviiclxvii, esp. pp. cliii-clix. 106. Bourdier assumes in his numerous articles on French science that Geoffroy was an evolutionist from the outset of his career. See, for example, Franck Bourdier, "Etienne Geoffroy SaintHilaire," DSB, 5: 355-358; idem, "Trois siecles d'hypotheses sur l'origine et la transformation des etres vivants (1550-1859)," Revue d'histoire des sciences 13 (1960): 1-44; and especially idem, "Geoffroy Saint-Hilaire versus Cuvier: The Campaign for Paleontological Evolution (18251838)," in Cecil J. Schneer, ed., Toward a History of Geology (Cambridge, Mass.: M.I.T. Press, 1969), pp. 36-61. This latter article contains an excellent account of Geoffroy's memoirs on evolution. In contrast, Laurent holds that Geoffroy's transformism represents a marked change from his earlier views on unity of composition, and that his shift to evolution was due to the influence of embryology (theory of arrests of development), teratology, and finally, paleontology. Goulven Laurent, "Le cheminement d'Etienne Geoffroy Saint-Hilaire (1772-1844) vers un transformisme scientiflque," Revue d'histoire des sciences 30 (1977): 43-70. See also Pietro Corsi, Oltre il mito: Lamarck e le scienze naturali del suo tempo (Bologna: il Mulino, 1983), pp. 315-364. 107. Bourdier, "Geoffroy Saint-Hilaire versus Cuvier," p. 46. Geoffroy published a major study of the bones of the skull of crocodiles in 1824 in which he provided a table comparing his and Cuvier's terminology and showing their anatomical disagreements. See Geoffroy, "Composition de la tete osseuse" (note 4), esp. p. 298. 108. Etienne Geoffroy Saint-Hilaire, "Recherches sur 1'organisation des Gavials; sur leurs affinites naturelles, desquelles resulte la necessite d'une autre distribution generique, Gavials, Teleosaurus et Steneosaurus; et sur cette question, si les Gavials (Gavialis), aujourd'hui repandus dans les parties orientales de l'Asie, descendent, par voie non interrompue de generation, des Gavials

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antediluviens, soit des Gavials fossiles, dits Crocodiles de Caen (Teleosaums), soit des Gavials fossiles du Havre et de Honfleur (Steneosaurus)," Memoires du Museum d'Histoire Naturelle 12 (1825): 97-155, esp. p. 144; idem, "Memoire ou l'on se propose de rechercher dans quels rapports de structure organique et de parente sont entre eux les animaux des ages historiques, et vivant actuellement, et les especes antediluviennes et perdues," Memoires du Museum d'Histoire Naturelle 17 (1828): 209-229, esp. p. 215. 109. Geoffroy, "Recherches sur 1'organisation des Gavials," p. 149. 110. Ibid., p. 154. 111. Ibid., pp. 150-152; Geoffroy, "Memoire ou 1'on propose de rechercher...," pp. 221-227. 112. Geoffroy, "Recherches sur 1'organisation des Gavials," quotations on pp. 150-151, 151n; Jean-Baptiste Lamarck, Philosophie zoologique (Paris, 1809), chapter 8, esp. p. 235. 113. Etienne Geoffroy Saint-Hilaire and E.R.A. Serres, "Rapport fait a 1'Academie des Sciences sur un memoire de M. Roulin ayant pour titre: Sur quelques changements observes dans les animaux domestiques transportes de 1'ancien monde dans le nouveau continent," Memoires du Museum d'Histoire Naturelle 17 (1828): 201-208, quotations on pp. 207-208. The report also appeared in Annales des sciences naturelles 16 (1829): 34-44. 114. Geoffroy, "Memoire ou l'on propose de rechercher. . .," p. 210. 115. Ibid., p. 210. 116. Ibid., pp. 216-219, quotation on p. 211. 117. On the comparison of Lamarck's and Geoffroy's evolutionary theories, see Franck Bourdier, "Lamarck et Geoffroy Saint-Hilaire face au probleme de l'evolution biologique," Revue d'histoire des sciences 25 (1972): 311-325; Corsi, Oltre il mito, pp. 315-364; and Reijer Hooykaas, Natural Law and Divine Miracle: The Principle of Uniformity in Geology, Biology and Theology (Leiden: E.J. Brill, 1963), pp. 80-89, 117-118. Hooykaas emphasizes the fragmentary nature of Geoffroy's ideas on evolution. 118. Etienne Geoffroy Saint-Hilaire, "Divers memoires sur de grands sauriens trouves a l'etat fossile vers les confins maritimes de la basse Normandie, attribues d'abord au crocodile, puis determines sous les noms de Teleosaums et Steneosaurus," Memoires de I'Academic des Sciences 12 (1833): 3-138, esp. p. 25. These memoirs were originally published as Recherches sur de grands sauriens. . . (Paris, 1831). 119. Etienne Geoffroy Saint-Hilaire, "Memoire sur le degre d'influence du monde ambiant pour modifier les formes animals; queslion interessant 1'origine des especes teleosauriennes et successivement celle des animaux de l'epoque actuelle," Memoires de 1'Academie des Sciences 12 (1833): 63-92, esp. p. 79. 120. Ibid., pp. 80-81. 121. Georges Cuvier, "Nature," in Dictionnaire des sciences naturelles, vol. 34 (Paris, 1825), pp. 261-268; Geoffroy, Fragment sur la Nature, Geoffroy, Cows de I'histoire naturelle des mammiferes (note 1); Cuvier and Valenciennes, Histoire naturelle des poissons (note 1). The first two volumes were published in 1828, and six more volumes appeared by the time of Cuvier's death. Valenciennes then continued the publication alone. 122. Cuvier, "Nature," p. 263.1 am grateful to William Coleman for making available to me his unpublished English translation of Cuvier's article. 123. Ibid., p. 265. 124. Ibid., p. 265. 125. Ibid., p. 266. 126. Ibid., p. 267. 127. Ibid., p. 267. 128. Cuvier and Valenciennes, Histoire naturelle des poissons, p. 1. 129. Georges Cuvier, "Memoire sur un ver parasite d'un nouveau genre (Hectocotylus octopodis)," Annales des sciences naturelles 18 (1829): 147-156, quotation on p. 149. 130. Etienne Geoffroy Saint-Hilaire, "De la 'theorie des analogues,' source de conceptions synthetiques d'un haut enseignement en histoire naturelle," Academie des Sciences, Comptes rendus 4 (1837): 537-546, quotation on p. 541; F. J. Cole, A History of Comparative Anatomy from Aristotle to the Eighteenth Century [1949](New York: Dover, 1975), pp. 28-33. On the theoryladen nature of supposed scientific facts, see Stephen Shapin, "History of Science and Its Sociological Reconstructions," History of Science 20 (1982): 157-211, esp. pp. 159-162.

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131. Geoffroy, Extrait d'un rapport fait a l'Academie, p. 5. 132. Ibid., p. 5. 133. Cuvier and Valenciennes, Histoire naturelle des poissons, pp. 275-277. 134. Ibid., p. 543. 135. Ibid., pp. 313-315. 136. Ibid., pp. 347, 348, 550. 137. Geoffroy, Cours de l'histoire naturelle des mammiferes, "Discours preliminaire," p. 33n; Geoffroy, Fragment sur la Nature, pp. 15-16, quotation on p. 16. 138. Geoffroy, Fragment sur la Nature, pp. 3, 4, 25, quotation on p. 4; Geoffroy, Cours de l'histoire naturelle des mammiferes, "Discours preliminaire," p. 32n. 139. Geoffroy, Fragment sur la Nature, p. 3. 140. Geoffroy, Cours de l'histoire naturelle des mammiferes, "Discours preliminaire," pp. 42, 43. 141. Ibid., pp. 41-42, quotation on p. 33n. 142. Ibid., pp. 7, 8. 143. Geoffroy, Fragment sur la Nature, p. 22. Chapter 6

1. Christoph Heinrich Pfaff, "Notice biographique sur Georges Cuvier," in idem, Lettres de Georges Cuvier a C.M. Pfaff, 1788-1792, sur l'histoire naturelle, la politique et la litterature, trans. Louis Marchant (Paris, 1858), pp. 11-44, 293-296, esp. pp. 37-44. 2. The political background to this chapter has been derived from Guillaume de Bertier de Sauvigny, The Bourbon Restoration, trans. Lynn M. Case (Philadelphia: University of Pennsylvania Press, 1966); and David H. Pinkney, The French Revolution of 1830 (Princeton: Princeton University Press, 1972). 3. Johann Wolfgang von Goethe, "Derniers pages de Goethe expliquant a l'Allemagne les sujets de philosophie naturelle controversees au sein de l'Academie des Sciences de Paris," Revue encyclopedique 53 (1832): 563-573 and 54 (1832): 54-68, esp. p. 567. 4. E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology (London: John Murray, 1916), reprint ed. (Chicago: University of Chicago Press, 1982), pp. 64-78; Jean Piveteau, "Les discussions entre Cuvier et Geoffroy Saint-Hilaire sur l'unite de composition du regne animal," Revue d'histoire des sciences 3 (1950): 343-363; J.H.F. Kohlbrugge, Historischekritische Studien uber Goethe als Naturforscher (Wurzburg: Curt Kabitzsch, 1913), pp. 61-113; and Wilhelm Lubosch, "Der Akademiestreit zwischen Geoffroy St.-Hilaire und Cuvier im Jahre 1830 und seine leitenden Gedanken," Biologisches Zentralblatt 38 (1918): 357-384, 397-456. For the interpretations of these and other historians, see the introduction to this book. The German authors have been helpful for their thorough research of the chronology of the controversy from 1830 to 1832. 5. See note 66. 6. Proces-verbaux, 9: 400. There is no manuscript of the memoir, "Quelques considerations sur l'organisation des mollusques," in the Archives of the Academie des Sciences. Laurencet appears to have been a physician of Lyon who was living in Paris in the 1820s and writing on the anatomy of the brain. 7. Etienne Geoffroy Saint-Hilaire, Principes de philosophie zoologique, discutes en mars 1830, au sein de 1'Academie Royale des Sciences (Paris, 1830), pp. 35-49. Another text of the report can be found in Proces-verbaux, 9: 403-406. According to Geoffroy, he and Latreille had dissected and observed together with Meyranx and Laurencet, attempting to verify the conclusions of the memoir. 8. For the text of the retracted portion, see Geoffroy, Principes de philosophie zoologique, pp. 50-52. The quotation was taken from Georges Cuvier, "Memoire sur les cephalopodes et sur leur anatomie," in idem, Memoires pour servir a l'histoire naturelle et a l'anatomie des mollusques (Paris, 1817), pp. 1-54, quotation on p. 43. 9. Revue bibliographique pour servir de complement aux Annales des sciences naturelles (1830): 19. This publication is a hard-to-find supplement to Annales des sciences naturelles which may be

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bound with the volumes of the journal. I used a copy at the Library of Congress. See also Gazette medicale de Paris 1 (1830): 72. 10. Meyranx to Cuvier, n.d., Fonds Cuvier, Ms. 3060, Bibliotheque de l'Institut de France. Meyranx was probably referring to Cuvier's work on mollusks (note 9). There is an additional undated letter from Meyranx to Cuvier in Ms. 3060 in which Meyranx complained that others were exaggerating and misrepresenting his conclusions. He stressed the factual nature of the work that he would soon present for Cuvier's judgment. According to Geoffroy's eulogy at Meyranx's funeral, Meyranx, moved by "the benevolent timidity of his character," at first abandoned his work because he feared to wound "the masters of science who, on these points of research, had other ideas than he." But when Meyranx realized that Geoffroy and Ampere supported him, he took courage. When he died in June 1832 of yellow fever, he left Geoffroy some of his manuscripts, one of which Geoffroy presented to the Academie in October 1832. Ampere reported in the name of a commission consisting of himself, Geoffroy, and Dumeril on the memoir which sought links between the cephalopods and reptiles. See Proces-verbaux, 10: 133, 359-360; Etienne Geoffroy Saint-Hilaire, "Discours de M. Geoffroy Saint-Hilaire prononce de 1er juillet 1832 aux funerailles de M. Meyranx," Revue encydopedique 59 (1833): 523-524. 11. Latreille to Cuvier, 18 May 1830, Fonds Cuvier, Ms. 3060, Bibliotheque de 1'Institut de France. 12. Pol Nicard, "Etude sur la vie et les travaux de M. de Blainville," in Henri Marie Ducrotay de Blainville, Osteographie, ou, Description iconographique comparee du squelette et du systeme dentaire des mammiferes recents et fossiles pour servir de base a la zoologie et a la geologie, 4 vols. (Paris, 1839-1864), 1: i-ccxxiii, esp. pp. lxxxiv-lxxxix; Revue encydopedique 45 (1830): 477, 761; Proces-verbaux, 9: 401, 402, 411-412, 413. Latreille reaffirmed his loyalty to Cuvier when he read on 31 May 1830 in his and Cuvier's name a report to the Academie on a memoir of Milne Edwards in which the latter had attempted to demonstrate a "uniformity of composition" between the mouthparts of chewing and sucking crustacea. Latreille praised the memoir for its factual content, emphasized that Milne Edwards was not a follower of Geoffroy because he did allow for the creation of new organs, and questioned Milne Edwards's use of the word "transformation" applied to structural elements as lacking in rigor. He concluded that Milne Edwards had failed to demonstrate his homologies because he could not point to intermediary forms. Revue encydopedique 46 (1830): 504-505. 13. Proces-verbaux, 9: 408. Cuvier's memoir, "Considerations sur les mollusques et en particulier sur les cephalopodes," was widely publicized. The original manuscript is located in Fonds Cuvier, Ms. 3063, Bibliotheque de 1'Institut de France. Cuvier's original drawings are in ibid., Ms. 3066, and the large colored drawings by Laurillard are in ibid., Ms. 3068. The memoir and drawings were published in Annales des sciences naturelles 19 (1830): 241-259. Shorter versions of the paper appeared in Revue encydopedique 46 (1830): 5-19 (followed by a critique written by Geoffroy for the occasion on pp. 20-22); Bulletin des sciences naturelles 20 (1830): 304-311; Gazette medicale de Paris 1 (1830): 81-82; Journal des debats, 23 February 1830; and Le Globe, 24 February 1830 (see note 52). Geoffroy reprinted the memoir from Journal des debats in Principes de philosophie zoologique, pp. 53-72. 14. Geoffroy, Principes de philosophie zoologique, quotations on pp. 63, 64-65, 66. 15. Geoffroy in his commentary claimed that mollusks had no brain corresponding to that of vertebrates and that they should be placed lower than the insects and Crustacea with respect to their nervous system rather than higher, as Cuvier placed them (ibid., pp. 68n-69n). 16. Ibid., pp. 73-80. See also Le Globe, 24 February 1830; Revue bibliographique (1830): 19-20. 17. Etienne Geoffroy Saint-Hilaire, "De la theorie des analogues pour etablir sa nouveaute comme doctrine, et son utilite pratique comme instrument," in idem, Principes de philosophie zoologique, pp. 81-108, quotation on p. 81; Proces-verbaux, 9: 413; Gazette medicale de Paris 1 (1830): 89-90; Journal des debats, 1 March 1830; Le Globe, 2 March 1830; Revue bibliographique (1830): 20-23. 18. Geoffroy, Principes de philosophie zoologique, pp. 87, 91. On the response of the audience at the Academie to Geoffrey's defense, see ibid., p. 86n. 19. Ibid., pp. 93-97. 20. Ibid., pp. 100-103.

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21. Ibid., p. 107. 22. Proces-verbaux, 9: 414; Gazette medicale de Paris 1 (1830): 110. Geoffrey's letter of excuse is preserved in the dossier of the meeting of 8 March 1830, Archives de l'Academie des Sciences, Institut de France. Otherwise the dossiers of the meetings from February to April 1830 contain no relevant information beyond what is published in Proces-verbaux. 23. Proces-verbaux, 9: 423. The manuscript of Cuvier's memoir, "Considerations sur l'os hyoid dans les animaux a poumons," is located in Fonds Cuvier, Ms. 3064, Bibliotheque de l'Institut de France. Extracts were published in Journal des debats, 23 March 1830; Le Globe, 23 March 1830; and Revue bibliographique (1830): 33-39. Geoffroy reprinted the extract from Journal des debats in Principes de philosophie zoologique, pp. 139-162. The titles vary. 24. Proces-verbaux, 9: 423; Etienne Geoffroy Saint-Hilaire, "De la theorie des analogues, appliquee a la connaissance de l'organisation des poissons," in idem, Principes de philosophie zoologique, pp. 109-138. See also Gazette medicale de Paris 1 (1830): 117-119; Journal des debats, 30 March 1830; Le Globe, 31 March 1830; and Revue bibliographique (1830): 25-29. 25. Geoffroy, Principes de philosophie zoologique, p. 142. Geoffroy replied in his commentary (p. 142n) that he had never used the expression "universal analogies" and that the words had no meaning. 26. Ibid., pp. 148-149. 27. Ibid., pp. 149-155n, quotation on p. 155n. 28. Ibid., p. 157. 29. Ibid., pp. 159-160. 30. Ibid., pp. 114-115. See also ibid., pp. 70-72n; and Gazette medicale de Paris 1 (1830): 118. 31. Geoffroy, Principes de philosophie zoologique, pp. 157-158. 32. Ibid., p. 138. 33. Proces-verbaux, 9: 426; Etienne Geoffroy Saint-Hilaire, "Sur les os hyoides," in idem, Principes de philosophie zoologique, pp. 163-190. See also Gazette medicale de Paris 1 (1830): 125126; Journal des debats, 30 March 1830; Le Globe, 31 March 1830; and Revue bibliographique (1830): 39-42. 34. Geoffroy, Principes de philosophie zoologique, pp. 165-166. 35. Ibid., pp. 174-183. 36. Ibid., p. 180. 37. Ibid., pp. 184-190, quotations on pp. 184-185, 187. Other examples of what Geoffroy later called "necessary facts" were Buffon's "law" of geographical distribution (that species of the Old and New Worlds differed), Lavoisier's theory of fermentation, and Montaigne's belief that monsters did not derive directly from God. See ibid., pp. 184, 185-186n. 38. Proces-verbaux, 9: 427; Georges Cuvier, "Suite des considerations sur l'hyoide," Fonds Cuvier, Ms. 3064, Bibliotheque de 1'Institut de France. This memoir was published in extract form in Journal des debats, 6 April 1830; Le Globe, 6 April 1830 (under the title "Considerations sur le sternum"); and Revue bibliographique (1830): 43-47. See also Gazette medicale de Paris 1 (1830): 135-136. Geoffroy did not include Cuvier's third memoir in Principes de philosophie zoologique. 39. Journal des debats, 6 April 1830. 40. Ibid. 41. Geoffroy, Principes de philosophie zoologique, pp. 72, 110, 147. 42. Ibid., p. 163. See also ibid., p. 20. 43. Ibid., p. 24. 44. Proces-verbaux, 9: 427; Geoffroy, Principes de philosophie zoologique, pp. 20-21, 29-34; Revue encyclopedique 46 (1820): 23; Gazette medicale de Paris 1 (1830): 136. 45. Georges Cuvier, "Analyse des travaux de I'Academie Royale des Sciences, pendant l'annee 1830. Partie physique," Memoires de I'Academie des Sciences 16 (1838): i-cxvii, esp. pp. lxi-lxiii, quotation on p. lxii. 46. Geoffroy, Principes de philosophie zoologique, p. 28; Gaspard-Joseph Martin Saint-Ange, Review of Geoffroy, Principes de philosophie zoologique, Revue encyclopedique 46 (1830): 707709; J.B.G.M. Bory de Saint-Vincent, Review of Geoffroy, Principes de philosophie zoologique, Le Moniteur universelle, 28 July 1830. 47. See, for example, Gazette medicale de Paris 1 (1830): 418; and Isidore Geoffroy Saint-Hilaire,

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Vie, travaux et doctrine scientifique d'Etienne Geoffroy Saint-Hilaire (Paris, 1847), reprint ed. (Brussels: Culture et Civilisation, 1968), p. 382. On Goethe's articles, see notes 3 and 58. 48. See note 13. 49. Gazette medicale de Paris 1 (1830): 81. The journal frequently published extensive accounts of Geoffroy's reports to the Academie, reports which often served as vehicles for propagating Geoffroy's own heretical views. One report (ibid., p. 7), for example, argued the case for spontaneous generation. 50. Jules Guerin, Review of Geoffroy, Principes de philosophie zoologique, Gazette medicale de Paris 1 (1830): 417-418, quotations on p. 417. 51. Gazette medicale de Paris 1 (1830): 283 (issue of 7 August 1830). 52. Le Globe. Journal philosophique et litteraire, reprinted, vol. 8 [15 February 1830-31 July 1830] (Geneva: Slatkine Reprints, 1974). 53. On Revue bibliographique, see note 9. On the editorial policy of Annales des sciences naturelles, see Annales des sciences naturelles 1 (1824): v-xvi. 54. Journal des debats politiques et litteraires, 30 March 1830. See also ibid., 23 February 1830, 2 March 1830. 55. Geoffroy, Principes de philosophie zoologique, pp. 191-204, 205-222. 56. See note 46. Another review, by Francois-Gabriel Boisseau, appeared in Journal universel des sciences medicates 58 (1830): 327-335. 57. Geoffroy, Principes de philosophie zoologique, pp. 205-206. 58. Johann Wolfgang von Goethe, "Reflexions de Goethe sur les debats scientifiques de mars 1830 dans le sein de 1'Academie des Sciences, publiees a Berlin dans les Annales de critique scientifique," Annales des sciences naturelles 22 (1831): 179-188; idem, "Derniers pages de Goethe" (note 3). The second article also appeared under the title "Les naturalistes francais, ou meditations de Goethe sur la marche et le caractere philosophique des sciences naturelles a Paris," Paris, ou Le livre des cent-et-un, vol. 5 (Paris, 1832), pp. 243-265. For a detailed study of Goethe's part in the debate in relation to his scientific and philosophical thought, see Dorothea Kuhn, Empirische und ideelle Wirklichkeit: Studien uber Goethes Kritik des franzosischen Akademiestreites (Graz: Hermann Bohlaus, 1967). 59. J.P. Eckermann, Conversations of Goethe with Eckermann and Soret, trans. John Oxenford (London, 1874), p. 479. 60. On Goethe and transcendental anatomy, see George A. Wells, "Johann Wolfgang von Goethe," DSB, 5: 442-446; idem, "Goethe and the Intermaxillary Bone," British Journal for the History of Science 3 (1967): 348-361; Russell, Form and Function, pp. 45-51; and Michel Guedes, "Goethe et Geoffroy Saint-Hilaire," Histoire et nature 1 (1973): 27-45. Goethe's morphological writings were republished in Johann Wolfgang von Goethe, Zur Morphologie, 2 vols. (Stuttgart, 1817-1823). 61. Johann Wolfgang von Goethe, "The Metamorphosis of Plants," in idem, Botanical Writings, trans. Bertha Mueller (Honolulu: University of Hawaii Press, 1952), pp. 30-78. 62. Etienne Geoffroy Saint-Hilaire, "Sur des ecrits de Goethe lui donnant les droits au titre de savant naturaliste," Annales des sciences naturelles 22 (1831): 188-193. 63. Eckermann, Conversations of Goethe, p. 480. 64. Goethe, "Derniers pages de Goethe," quotation on p. 567. 65. Goethe, "Reflexions de Goethe," p. 179. 66. Eckermann, Conversations of Goethe, p. 480. 67. On this episode, see Goethe, "Derniers pages de Goethe," pp. 65-67, quotation on p. 66; Le Globe, 15 July 1830, 21 July 1830; Journal des debats, 14 July 1830, 20 July 1830; Gazette medicale de Paris 1 (1830): 271; and Revue encydopedique 46 (1830): 240-241. There is no account of the argument in Proces-verbaux. 68. Eckermann, Conversations of Goethe, p. 480. 69. Antoine Duges, Memoire sur la conformite organique dans l'echelle animale (Montpellier, 1832); idem, "Memoire sur la conformite organique dans l'echelle animale, lu a 1'Academie des Sciences, seance du 18 octobre 1831," Annales des sciences naturelles 24 (1831): 254-260, also published in Gazette medicale de Paris 2 (1831): 370-372. The manuscript of Duges's memoir, on which Cuvier based his report, is located in Fonds Cuvier, Ms. 3062, Bibliotheque de l'Institut de

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277

France. On Duges's career, see Etienne Frederic Bouisson, "Eloge de Duges, prononce a la seance de rentree des facultes," Journal de la Societe de Medecine-Pratique de Montpellier 2 (1840): 81107; and Hubert Rodrigues, "Antoine-Louis Duges," Nouvelle biographie generate, 15: columns 87-89. On Duges's biological thought, see Edmond Perrier, La philosophie zoologique avant Darwin, 3rd ed. (Paris, 1896), pp. 149-159; and Russell, Form and Function, pp. 86-88. 70. Duges, "Memoire sur la conformite organique," p. 255. 71. Ibid., p. 256. On Moquin-Tandon and the concept of "zoonite," see Alfred Moquin-Tandon, Monographie de la famille des Himdinees, 2nd ed. (Paris, 1846), pp. 195-208; and idem, Un naturaliste a Paris sous Louis-Philippe: journal de voyage inedit (1834), 2nd ed., ed. Marcel Roland (Paris: Mercure de France, 1944), "Introduction," pp. 8-47. 72. The discussion in this and the preceding paragraph is based upon the manuscript version of Duges's memoir. 73. Proces-verbaux, 9: 706; Etienne Geoffroy Saint-Hilaire, "Lettre sur quelques points du memoire ayant pour titre: De la conformite organique dans l'echelle animale," Gazette medicale de Paris 2 (1831): 381-382. Geoffroy's letter, dated 31 October 1831, is followed by Duges's response (p. 382). Geoffroy's undated and unsigned draft of a letter to the Academie des Sciences resigning from the commission is located in Dossier Etienne Geoffroy Saint-Hilaire, Archives de 1'Academie des Sciences, Institut de France. 74. Duges, Memoire sur la conformite organique, p. v. 75. Georges Cuvier, "Rapport sur le memoire de M. Duges concernant la conformite organique," 36pp., Fonds Cuvier, Ms. 3062, Bibliotheque de l'Institut de France, p. 32. 76. Ibid., pp. 33, 34-35. 77. Ibid., pp. 33-35. 78. Duges, Memoire sur la conformite organique, pp. v-vi. 79. Proces-verbaux, 10: 155, 258. 80. Ibid., p. 653; Moquin-Tandon, Un naturaliste a Paris, p. 75; Antoine Duges, Traite de physiologie comparee de l'homme et des animaux, 3 vols. (Paris: 1838-1839). See especially the "Tableau des homologues" in vol. 1. 81. I. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 389-391. Geoffroy apparently considered becoming a deputy for Etampes after the July Revolution (p. 200n). 82. The Reverend Richard Owen, The Life of Richard Owen, 2 vols. (London, 1894), 1: 48-49. 83. Etienne Geoffroy Saint-Hilaire, "Divers memoires sur de grands sauriens trouves a l'etat fossile vers les confins maritimes de la basse Normandie, attribues d'abord au crocodile, puis determines sous les noms de Teleosaurus et Steneosaurus," Memoires de l'Academie des Sciences 12 (1833): 3-138, esp. pp. 3-26, 43-61. Perhaps Geoffroy was able to have this long and controversial series of memoirs published in the Academie's journal because he was President of the Academie in 1833. The memoirs were originally published separately as idem, Recherches sur de grands sauriens. . . (Paris, 1831). 84. Gazette medicale de Paris 1 (1830): 375, 384, quotations on p. 384. 85. Ibid., p. 399; Etienne Geoffroy Saint-Hilaire, "Sur quelques considerations generales des rochers et la specialite de cet organe chez le crocodile," ibid., 391-393, esp. p. 391. 86. Geoffroy, ibid., p. 392. 87. Gazette medicale de Paris 2 (1831): 18; Etienne Geoffroy Saint-Hilaire, "Sur la theorie physiologique designee sous le nom de vitalisme," ibid., pp. 8-12; idem, "Divers memoires sur de grands sauriens," pp. 63-92. Geoffroy's article attacking vitalism was countered by the physician J.-J. Virey, "Des vraies fondemens de la theorie du vitalisme," ibid., pp. 29-31. Virey claimed that one could recognize under the veil of matter, active and intelligent forces which bring about vital movement. Any other physiology was claimed to be materialistic. Geoffroy continued this little polemic with "Reponse a un article de M. J. -J. Virey sur le doctrine de vitalisme," ibid., pp. 6263. Geoffroy had previously attacked vitalism in print in 1926 in a review of Hippolyte Cloquet, Traite complet de l'anatomie de I'homme, Revue encyclopedique 19 (1826): 188-191. 88. Georges Cuvier, "Memoire sur le progres de l'ossification dans le sternum des oiseaux," Annales des sciences naturelles 25 (1832): 260-272, also published in Gazette medicale de Paris 3 (1832): 8-11, and in Revue encyclopedique 53 (1832): 213-215; Etienne Geoffroy Saint-Hilaire, "Memoire sur les observations communiquees par M. le baron Cuvier a 1'Academie des Sciences

278

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(seance du 2 Janvier 1832) au sujet des sternums des oiseaux, et sur leur immediate application a la theorie des analogues," Annales des sciences naturelles 27 (1832): 189-200. 89. Georges Cuvier, "Extrait d'un memoire sur les oeufs de la seiche," Annales des sciences naturelles 26 (1832): 69-75, also published in Nouvelles annales du Museum d'Histoire Naturelle 1 (1832): 153-160. 90. Georges Cuvier, "Sur la variete de composition des animaux," n.d., Fonds Cuvier, Ms. 3065, Bibliotheque de l'Institut de France. Cuvier apparently intended to include his memoirs from the debate in this work. He also began a piece entitled "Considerations sur les osselets de l'oreille des animaux a poumon et sur les opercules des ouies des poissons osseux," Fonds Cuvier, Ms. 3064, Bibliotheque de l'Institut de France. Cuvier announced his intention to publish "Sur la variete de composition" in "Analyse des travaux de 1'Academie . . . 1830," pp. lxii-lxiii. 91. See, for example, Richard W. Burkhardt, Jr., The Spirit of System: Lamarck and Evolutionary Biology (Cambridge, Mass.: Harvard University Press, 1977), pp. 196-198. 92. Georges Cuvier, "Eloge de M. de Lamarck," Memoires de 1'Academie des Sciences 13 (1835): i-xxxi, quotations on pp. i-ii. Originally the eloge was to be read on 26 July 1831, the same day that Arago read an eloge of Volta. 93. Ibid., p. xix. 94. According to Blainville, the Academie committee concerned with the reading of the eloge insisted that Cuvier make changes which Cuvier refused. The eloge was not read until 26 November 1832 by Baron Sylvestre and then in an edited form. See Henri Marie Ducrotay de Blainville, Histoire des sciences de l'organisation et de lews progres comme base de la philosophie, ed. F.L.M. Maupied, 3 vols. (Paris, 1858), 3: 342. Lamarck's oldest son, who, with his brothers and sisters, was invited to attend the public meeting at which the eloge was read, protested to the Academie in two letters and requested that the eloge not be printed. Refusing to take responsibility for the work of a member, the Academie went ahead with the publication. According to Robert Courrier, Georges Cuvier (1769-1832): certains aspects de sa carriere (Paris: Palais de l'Institut, 1970), pp. 15-17, the published eloge is the original rather than a truncated version. 95. Georges Cuvier, "Avertissement," Nouvelles Annales du Museum d'Histoire Naturelle 1 (1832): i-iv, quotation on p. ii. This introduction was written on 15 March 1832, two months before Cuvier's death. 96. Ibid., p. iii. 97. Etienne Geoffroy Saint-Hilaire, "Observations sur la concordance des parties de l'hyoide dans les quatre classes des animaux vertebres," Nouvelles Annales du Museum d'Histoire Naturelle 1 (1832): 321-356. One of the most senior professors at the Museum, Geoffroy was not denied the right of publication in its journals until 1834 when Blainville objected to one of Geoffroy's tendentious reports to the Academie. See below, chapter 7. 98. Georges Cuvier, Histoire des sciences naturelles, depuis leur origins jusqua nos jours, chez tons les peuples connus, professee au College de France, par Georges Cuvier, completee, redigee, annotee et publiee par M. Magdeleine de Saint-Agy, 5 vols. (Paris, 1841-1845), reprint ed. (Brussels: Culture et Civilisation, 1969), 2: 537-538. 99. Quoted by John Vienot, Le Napoleon de l'intelligence: Georges Cuvier, 1769-1832 (Paris: Fischbacher, 1832), p. 210. A shorter variation of this quotation appears in Etienne Pasquier, Eloge de M. le baron Georges Cuvier (Paris, 1832), 55pp., p. 52; Georges Cuvier, "Derniere lecon de M. Cuvier, professee au College de France le 8 mai 1832," Le Temps, 5 June 1832. I am grateful to Jean Chandler Smith for locating this lecture. 100. Louis Agassiz, Address Delivered on the Centennial Anniversary of the Birth of Alexander von Humboldt (Boston: Boston Society of Natural History, 1869), pp. 42-43; Francois Arago, "Ampere," in idem, Oeuvres de Francois Arago, vol. 2 (Paris, 1854), pp. 1-116, esp. pp. 70-75. On Humboldt's appreciation of Geoffroy, see Use Jahn, "Etienne Geoffroy Saint-Hilaire an Alexander von Humboldt uber Goethes Stellungnahme zum Pariser Akademiestreit," NTM: Zeitschrift fur Geschichte der Naturwissenschaft, Technik und Medizin 10 (1973): 59-67; and Jean Theodorides, "Une lettre inedite de A. de Humboldt concernant Etienne Geoffroy Saint-Hilaire," Histoire et nature 1 (1973): 67-69. 101. E.R.A. Serres, "Recherches sur l'anatomie comparee des animaux invertebres. Premier memoire. Que sont par rapport aux vertebres a l'homme les animaux invertebres?" Annales des sciences naturelles, series 2, 2 (1834): 238-248, quotations on pp. 239, 247.

NOTES

279

102. Henri Dutrochet, "Considerations sur l'opposition qui existe entre les animaux vertebres et les animaux invertebres, presentees a l'Academie des Sciences de Paris, le 10 mars 1834," L'lnstitut 2 (1834): 90-92. Dutrochet envisioned the vertebrates on the one hand and the articulates and mollusks on the other as two "opposite poles," or two branches united by a common trunk consisting of the radiates. See also Joseph Schiller and Tetty Schiller, Henri Dutrochet (Henri du Trochet, 1776-184 7): le materialisme mecaniste et la physiologie generate (Paris: Albert Blanchard, 1975), pp. 114, 116, 135, 142-143. 103. Cuvier, "Analyse des travaux de l'Academie . . . 1830," pp. lxi-lxiii.

Chapter 7

1. Honore de Balzac, Louis Lambert [1832], trans. Clara Bell, in idem, Comedie Humaine, ed. George Saintsbury, 40 vols. (London, 1895-1900), 39: 158-280, quotation on p. 233. 2. Honore de Balzac, A Distinguished Provincial at Paris [ Un grand homme de province a Paris] [1839], trans. Ellen Marriage, in idem, Comedie Humaine, vol. 12, quotation on p. 78. 3. Honore de Balzac, Pere Goriot (New York: Washington Square Press, c. 1962, 1966), p. v. Any French or English edition after 1842 will include this dedication. 4. The distinction between those who are in a position to judge the arguments based on personal experience ("the core-set") and those who are not has been sharpened for me by Harry M. Collins, "The Place of the 'Core-Set' in Modern Science: Social Contingency with Methodological Propriety in Science," History of Science 19 (1981): 6-19. 5. Among those who attended Geoffroy's salon were Jean Reynaud, Victor Considerant, Pierre Leroux, Stephane Esquiros, David Richard, and Charles Didier. Other more celebrated intellectuals—Edgar Quinet, Alfred de Musset, David d'Angers, and Victor Hugo—also on occasion took part in the festivities, which were presided over by Geoffroy, his wife, his daughter Stephanie, his son Isidore, and his daughter-in-law. On Geoffroy's relationship to these and several other advanced thinkers, a number of whom participated in the Revolution of 1848 and subsequently left the country, see the excellent article by Franck Bourdier, "Le prophete Geoffroy Saint-Hilaire, George Sand et les saint-simoniens," Histoire et nature 1 (1973): 47-66. On the salon, see also Les belles femmes de Paris et de la province, 2 vols. (Paris, 1839-1840), 2: 49-54.I consulted a copy of this obscure work, brought to my attention by Bourdier, in the Bibliotheque Nationale. On the medical student Maxime Vernois (1809-1877) and the cleric Julien Danielo (1806-1866), see Etienne Geoffrey Saint-Hilaire, Notions synthetiques, historiques et physiologiques de philosophie naturelle (Paris, 1838), pp. 61-62n, 123-129. 6. Etienne Geoffroy Saint-Hilaire, Etudes progressives d'un naturaliste pendant les annees 1834 et 1835, faisant suite a ses publications dans les 42 volumes des Memoires et Annales du Museum d'Histoire Naturelle (Paris, 1835), p. xiii. Speaking at Cuvier's funeral, Geoffroy recalled the "inexpressible happiness" of having been first to recognize Cuvier's genius and reveal it to the scientific world. "How much at this moment of final parting ... my thoughts are brought back to that common life of our youthful years, to those relations so intimate and devoted, to that community of work so sweet to us both." "Principaux discours prononces sur la tombe de Cuvier, le 16 mai 1832," Annales des sciences naturelles 26 (1832): 394-415, quotations on pp. 403, 407-408. 7. Proces-verbaux, 10: 65, 81-82, 84, 324, 331, 334, 346. 8. Revue encyclopedique 53 (1832): 213, 213n. 9. Etienne Geoffroy Saint-Hilaire, Fragments biographiques reproduisant les discours prononces aux funerailles de MM. Thouin, de Lacepede, Pinel, de Lamarck, et Cuvier, 32pp. (Paris, 1832); idem, Fragments biographiques, precedes d'Etudes sur la vie, les ouvrages et les doctrines de Buffon (Paris, 1838). See above, notes 5 and 6. 10. Etienne Geoffroy Saint-Hilaire, "Note sur la repulsion, consideree comme caracteristique de l'essence des choses," Academie des Sciences, Comptes rendus (hereafter Comptes rendus) 1 (1838): 551-554. 11. Etienne Geoffroy Saint-Hilaire, "Puissance du monde ambiant governant l'engendrement

280

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des causes auxquelles se rapportent les formes differentes et a la multiplication des especes vegetales," Dossier Etienne Geoffroy Saint-Hilaire, Archives de l'Academie des Sciences, Institut de France. An extract appeared in L'Institut 2 (1834): 314-315. 12. Alfred Moquin-Tandon, Un naturaliste a Paris sous Louis-Philippe: journal de voyage inedit (1834), 2nd ed., ed. Marcel Roland (Paris: Mercure de France, 1944), pp. 105, 116. 13. Ibid., p. 116. Isidore Geoffroy, in his biography of his father, preferred to ignore Geoffroy's work of the 1830s. As far as he was concerned, his father's work was complete as of 1828. After 1828, his father's writings served only to defend his views, and therefore utilized a new procedure—that of taking up the most diverse subjects at short intervals. Isidore Geoffroy Saint-Hilaire, Vie, travaux et doctrine scientifique d'Etienne Geoffroy Saint-Hilaire (Paris, 1847), reprint ed. (Brussels: Culture et Civilisation, 1968), pp. 366-367. 14. Moquin-Tandon, Un naturaliste a Paris, p. 106, quotation on p. 116. In 1836 the naturalistvoyager J.-R.-C. Quoy described Geoffroy to a friend: "M. Geoffroy, poor man! Loving commotion so that others will talk about him, who was completely beaten in every way, both in form and in substance, with his extravagant idealism, his insufferable elocution and his style of a cook, to use the mocking expression of his eloquent adversary [Cuvier]. Now, on certain matters, he is considered nearly half crazy." E.T. Hamy, "Notes intimes sur Georges Cuvier redigees en 1836 par le Dr Quoy pour son ami J. Desjardins, de Maurice," Archives de medecine navale 86 (1906): 450-475, quotation on p. 472. 15. Jourdan's memoir, read 30 June 1834, was entitled "Description d'un appareil dentaire particulier appartenant a la colonne vertebrate." Jourdan suggested that in the grass snake of Africa there were peculiar vertebral processes that were analogous to teeth. Geoffroy and Serres were charged with the report which Geoffroy presented on 7 July 1834. Proces-verbaux, 10: 543-547, esp. p. 547. The report was also published in L'Institut 2 (1834): 222-223, and in Geoffroy, Etudes progressives, pp. 67-80. 16. Geoffroy, Etudes progressives, p. 78n. On Blainville's skepticism with regard to teratology, see Henri Marie Ducrotay de Blainville, Cuvier et Geoffroy Saint-Hilaire: biographies scientifiques (Paris, 1890). 17. Proces-verbaux, 10: 646. 18. Geoffroy, Etudes progressives, pp. vii-viii. 19. Geoffroy supplied each epoch with a leader, but pointedly left the leader of the final age blank. Ibid., p. ix. 20. On the circumstances of writing this work, see Geoffroy, Notions synthetiques, pp. 19-20n. 21. Geoffroy, Etudes progressives, p. xii. 22. Ibid., pp. 84-85. 23. Geoffroy, Notions synthetiques, pp. 46, 56, 61, quotation on p. 56. The epigraph from Balzac, "La science est une, et vous l'avez partagee," appears on the frontispiece of Notions synthetiques and several times in the text. 24. Etienne Geoffroy Saint-Hilaire, "Il n'est qu'une seule physique dans l'universe ...," Comptes rendus 9 (1839): 194-200, 228-233, 268-274, 290-295, 305-315, quotation on p. 228. 25. Etienne Geoffroy Saint-Hilaire, "Monstre," in Dictionnaire classique d'histoire naturelle, vol. 11 (Paris, 1827), pp. 108-151, esp. pp. 139-141. 26. Proces-verbaux, 10: 621; Etienne Geoffroy Saint-Hilaire, "Loi universelle (attraction de soi pour soi), ou clef applicable a l'interpretation de tous les phenomenes de philosophie naturelle," in Geoffroy, Etudes progressives, pp. 125-189, esp. p. 125. 27. On Napoleon, see Geoffroy, Notions synthetiques, pp. i-xlix. 28. Geoffroy, Etudes progressives, p. 144. 29. For the most extended accounts of soi pour soi, see Geoffroy, Etudes progressives, pp. 125189; idem, Notions synthetiques; idem, "De la loi d'attraction de soi pour soi; et nouveaux efforts de l'inventeur pour en presenter le principe comme une annexe etendant les vues de la gravitation universelle de Newton," Comptes rendus 6 (1838): 766-769; idem, "Il n'est qu'une seule physique dans 1'universe ..."; and idem, "D'un nouvel argument de physique intrastellaire ...," Comptes rendus 9 (1839): 439-446, 489-491. 30. On the reaction of savants to soi pour soi, see Geoffroy, Notions synthetiques, pp. 3-10. 31. Geoffroy, "Il n'est qu'une seule physique," p. 295.

NOTES

281

32.I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 303. For example, Journal des savants, which ordinarily reviewed scientific works, offered no reviews of Principes de philosophie zoologique, Etudes progressives, or Notions synthetiques. 33. Geoffroy, Notions synthetiques, pp. 122-123. 34. Geoffroy, Etudes progressives, pp. 188-189. 35. Ibid., p. 178. 36. Geoffroy, Notions synthetiques, p. 82. Geoffroy also combated allegations of irreligion in idem, "D'un nouvel argument de physique intrastellaire," pp. 489-491. 37. Geoffroy, Notions synthetiques, pp. 26, 33. 38. Etienne Geoffroy Saint-Hilaire, "De la necessite d'embrasser dans une pensee unitaire, les plus subtiles manifestations de la psychologie, et des difficultes de la solution de ce probleme," Comptes rendus 4 (1837): 259-263. 39. Geoffroy, Notions synthetiques, p. 110. 40. Etienne Geoffroy Saint-Hilaire, "Heresies pantheistiques," in Dictionnaire de la conversation et de la lecture, vol. 31 (Paris, 1836), pp. 484-489, quotation on p. 487, reprinted in idem, Fragments biographiques (1838), pp. 337-351; I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 406. 41. Revue zoologique 1 (1838): 206-207. On Geoffroy's beliefs concerning marsupials and monotremes, see Kathleen G. Dugan, "Marsupials and Monotremes in Pre-Darwinian Theory," Ph.D. Dissertation, University of Kansas, 1980, pp. 51-54, 90-91, 211-225. 42. Etienne Geoffroy Saint-Hilaire, "Geologie et palaeontographie. Decouverte d'ossements fossiles dans les bassins de l'Auvergne, et considerations sur les deux sortes de zoologie, Tune revelee par des vestiges antediluviens, et l'autre formee par la serie animale actuellement vivante," in idem, Etudes progressives, pp. 87-103, quotation on p. 97. 43. Geoffroy, Etudes progressives, p. xi (note). 44. Geoffroy, "Geologie and palaeontographie," p. 100. 45. Ibid., p. 110. See also Etienne Geoffroy Saint-Hilaire, "De l'influence des circonstances exterieures sur les etres organises," L'Institut 1 (1833): 12-13, p. 12n. As president of the Academie in 1833, Geoffroy was able to read this paper at the annual meeting of the five Academies of the Institut on 2 May 1833. 46. Geoffroy, Fragments biographiques (1838), p. 333. On the progress of humanity, see also idem, "Heresies pantheistiques." 47. On the orangutan of Sumatra, see Etienne Geoffroy Saint-Hilaire, "Considerations sur les singes les plus voisins de l'homme," Comptes rendus 2 (1836): 92-95; idem, "Memoire sur l'orangoutang," Comptes rendus 2 (1836): 581-585, 601-603, and 3 (1836): 1-8, 27-31. 48. In at least one case Geoffroy was drawn into a public dispute over science and religion. In 1837 during a discussion of paleontological evolution at the Academie, Geoffroy intimated that Cuvier may have made deliberate concessions to the religious authorities. Cuvier's recognition that fossil species differed from living species might have been offensive to the Church, if Cuvier, out of political expediency, had not mitigated the impact of the new ideas by being "careful to limit himself in his communications to approximations of biblical and historical truths." Frederic Cuvier strongly reproached Geoffroy before the Academie for insulting his brother's memory. Geoffroy claimed to be deeply hurt by the rebuke. See Etienne Geoffroy Saint-Hilaire, "Des changements a la surface de la terre, qui paraissent dependre originairement et necessairement de la variation preexistante, incessante, lente et successive des milieux ambiants divers et consecutifs du globe terrestre," Comptes rendus 5 (1837): 183-194, quotation on p. 193; "Observations de M. Frederic Cuvier, sur un paragraphe du memoire de M. Geoffroy Saint-Hilaire, insere dans le compte rendu de la seance de 14 aout 1837," Comptes rendus 5 (1837): 305-306; and I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 404. 49. For a critique of Comptes rendus, see [Guglielmo B.I.T. Libri] "Lettres a un Americain sur l'etat des sciences en France. I. L'Institut," Revue des deux mondes, 4th series, 21 (1840): 789-818. Some of Geoffroy's correspondence with Arago, editor of the journal, is in the Butler Library, Columbia University. Arago, though an admirer of Geoffroy, was impatient with his excessive demands for publication of his articles. 50. Geoffroy, "Il n'est qu'une seule physique dans l'universe" (note 24); idem, "D'un nouvel argument de physique intrastellaire" (note 29). Geoffroy was not the only one to be presenting

282

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such inappropriate papers to the Academie. Moquin-Tandon described in his journal of 1834 a "vieux original' with a bald head who presented to the Academie a machine made of copper and began reading a memoir titled, "New System of the World." The author supposed that there were only three "stars" in nature—the earth, the moon, and the sun—and supported this notion by reference to the Trinity. At first the academicians laughed, and then at the request of Poisson, the author was told to cease reading. Moquin-Tandon, Un naturaliste a Paris, pp. 79-80. 51. Etienne Geoffroy Saint-Hilaire, "De la nature et de lage des ossements fossiles, sous des temps antediluviens, et d'abord d'un essai de polemique ayant commence dans la derniere seance," Comptes rendus 5 (1837): 365-372, quotation on p. 366. 52. On this episode, see Geoffroy, Fragments biographiques (1838), "Vieilesse outragee," pp. 137-157; Geoffroy, Notions synthetiques, pp. 111-120; and [Jean Reynaud] "Fondation de la menagerie du Museum d'Histoire Naturelle," Magasin pittoresque 6 (1838): 106-108. 53. [Etienne Geoffroy Saint-Hilaire] "Necrologie. M. de Lamarck," Gazette medicate de Paris 1 (1830): 13-16, quotation on p. 16. 54. Ibid., pp. 15, 16. 55. Geoffroy, Fragments biographiques (1838), pp. 81-82. Through the 1830s Geoffroy continued to expand upon the doctrine of "necessary facts" (see chapters 5 and 6), which he associated with Lamarck and Buffon. See, for example, Geoffroy, Etudes progressives, p. 175n; and idem, Notions synthetiques, pp. 27, 29. 56. On Buffon's statue, see Etienne Geoffroy Saint-Hilaire, "De la statue de Buffon, afin de lui faire recouvrir ses anciens honneurs," Comptes rendus 1 (1838): 681-687; idem, "Sur Buffon et sa statue," lp., n.d., Dossier Etienne Geoffroy Saint-Hilaire, Archives de l'Academie des Sciences, Institut de France; and Geoffroy to Abel-Francois Villemain, 5 September 1837, Special Collections Branch, Smithsonian Institution Libraries. 57. Geoffroy, Fragments biographiques (1838), "Dedicace." 58. See note 56. For citations to these publications, see I. Geoffroy, Etienne Geoffroy SaintHilaire, pp. 468-469. 59. Geoffroy, Fragments biographiques (1838), pp. 4, 79; Etienne Geoffroy Saint-Hilaire, "De la Theorie des Analogues, source de conceptions synthetiques d'un haul enseignement en histoire naturelle," Comptes rendus 4 (1837): 537-546. 60. Geoffroy, "De la theorie des analogues," p. 544. 61. Geoffroy, "Memoire sur l'orang-outang," p. 582. 62. Geoffroy, Notions synthetiques, p. 63. 63. Ibid., p. 28. 64. "Obseques de M. Geoffroy Saint-Hilaire," Gazette medicale de Paris 12 (1844): 409-422, esp. p. 416. 65. Ibid., p. 409; I. Geoffroy, Etienne Geoffroy Saint-Hilaire, p. 416. 66. On Geoffroy and Sand, see Bourdier, "Le prophete Geoffroy Saint-Hilaire," pp. 56-63. 67. George Sand, Correspondance, ed. Georges Lubin (Paris: Gamier freres, 1964— ), 2: 892n, 924 (editor's note), 3: 54, 638-639. 68. Sand to Eliza Tourangin, 30 December 1836, Correspondance, 3: 622. 69. Sand to Geoffroy, 30 April 1837, Correspondance, 3: 831-835 and notes, quotations on pp. 832, 833; George Sand, Lelia, ed. Pierre Reboul (Paris: Gamier freres, 1960), p. 552 (editor's note). 70. George Sand, "Soi pour soi. La science," in idem, Lelia, pp. 545-553, quotation on p. 546. 71. Ibid., p. 548. 72. Ibid., pp. 547, 548-549. 73. Ibid., p. 547. 74. Honore de Balzac, Correspondance, ed., Roger Pierrot, 5 vols. (Paris: Gamier freres, 19601969), 2: 691-693, 766-770, 772-775. On Geoffroy and Balzac, see Helene d'Also, "Balzac, Cuvier et Geoffroy Saint-Hilaire," Revue d'histoire de la philosophie, n.s. 2 (1934): 339-354; Silvestre de Sacy, "Balzac, Geoffroy Saint-Hilaire et l'unite de composition," Mercure de France, No. 303 (1948): 292-305, 469-480; Theophile Cahn, La vie et l'oeuvre d'Etienne Geoffroy Saint-Hilaire (Paris: Presses Universitaires de France, 1962), pp. 243-248. 75. Honore de Balzac, The Magic Skin [La peau de chagrin] [1831], trans. Katherine Frescott Wormeley, in La Comedie Humaine de Honore de Balzac, 30 vols. (New York, 1896), 1: 1-323, quotation on pp. 25-26.

NOTES

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76. Balzac to Laure Surville, August? 1832, Correspondence, 2: 89. 77. Balzac, Louis Lambert, p. 233. 78. Ibid., pp. 235, 236. 79. Ibid., p. 229. 80. Balzac to Geoffroy, circa 1 December 1835, Correspondance, 1: 766; Geoffroy to Balzac, 4 December 1835, ibid., 2: 767-768; Geoffroy to Balzac, 8 December 1835, ibid., 2: 772-775. See also above, note 23. 81. Honore de Balzac, La comedie humaine, vol. 1 (Paris: Editions du Seuil, 1965), "Avantpropos," pp. 51-56, quotation on p. 51. 82. Like Quinet, Michelet turned to natural history not as an end in itself, but for its moral and social ramifications. In the 1840s, he attended the lectures of Isidore Geoffroy, Milne Edwards, Serres, and Felix Pouchet and recalled in his journal of 1865 the impact of philosophical anatomy on his thought: "[Serres's] book on embryogenesis lifted the veil of Isis for me, and made me to see the enormous moral bearing of what we believed to be physical." In the notes of one of his popular, romantic works on nature, The Insect, Michelet recalled "the great duel between Cuvier and Geoffroy," as well as Goethe's famous conversation with Soret. His wife had prepared a discussion of the debate for the book, but he did not in the end include it because he was not sure that the audience of a work of popular science would appreciate the technicalities. See Robert van der Elst, Michelet naturaliste (Paris, 1914), quotation on pp. 45-46; and Jules Michelet, L'Insecte, 2nd ed. (Paris, 1858), pp. 379-380. 83. "Obseques de M. Geoffroy Saint-Hilaire," "Discours de M. Edgard Quinet," pp. 421-422, quotation on p. 421. 84. Ibid., p. 422. 85. Honore de Balzac, "Guide-ane a l'usage des animaux qui veulent parvenir aux honneurs," in P.J. Stahl (pseud, for Pierre-Jules Hetzel) ed., Scenes de la vie privee et publique des animaux, 2nd ed. (Paris, 1867) [1st ed. 1842], pp. 267-268. The story has recently been republished in paperback in J.J. Grandville, Un renard pris au piege. Scenes de la vie privee et publique des animaux, vol. 2 (Editions Gallimard, 1978), pp. 52-75. 86. Ibid., p. 271. 87. Ibid., p. 271. 88. Ibid., p. 272. 89. Ibid., pp. 273-274. 90. Ibid., p. 280. 91. Ibid., p. 285. 92. On Raspail, see Dora B. Weiner, Raspail: Scientist and Reformer (New York: Columbia University Press, 1968); and Marc Klein, "Francois-Vincent Raspail," DSB, 11: 300-302. 93. Weiner, Raspail, offers a penetrating analysis of Raspail's character and underlines Raspail's own role in assuring his lack of recognition by the academicians. 94. Weiner, Raspail, p. 70. 95. Quoted in ibid., p. 73. 96. In the 1830s and 1840s, two cliques held sway at the Academie—the Arago and Brongniart factions. They divided to some extent along political lines (Arago was considerably more liberal in politics than Brongniart), but not particularly according to lines laid out by the Cuvier-Geoffroy debate. Some of Cuvier's disciples (e.g., Valenciennes) were supported by Arago, while others (e.g., Duvernoy) were supported by Brongniart. To critics of the prevailing scientific system, Brongniart was especially despised because he used his considerable influence to promote the careers of his relatives, Adolphe Brongniart, Dumas, and Audouin, as well as their mutual friend, Milne Edwards. 97. Four volumes appeared in 1829 and 1830. 98. Francois-Vincent Raspail, "Coteries scientifiques," Annales des sciences d'observation 3 (1830): 151-159, quotation on p. 154. 99. Ibid., p. 156. 100. Francois-Vincent Raspail, Nouveaux coups defouet scientifique, 33pp. (Paris, 1831), "Theorie des analogues; et discussion entre MM. Cuvier et Geoffroy," pp. 1-24, esp. pp. 8-9. I used a copy of this obscure work at the Bibliotheque Nationale, Paris. 101. Ibid., p. 24.

284

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102. Ibid., pp. 23-24. 103. Raspail's article on scientific cliques was followed by a comparison of the fates of Lamarck and the chemist Nicolas Louis Vauquelin, both recently deceased. Vauquelin died rich and honored because he knew how to play politics, while Lamarck, who cared only for science, died poor, blind, and paralyzed, with none but his daughters to cry. Francois-Vincent Raspail, "Necrologie; parallele," Annales des sciences d'observation 3 (1830): 159-160. 104. Geoffroy to Raspail, 5 July 1833, No. 130, and Raspail to Geoffroy, 9 July 1833, No. 131, Fonds Raspail, Bibliotheque Centrale du Museum National d'Histoire Naturelle. Francois-Vincent Raspail, Nouveau systeme de chimie organique (Paris, 1833), p. 10. 105. Weiner, Raspail, p. 127. 106. Geoffroy to Raspail [7 July 1833], No. 126, Fonds Raspail. 107. Raspail to Geoffroy, 9 July 1833, No. 131, Fonds Raspail. 108. Isidore de Gosse (pseud, for Bertrand-Isidore de Salles), Histoire naturelle drolatique et philosophique des professeurs du Jardin des Plantes, des aide-naturalistes, preparateurs, etc., attaches a cet etablissement, accompagnee d'episodes scientifiques et pittoresques (Paris, 1847). This work was published "with annotations by" Frederic Gerard, who wrote the introduction and signed many of the articles. On Gerard, see Franck Bourdier, "Geoffroy Saint-Hilaire versus Cuvier: The Campaign for Paleontological Evolution (1825-1838)," in Cecil J. Schneer, ed., Toward a History of Geology (Cambridge, Mass.: M.I.T. Press, 1969), pp. 36-61, esp. pp. 59-60. 109. Alexandre Brongniart was described as follows: "Arrogant, imperious, skillful to take advantage of every talent and every opportunity to arrive at his ends, the old porcelain maker of the manufactury of Sevres is the chief of the great coterie which dominates the aristocratico-scientific republic of the Museum. It is he who can at his whim make or unmake a candidate. Everywhere he has disseminated his sons, his relatives, and his vassals, and except for some professors such as MM. Isidore Geoffroy, de Blainville, Cordier, and Gay-Lussac, everyone bows to his despotic sway." See ibid., pp. 63-64, quotation on p. 63. On Adolphe Brongniart and Milne Edwards, see ibid., pp. 89-91, 144-145. In one "episode" entitled "Le Banquet" (ibid., pp. 93-98), the "Aragotists" and the "Brongniardists," said to be a repetition in miniature of the Guelfs and the Ghibellines, were reconciled on condition that Adolphe be made vice-president of the Academie. 110. Ibid., p. 33. Deianira sent to her husband Hercules a splendid robe stained with the blood of the centaur Nessus as a charm against his preferring another woman to her. The garment brought Hercules to his death. 111. The bookcover of Histoire naturelle drolatique advertised other works by the same publisher, Gustave Sandre, including Frederic Gerard, Geoffroy Saint-Hilaire et son epoque: histoire d'ecole de philosophie naturelle. I used a copy of Histoire naturelle drolatique with its original cover in the Bibliotheque Nationale, Paris. 112. De Gosse, Histoire naturelle drolatique, p. 31. 113. Ibid., pp. 29, 39, 134, 137-138. 114. Ibid., p. 166. 115. Ibid., "Funerailles de Geoffroy Saint-Hilaire ler," pp. 168-181, quotations on pp. 170, 181. 116. Ibid., pp. 19-20. 117. Ibid., p. 21. 118. Ibid., pp. 21-22. 119. Ibid., p. 23. 120. Ibid., pp. 231-240. Chapter 8

1. Pierre Flourens, "Eloge historique d'Etienne Geoffroy Saint-Hilaire, lu dans la seance publique annuelle du 22 mars 1852," in idem, Recueil des eloges historiques, lus dans les seances publiques de I'Academie des Sciences, 3 vols. (Paris, 1856-1862), 1: 229-284, quotation on p. 257. 2. Isidore Geoffroy Saint-Hilaire, Vie, travaux et doctrine scientifique d'Etienne Geoffroy SaintHilaire (Paris, 1847), reprint ed. (Brussels: Culture et Civilisation, 1968), pp. 125-126, 378, quotation on p. 305. 3. Georges Louis Duvernoy, "Anatomie," in Dictionnaire universel d'histoire naturelle, ed.

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Charles Dessalines D'Orbigny, 13 vols. (Paris, 1842-1861), 1: 438-454, esp. section on "Anatomie philosophique, transcendante et speculative," pp. 445-449. Duvernoy's student at Strasbourg, Dominique-Auguste Lereboullet, took a similar view in his article "Anatomie philosophique," in Dictionnaire encyclopedique des sciences medicates, ed. Amedee Dechambre, 100 vols. (Paris, 1864-1889), 1st series, 4: 250-278. 4. This point is made by E. S. Russell, Form and Function: A Contribution to the History of Animal Morphology (London: John Murray, 1916), reprint ed. (Chicago: University of Chicago Press, 1982). 5. Duvernoy, "Anatomie," pp. 445-449. 6. Dov Ospovat, The Development of Darwin's Theory: Natural History, Natural Theology, and Natural Selection, 1838-1859 (Cambridge: Cambridge University Press, 1981), pp. 6-38. See also idem, "Perfect Adaptation and Teleological Explanation: Approaches to the Problem of the History of Life in the Mid-nineteenth Century," in Studies in History of Biology, vol. 2, ed. William Coleman and Camille Limoges (Baltimore: Johns Hopkins University Press, 1978), pp. 33-56. 7. On von Baer and the embryological criterion for the determination of homologies, see Russell, Form and Function, pp. 113-132; and Timothy Lenoir, The Strategy of Life: Teleology and Mechanics in Nineteenth-Century German Biology (Dordrecht: D. Reidel, 1982), pp. 72-95. Lenoir sees developmental morphology as an improved phase of the program of vital materialism of the school of Blumenbach. He recognizes Cuvier's contribution to this program, but not Geoffroy's (ibid., pp. 54-111). 8. On von Baer, recapitulation, and the branching conception of development, see Dov Ospovat, "The Influence of Karl Ernst von Baer's Embryology, 1828-1859: A Reappraisal in Light of Richard Owen's and William B. Carpenter's 'Paleontological Application of "von Baer's Law'"," Journal of the History of Biology 9 (1976): 1-28, esp. pp. 10-12; idem, Darwin's Theory, pp. 117-124; and Stephen Jay Gould, Ontogeny and Phylogeny (Cambridge, Mass.: Harvard University Press, 1977), pp. 52-63. German embryology, while it lent support to the four embranchements, also confirmed some of the more striking results put forth by Geoffroy and his followers. Rathke, for example, in his 1829 memoir on the embryology of the crayfish, confirmed Savigny's belief that limbs and jaws of arthropods belonged to a single type of structure. Russell, Form and Function, p. 137. 9. Richard Owen, Lectures on the Comparative Anatomy and Physiology of the Vertebrate Animals, Delivered at the Royal College of Surgeons of England, in 1844 and 1846. Part 1. Fishes (London, 1846), pp. 137-141. 10. Russell, Form and Function, pp. 134-141. See also Lenoir, Strategy of Life, pp. 95-102; and Vern L. Bullough, "Martin Heinrich Rathke," DSB, 11: 307-308. According to Lenoir, Rathke deliberately sought to refute by embryological argument Geoffroy's contention that the gill system was related to the ribs or larynx of higher vertebrates. Instead, Rathke showed that the gill system was a more highly developed form of the hyoid of higher vertebrates. 11. Russell, Form and Function, pp. 144-146; Vladislav Kruta, "Karl Bogislaus Reichert," DSB, 11:360-361. 12. See, for example, the term "homologous" in A Dictionary of Biology, 6th ed., ed. M. Abercrombie, CJ. Hickman, and M.L. Johnson (Baltimore: Penguin Books, 1973), p. 139. 13. This is the main argument of Ospovat, Darwin's Theory. 14. On the career of Isidore Geoffroy Saint-Hilaire, see Armand de Quatrefages, Eloge historique de M. Isidore Geoffroy Saint-Hilaire, 22pp., extracted from Bulletin de la Societe Imperiale Zoologique d'Acclimitation 9 (1862): 257-278; J.-B. Dumas, Eloge historique de Isidore Geoffroy SaintHilaire (Paris, 1872), 36pp., also in Memoires de l'Academie des Sciences 38 (1873): 178-212; Nicholas Joly, Eloge historique d'lsidore Geoffroy Saint-Hilaire (Toulouse, 1863), 24pp., extracted from Memoires de I'Academie Imperiale des Sciences de Toulouse (1863); and Franck Bourdier, "Isidore Geoffroy Saint-Hilaire," DSB, 5: 358-360. 15. Apparently, Isidore Geoffroy was astonished to be elected on his first attempt. See the correspondence between F.-V. Raspail and Etienne Geoffroy Saint-Hilaire, Fonds Raspail, Bibliotheque Centrale du Museum National d'Histoire Naturelle. 16. Joly, Isidore Geoffroy, p. 21. Joly, one of Isidore Geoffroy's few disciples, became professor of zoology at the Faculte des Sciences of Toulouse. 17. Quatrefages, Isidore Geoffroy, p. 9; Joly, Isidore Geoffroy, p. 14.

286

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18. On homologies between embranchements, see I. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 217-224; on soi pour soi, ibid., p. 303; and on transformism, ibid., pp. 344-358; and Quatrefages, Isidore Geoffroy, pp. 9-10. 19. Isidore Geoffroy Saint-Hilaire, Essaies de zoologie generate, ou, Memoires et notices sur la zoologie generale, l'anthropologie, et l'histoire de la science (Paris, 1841), pp. viii-ix. 20.I. Geoffroy, Etienne Geoffroy Saint-Hilaire, pp. 265-267, quotation on p. 326; Isidore Geoffroy Saint-Hilaire, Histoire generate et particuliere des anomalies de I'organisation... ou, Traite de teratologie, 3 vols. (Paris, 1832-1836), 1: vii; idem, Essaies de zoologie, pp. 55-56. 21. Ibid., pp. 336, 340. 22. Pierre Flourens, "Eloge historique de G. Cuvier," in idem, Recueil des eloges, 1: 105-196. 23. Flourens, "Eloge . . . Geoffroy," quotation on p. 257. 24. Ibid., pp. 255, 256. 25. Ibid., pp. 246-247. 26. Ibid., p. 260. 27. Pierre Flourens, Buffon: histoire de ses travaux et de ses idees (Paris, 1844), chapter 4. 28. Flourens, "Eloge ... Geoffroy," pp. 260-261. 29. Pierre Flourens, "De Punite de composition et du debat entre Cuvier et Geoffroy SaintHilaire," Journal des savants (1864): 265-274, 380-388, 417-425, 587-595, 719-726, quotation on p. 721. This series of articles was also published separately in 1865. 30. The authors of Museum satire dubbed Blainville "Anatomicus erinaceus" (hedgehog anatomist) and described him: "Of a misanthropic character, he sets himself in opposition on all issues because he is the enemy of the entire universe. He spends his life alone with a parrot, a dog, and a housekeeper..." Isidore de Gosse (pseud, for Bertrand-Isidore de Salles), Histoire naturelle drolatique et philosophique des professeurs du Jardin des Plantes (Paris, 1847), p. 125. 31. This section is largely based on my paper, "Henri de Blainville and the Animal Series: A Nineteenth-Century Chain of Being," Journal of the History of Biology 13 (1980): 291-319. On the series doctrine and the doctrine of a unique creation, see also Goulven Laurent, "La paleontologie de de Blainville (1777-1850): support ou ruine du fixisme?," Histoire et nature 12-13 (1978): 8396; and Gabriel Gohau, "L'unite de la creation chez Blainville," Revue d'histoire des sciences 32 (1979): 43-58. The first issue of volume 32 of this journal is devoted to a collection of articles on Blainville. 32. Pierre Flourens, "Eloge de Marie-Henri Ducrotay de Blainville," Memoires de l'Academie des Sciences 27, 2nd part (1860): i-lx, esp. pp. xiii-xxi, also in idem, Recueil des eloges, 1: 285341; idem, Review of Blainville, Osteographie, Journal des savants (1850): 321-333, 415-429, 449459, (1851): 115-125, 206-217, 273-284. Flourens believed that Blainville had demonstrated "unity of the kingdom," but that the evidence was not yet available to prove "unity of creation" (p. 284). 33. For the final version of Blainville's animal series, see Henri Marie Ducrotay de Blainville, Considerations generates sur les animaux et leur classification (n.p., n.d.), 37pp., extracted from idem, "Animal," in Supplement to Dictionnaire des sciences naturelles (Paris and Strasbourg, 1840) (Only one volume, A-Aye, of the Supplement was ever published. The extracted version that I consulted was obtained from the Museum of Comparative Zoology, Harvard University.); idem, Sur les principes de zooclassie, ou de la classification des animaux (Paris, 1847); and idem, Osteographie, ou, Description iconographique comparee du squelette et du systeme dentaire des mammiferes recents et fossiles pour servir de base a la zoologie et a la geologie, 4 vols. (Paris, 18391864). 34. Henri Marie Ducrotay de Blainville, Cuvier et Geoffroy Saint-Hilaire; biographies scientifiques (Paris, 1890), pp. 199-200. 35. Blainville, Considerations generates sur les animaux, pp. 8-9, quotation on p. 8. On Blainville and philosophical anatomy, see Appel, "Blainville and the Animal Series," pp. 302-307. 36. Henri Marie Ducrotay de Blainville, Histoire des sciences de l'organisation et de leursprogres comme base de la philosophie, ed. F.L.M. Maupied, 3 vols. (Paris, 1858). This work first appeared in 1845. On Cuvier, see ibid., 3: 370-411, on Lamarck, ibid., 3: 411-466, and on Oken, ibid., 3: 466-514, 527-528. For examples of criticism, see Pol Nicard, "Etude sur la vie et les travaux de M. de Blainville," in Blainville, Osteographie, 1: i-ccxxiii, esp. p. cxvii; Flourens, "Eloge .. . Blain-

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ville," p. xxiv; and Blainville, Cuvier et Geoffroy, pp. 2-3. On Blainville's history and his attempt to support Catholicism by biology, see Georges Canguilhem, "L"Histoire des sciences de 1'organisation' de Blainville et l1'abbe Maupied," Revue d'histoire des sciences 32 (1979): 73-91; and Evelyne Shuster-Aziza, "Note sur H. de Blainville historien de la biologie," Revue d'histoire des sciences 25 (1972): 191-200. 37. Blainville, Cuvier et Geoffroy, p. 4. 38. Blainville, Histoire des sciences de l'organisation, 1: vi, 3: 336-338. Despite Blainville's disapproval of Cuvier's use of political power to dominate science, Blainville had little sympathy with those who wanted to appeal over the heads of the academicians to the public. In his opinion, even Cuvier had overpopularized science. He held that Goethe should never have involved himself in the Cuvier-Geoffroy debate because he had insufficient knowledge of science, and accused the editors of Le Temps and Le National of writing their pro-Geoffroy analyses "in the manner of clever men who speak on a subject that they truly do not know and for a public that cannot be a competent judge ..." Blainville distrusted the public and the public in turn disliked him. Blainville, Cuvier et Geoffroy, p. 377. 39. Flourens, "Eloge ... Blainville," pp. i, vii, ix, xxlii. 40. For Blainville's comments on Geoffroy, see, for example, Blainville, Cuvier et Geoffroy, pp. 293, 312-313, 432-433, 435-436, quotation on p. 435. 41. Ibid., pp. 312-313. 42. On Milne Edwards, see Marcelin Berthelot, "Notice historique sur Henri Milne Edwards," Annales des sciences naturelles (Zoologie), series 7, 13(1892): 1-30; Armand de Quatrefages, "Discours de M. A. de Quatrefages. Funerailles de Henri-Milne Edwards," ibid., series 6, 19 (1885): iix; and Jean Anthony, "Henri Milne-Edwards," DSB, 9: 407-409. 43. On Milne Edwards's role in the training of zoologists, see Berthelot, "Milne Edwards," pp. 10-12; and Henri de Lacaze-Duthiers, "Dix-sept annees d'enseignement de la zoologie en Sorbonne," Revue scientifique, 3rd series, 37 (1886): 737-748. 44. Jean Theodorides, "Les debuts de la biologie marine en France: Jean-Victor Audouin et Henri-Milne Edwards, 1826-1829," Actes du ler Congres International de l'Oceanographie (Monaco, 1966) (1968), 2: 417-437. 45. Jean-Victor Audouin and Henri Milne Edwards, "Recherches anatomiques et physiologiques sur la circulation dans les crustaces," Annales des sciences naturelles 11 (1827): 283-314, 352-399. Cuvier and Dumeril's report appeared in Annales des sciences naturelles 10 (1827): 394-399. 46. In his history of crustaceans, for example, he wrote that he would not discuss unity of composition applied to the entire animal kingdom, or pronounce judgment on Geoffroy's homologies between Crustacea and vertebrates. Henri Milne Edwards, Histoire naturelle des crustaces, comprenant l'anatomie, la physiologie et la classification de ces animaux, 3 vols. (Paris, 1834-1840), 1: 50, 52. 47. Jean-Victor Audouin and Henri Milne Edwards, "Troisieme memoire sur l'anatomie et la physiologie des crustaces; Recherches anatomiques sur le systeme nerveux," Annales des sciences naturelles 14 (1828): 77-102, quotation on p. 78. 48. Ibid., pp. 98-99. 49. Etienne Geoffroy Saint-Hilaire, "Rapport sur le travail de MM. Audouin et Henri Milne Edwards, ayant pour litre: 'Recherches anatomiques sur le systeme nerveux chez les crustaces,'" Annales des sciences naturelles 13(1828): 218-224. The apparent difference, in Geoffroy's opinion, could be more simply explained as the result of the elongated conformation of the crayfish and the massive orbicular form of the crab. 50. Milne Edwards, Histoire naturelle des crustaces, 1: 5-6. Division of labor was first discussed in Henri Milne Edwards, "Organisation," in Dictionnaire classique d'histoire naturelle, vol. 12 (Paris, 1827), pp. 332-344, esp. pp. 339-341. For later versions, see idem, Lecons sur la physiologie et l'anatomie comparee de l'homme et des animaux faites a la Faculte des Sciences de Paris, vol. 1 (Paris, 1857), p. 17ff; and idem, Introduction a la zoologie generale, ou considerations sur les tendances de la nature dans la constitution du regne animal (Paris, 1853), chapters 3 and 4. Milne Edwards's biological philosophy is discussed by Russell, Form and Function, pp. 195-200, 204205; Camille Limoges, "L'economie naturelle et le principe de correlation chez Cuvier et Darwin," Revue d'histoire des sciences 23 (1970): 34-45; and Ospovat, Darwin's Theory, pp. 124-129.

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51. Milne Edwards, Histoire naturelle des crustaces, 1: 14-15, 41-42. 52. Henri Milne Edwards, "Observations sur les changemens de forme que divers Crustaces eprouvent dans le jeune age," Annales des sciences naturelles (Zoologie), 2nd series, 3 (1835): 321334, quotation on p. 327. See also Ospovat, Darwin's Theory, p. 126. 53. Milne Edwards, Histoire naturelle des crustaces, 1: 24-26, 42-45. 54. For other examples of the use of homologies in classification, see Mary P. Winsor, "Barnacle Larvae in the Nineteenth Century: A Case Study in Taxonomic Theory," Journal of the History of Medicine 24 (1969): 294-309; and idem, Starfish, Jellyfish, and the Order of Life: Issues in Nineteenth-Century Science (New Haven: Yale University Press, 1976). 55. Milne Edwards, Histoire naturelle des crustaces, 1: 227. 56. Ibid., 1: 229. 57. Henri Milne Edwards, "Considerations sur quelques principes relatifs a la classification naturelle des animaux, et plus particulierement sur la distribution methodique des mammiferes," Annales des sciences naturelles (Zoologie), 3rd series, 1 (1844): 65-99, esp. pp. 67, 70; Ospovat, "Von Baer's Embryology," pp. 11-12; Russell, Form and Function, p. 205. Milne Edwards recalled that he had been attracted to the theory of arrests of development earlier in his career. See Milne Edwards, Introduction a la zoologie generale, pp. 99-100. But, even then, his attitude toward the theory was ambivalent. See, for example, idem, "Observations sur les changemens de forme." Isidore Geoffroy, reporting on this paper to the Academie in 1833, associated Milne Edwards with Serres and philosophical anatomy, yet it seemed to him that Milne Edwards was thinking more in terms of excess of development than of arrest of development. Isidore Geoffroy Saint-Hilaire, "Rapport fait a l'Academie Royale des Sciences sur un memoire de M. Milne Edwards, intitule: Observations sur les changemens de forme que les crustaces eprouvent dans le jeune age," Annales des sciences naturelles 30 (1833): 360-372. 58. Milne Edwards, Lefons, 31-32. 59. For references to Geoffroy, see Milne Edwards, Lefons, pp. 28-29n; and idem, Introduction a la zoologie generale, pp. 95-96, 111-112, 135, 148. 60. Henri Milne Edwards, "Recherches zoologiques faites pendant un voyage sur les cotes de la Sicile," Annales des sciences naturelles (Zoologie), 3rd series, 3 (1845): 129-182, quotation on p. 130. 61. Milne Edwards, Lefons, p. 9; idem, Introduction a la zoologie generale, pp. i-vi. 62. Milne Edwards, Lecons, pp. 2-4. Milne Edwards conceived of a vital force not as a force that contravened physical and chemical forces, but rather as a distinct force superadded to physical and chemical forces. 63. Ibid., p. 26. See also idem, Introduction a la zoologie generale, pp. 81-85. 64. Milne Edwards, Lecons, p. 13. See also idem, Introduction a la zoologie generale, pp. 7-12. 65. Milne Edwards, Lemons, p. 2. 66. Ibid., p. 4. See also idem, Introduction a la zoologie generale, pp. 12-19. 67. Milne Edwards, Lecons, p. 13. See also idem, Introduction a la zoologie generale, p. 57ff. 68. Milne Edwards, Introduction a la zoologie generale, p. 62. 69. On philosophical anatomy in Britain, see Ospovat, Darwin's Theory, esp. pp. 6-38; idem, "Perfect Adaptation"; Philip F. Rehbock, The Philosophical Naturalists: Themes in Early Nineteenth-Century British Biology (Madison: University of Wisconsin Press, 1983), pp. 15-114; and Michael Ruse, The Darwinian Revolution: Science Red in Tooth and Claw (Chicago: University of Chicago Press, 1979), pp. 94-131. 70. William Whewell, History of the Inductive Sciences, 3 vols. (London, 1837), 3: 451-478. Whewell's struggle with homology versus teleology is discussed by Michael Ruse, "William Whewell and the Argument from Design," The Monist 60 (1977): 244-268; and idem, Darwinian Revolution, pp. 125-127. 71. Whewell, History of the Inductive Sciences, 3: 457. Whewell's discussion of the debate is confusing, for he tended to interpret the controversy in terms of four plans versus one. 72. Ibid., 3: 468. 73. William Whewell, History of the Inductive Sciences, 3rd ed., 3 vols. (London, 1857), reprint ed. (London: Frank Cass & Co., 1967), pp. 558-562, quotation on pp. 560-561. Whewell's change of position dated from the mid-1840s.

NOTES

289

74. Ruse, "Whewell and the Argument from Design," p. 245. 75. Martin Barry, "On the Unity of Structure in the Animal Kingdom," Edinburgh New Philosophical Journal 2 ( 1836-1837): 116-141, 345-364, esp. pp. 116-118, 136-137, 363. On Barry, see Ospovat, Darwin's Theory, pp. 11-22; and Rehbock, Philosophical Naturalists, pp. 59-61. 76. J. Estlin Carpenter, "William Benjamin Carpenter: A Memorial Sketch," in William B. Carpenter, Nature and Man: Essays Scientific and Philosophical (London, 1888), pp. 1-152, esp. p. 10. On Carpenter, see Ospovat, Darwin's Theory, pp. 12-14, 17, 20; and Rehbock, Philosophical Naturalists, pp. 61-68. 77. William B. Carpenter, Principles of General and Comparative Physiology, 2nd ed. (London, 1841) [1st ed., 1839], p. 190. For Carpenter's discussion of design in nature, see ibid., pp. 183-202, 559-563. 78. Ibid., p. 560. 79. Ibid., p. 192. 80. Ibid., p. 202. 81. On Owen's career, see the Rev. Richard Owen, The Life of Richard Owen, 2 vols. (London, 1894); and Wesley C. Williams, "Richard Owen," DSB, 10: 260-263. On Owen and philosophical anatomy, see Russell, Form and Function, pp. 102-112; Ospovat, Darwin's Theory, pp. 6-23,129139; Ruse, Darwinian Revolution, pp. 116-125; and Rehbock, Philosophical Naturalists, pp. 7384. 82. Owen, Life of Richard Owen, 1: 48-58. The debate was also a stimulating intellectual experience for the Swiss-born Louis Agassiz (1807-1873) who, like Owen, arrived in Paris in 1831 near the commencement of his professional career to meet Cuvier and to work with the collections at the Museum. See Toby A. Appel, "Jeffries Wyman, Philosophical Anatomy and the Scientific Reception of Darwin in America," Journal of the History of Biology (in press). 83. Richard Owen, On the Anatomy of Vertebrates, 3 vols. (London, 1866-1868), 3: 786-787. Owen was not unaware of the political dimension of the debate. He drew a parallel between this confrontation and the Pasteur-Pouchet debate: "Besides the superiority in fact and argument, Pasteur, like Cuvier, had the advantage of subserving the prepossessions of the 'party of order' and the needs of theology." Ibid., 3: 814, also quoted in John Farley and Gerald L. Geison, "Science, Politics and Spontaneous Generation in Nineteenth-Century France: The Pasteur-Pouchet Debate," Bulletin of the History of Medicine 48 (1974): 161-198, on p. 167. 84. Ibid., 3: 789. Before 1859 Owen had been sympathetic to the notion of the formation of new species by secondary law. He publicly announced a designed or theistic theory of evolution in 1863 which he called "derivation." The most probable nature of the law accounting for "derivation" was "an innate tendency to deviate from parental type." Ibid., 3: 799-809, esp. p. 807; Appel, "Wyman,"; Roy M. MacLeod, "Evolutionism and Richard Owen, 1830-1868: An Episode in Darwin's Century," Isis 56 (1965): 259-280. 85. Owen, Anatomy of Vertebrates, 3: 787-789. 86. Richard Owen, Lectures on the Comparative Anatomy and Physiology of the Invertebrate Animals, Delivered at the Royal College of Surgeons in 1843 (London, 1843); idem, Vertebrate Animals (note 9); idem, On the Archetype and Homologies of the Vertebrate Skeleton (London, 1848); idem, On the Nature of Limbs. A Discourse Delivered on Friday, February 9, at an Evening Meeting of the Royal Institution of Great Britain (London, 1849). 87. Owen, Invertebrate Animals, pp. 374, 379; idem, On the Archetype, p. 7. The British naturalist William Sharp Macleay (1792-1865), author of the quinarian system of classification, had previously distinguished between "affinities" (forms with anatomical similarities that were placed close together in his system) and "analogies" (forms with some anatomical similarities but that were otherwise very different). On Macleay, see Winsor, Starfish, Jellyfish, pp. 81-87; Ospovat, Darwin's Theory, pp. 101-113; and Rehbock, Philosophical Naturalists, pp. 26-30. 88. Owen, On the Archetype, pp. 7-8. 89. See Ospovat, "Perfect Adaptation," p. 36. 90. Owen, On the Nature of Limbs, pp. 9-10. 91. Owen, On the Archetype, p. 73. 92. Owen, Vertebrate Animals, pp. 36-61; idem, On the Archetype, pp. 72-73; and idem, On the Nature of Limbs, pp. 39-40, 84-86.

290

NOTES

93. Owen, Anatomy of Vertebrates. 94. Owen, Invertebrate Animals, pp. 367-371; idem, Vertebrate Animals, pp. 10-11; Ospovat, "Von Baer's Embryology," pp. 17-23. Before Ospovat's article, Owen's views were usually confounded with the Meckel-Serres theory of recapitulation. 95. Owen, Vertebrate Animals, pp. 42-46; idem, On the Nature of Limbs, pp. 83-84. 96. Owen, Vertebrate Animals, pp. 137-140; idem, On the Archetype, pp. 63-68. 97. Owen, On the Archetype, pp. 171-172; idem, On the Nature of Limbs, pp. 84-86. 98. Ibid., pp. 85-86. 99. Ospovat, "Perfect Adaptation," esp. pp. 33-35, 51-52. 100. Charles Darwin, Darwin's Notebooks on Transmutation of Species. Part I. First Notebook (July 1837-February 1838), ed. Gavin de Beer, Bulletin of the British Museum (Natural History). Historical Series, vol. 2, no. 2 (London, 1860), pp. 23-73, esp. pp. 53-55. 101. Edward Manier, The Young Darwin and His Cultural Circle: A Study of the Influences Which Helped Shape the Language and Logic of the First Drafts of the Theory of Natural Selection (Dordrecht: D. Reidel, 1978), pp. 52-55. 102. Ospovat, Darwin's Theory, pp. 145-146, 149. 103. Charles Darwin, On the Origin of Species: A Facsimile of the First Edition (Cambridge, Mass.: Harvard University Press, 1964), p. 206. 104. Ospovat, Darwin's Theory, pp. 115-189, esp. p. 146. 105. Darwin, Origin of Species, chapter 13, "Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Organs," pp. 411-458, quotations on pp. 434, 435. On Darwin and morphology, see also George Gaylord Simpson, "Anatomy and Morphology: Classification and Evolution: 1859 and 1959," Proceedings of the American Philosophical Society 103 (1959): 286306; Ruse, Darwinian Revolution, pp. 196-197. It is known that Darwin read Isidore Geoffroy's biography of his father, all the other major works of Isidore Geoffroy, and Robert Knox's Great Artists and Great Anatomists (London, 1852), which included a highly flattering account of Geoffroy. Peter J. Vorzimmer, "The Darwin Reading Notebooks (1838-1860)," Journal of the History of Biology 10 (1977): 107-153, esp. pp. 126, 133, 146, 148, 152. 106. Darwin, Origin of Species, p. 435. 107. Ruse, Darwinian Revolution, suggests (pp. 141, 196) that serial homologies were not as important to Darwin's theory as they were to Owen's. Darwin accepted a modified vertebral theory of the skull (that both vertebrae and segments of the skull were formed on a common plan), and also noted serial homologies between the anterior and posterior limbs in vertebrates, and between leaves, sepals, petals, stamens, and pistils in plants. 108. Jeffries Wyman, "On Symmetry and Homology in Limbs," Proceedings of the Boston Society of Natural History 11 (1868): 246-278. Wyman suggested that Owen's Archetype should be modified to reflect a fore and hind or mirror image symmetry as well as bilateral symmetry. See Appel, "Wyman." On the variety of non-Darwinian theories of evolution, see Peter Bowler, The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades Around 1900 (Baltimore: Johns Hopkins University Press, 1983). 109. Patrick Geddes, "Morphology," in Encyclopedia Brittanica, 9th ed., vol. 16 (1890), pp. 837846 [Geddes claimed that Geoffroy was "by far the most distinguished anatomist of the transcendental school" and "had enormously greater influence than Goethe or Oken" (p. 839)]; Russell, Form and Function, pp. 168, 230-241, 247. 110. See, for example, John Farley, "The Initial Reactions of French Biologists to Darwin's Origin of Species" Journal of the History of Biology 1 (1974): 275-300. On the reception of Darwin in France, see also Robert E. Stebbins, "French Reactions to Darwin, 1859-1882," Ph.D. Dissertation, University of Minnesota, 1965; idem, "France," in Thomas F. Glick, ed. The Comparative Reception of Darwinism (Austin: University of Texas Press, 1974), pp. 117-163; Yvette Conry, L'introduction du darwinisme en France au X I X e siecle (Paris: J. Vrin, 1974); Joy Harvey, "Evolutionism Transformed: Positivists and Materialists in the Societe d'Anthropologie de Paris from Second Empire to Third Republic," in David Oldroyd and Ian Langham, eds. The Wider Domain of Evolutionary Thought (Dordrecht: D. Reidel, 1983), pp. 289-310; and Pietro Corsi and Paul J. Weindling, "Darwinism in Germany, France and Italy," in David Kohn, ed., The Darwinian Heritage (Princeton: Princeton University Press, 1985), pp. 683-729, esp. pp. 698-711. Corsi empha-

NOTES

291

sizes the continuity between pre-1860 and post-1860 debates in France over evolution and suggests the need for more study of the French reaction to the evolutionary theories of Geoffroy and others in the period 1830-1860. 111. Charles Darwin, De l'origine des especes, ou des lois du progres chez les etres organises, trans. Clemence Auguste Royer (Paris, 1862), "Preface du traducteur," pp. v-lxiv. 112. Flourens, "De 1'unite de composition" (note 29), p. 274. 113. Ibid., p. 424. See also ibid., p. 587. 114. Armand de Quatrefages, Darwin et ses precurseurs francais: etude sur le transformisme, 2nd ed. (Paris, 1892), p. 69. On Quatrefages, see Camille Limoges, "Jean-Louis-Armand de Quatrefages de Breau," DSB, 11: 233-235. 115. Quatrefages, Darwin et ses precurseurs francais, pp. 291-292. 116. Edmond Perrier, La philosophie zoologique avant Darwin, 3rd ed. (Paris, 1896), pp. 129141. Perrier, appointed a professor of the Museum in 1876, announced his support for evolution in 1879. On Perrier's career, see Camille Limoges, "Edmond Perrier," DSB, 10: 522-523. 117. This is the inscription on the statue of Lamarck at the entrance of the Museum. 118. On Russell's interpretation of Cuvier and Geoffroy, see Russell, Form and Function, pp. 31-44, 52-78. On his Lamarckian leanings, see ibid., pp. v-vi; and E.S. Russell, The Interpretation of Development and Heredity: A Study in Biological Method (Oxford: Clarendon Press, 1930). Recent studies of Russell include George V. Lauder, "Introduction" to E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology (Chicago: University of Chicago Press, 1982), pp. xi-xlv; and Nils Roll-Hansen, "E.S. Russell and J.H. Woodger: The Failure of Two Twentieth-Century Opponents of Mechanistic Biology," Journal of the History of Biology 17 (1984): 399-428.

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Index

Academie de Medecine, 102, 156, 161 Academie des Sciences: in ancien regime, 13, 15, 18, 20, 21, 38; section of anatomy and zoology, 38, 243-44, 252n79, 262n57; recreated as First Class of the Institut, 3839; function of in nineteenth century, 6465, 258n95, 258n96; Cuvier-Geoffroy debate before, 143-55; continuation of debate before (1830-1832), 166-67; list of zoologists who were members of, 243-44 permanent secretaries: institution and function of, 42, 64; elections of, 120-21, 178-79; list of, 244 politics of science at: and election of Blainville, 67, 119-21; Arago and Brongniart as rival leaders of factions, 67, 200, 283n96; factions and control of patronage at, 107, 120-21, 144; conflict between Arago and Cuvier, 121, 160-61; conflict on openness of Academie, 145, 149, 160-61, 269n59; critique of Academie by outsiders, 177, 182, 195, 198, 199-200. See also Politics of science Adanson, Michel, 201 Agassiz, Louis, 170, 206, 225, 289n82 Ampere, Andre-Marie, 111, 170, 267nl8, 274nl0 Analogy: Geoffroy's use of term, 70, 225; distinguished from homology by Owen, 70-71, 227; as used by Macleay,.289n87. See also Homology Anatomy, comparative. See Comparative anatomy Anatomy, philosophical. See Philosophical anatomy Annales des sciences naturelles: founding of and editors, 37, 63, 107, 188, 216; and debate, 156, 157, 167; and Geoffroy in 1830s, 185

Arago, Francois: as a patron of Blainville, 67, 121; as leader of a faction at the Academie, 67, 200, 283n96, 284nl09; rise to power and conflict with Cuvier, 107, 120-21, 144, 160-61; and Geoffroy, 281n49 Archetype: Goethe and, 159; Milne Edwards on, 218; Owen's, 226, 227, 228-29, 232; Darwin and, 231; persistence of after Darwin, 232 Aristotle, 3, 44-45, 46, 158, 235; homologies in work of, 70, 86; Cuvier attributes concept of homology to, 91, 148; Geoffroy distinguishes his doctrine from that of, 149-50, 153; Cuvier contrasts to Plato, 170 "Arrests of development": in work of Geoffroy, 88, 152, 153, 167, 203; in work ofSerres, 123-24, 167, 171-72; applied to teratology, 124, 128; and evolution, 135; opposed by Cuvier, 140, 152, 167, 168; Milne Edwards on, 217, 219, 220, 288n57 Articulates. See Homology Athenee, 63, 66, 195, 254n39 Audouin, Jean-Victor, 172, 203; at Museum, 60, 118, 147, 216; early career and support from Brongniart, 63, 114, 257n90; and Cuvier, 65, 113, 115-16, 117-18; and philosophical anatomy, 94, 106, 110, 11418, 217; collaboration with Milne Edwards in marine biology, 216-17 Autenrieth, Johann von, 30, 39 Aye-aye, 27 Baer, Karl Ernst von, 205-6, 207, 219, 224, 225, 228, 231 Balance of organs, principle of, 116, 123, 159; in work of Geoffroy, 76, 83, 151 Balzac, Honore de: on Cuvier and Geoffroy, 58, 175, 176, 188, 190-92; satire of debate, 108, 177, 193, 194-95

293

294

INDEX

Barry, Martin, 223, 224-25 Belon, Pierre, 70 Bell, Charles, 56 Bernardin Saint-Pierre, Jacques-Henri, 21 Berthollet, Claude Louis, 20, 72, 73, 77, 78, 80 Bichat, Xavier, 50 Billarderie, marquis de la, 17 Biot, Jean-Baptiste, 114, 120, 196 Blainville, Henri Marie Ducrotay de career: relations with Cuvier, 52, 65, 66-67, 119, 147, 168; at Museum, 60, 66, 67, 118, 144, 147, 212; at Faculte des Sciences, 62, 66, 114, 208, 212, 214; early career and patronage of Cuvier, 63, 6667; personality, 66-67, 200, 212, 215, 286n30; election to Academie, 67, 107, 119-21; as leading French zoologist, 212; as historian of science, 214-15 and Cuvier, Geoffroy, and the debate: in argument over openness of Academie, 160-61; and Duges, 163, 164, 165; borrows from both Cuvier and Geoffroy, 171, 203, 213; on the debate, 172, 177, 208, 212-15, 287n38; conflict with Geoffroy, 180; critique of Cuvier and Geoffroy, 203, 213, 214-15, 252n82, 287n38 doctrine: five "types" and animal series, 45, 58, 119-20, 133, 138, 212-14; philosophical anatomy, 45, 72, 93, 94-95, 97-98, 120, 213-15; relations of science and religion, 57-58, 213; satire of his terminology, 200 Blumenbach, Johann Friedrich, 39, 90, 126, 127, 159, 250n46, 270n85 Bonnet, Charles: chain of being of rejected, 25, 26, 28, 49, 89; as possible stimulus to Geoffroy, 89; Cuvier on, 49, 146 Bory de Saint-Vincent, Jean Baptiste George Marie, 63, 158, 171 Bosc, Louis Augustin Guillaume, 91, 115 Botany, 13, 15, 16, 96; Geoffroy on, 76-77, 179 Bourdier, Franck, 2, 90, 131 Bridgewater Treatises, 56, 224 Brisson, Mathurin-Jacques, 19, 20, 30 Britain, biology in: religion and science, compared to France, 56-58, 141; debate publicized, 144-45, 223; synthesis of teleology and morphology in, 205, 22230; debate and evolution in, 230-32 Brongniart, Alexandre: early career, 16, 21, 61, 63, 84; and Geoffroy's entrance to Museum, 21-22; and Cuvier, 38, 43; as head of a faction, 67, 200, 283n96,

284nl09; proteges of, 114, 157, 218; and Raspail, 196, 198; and nepotism, 198, 199, 283n96, 284nl09 Brongniart, Adolphe, 180, 196 Brongniart, Antoine-Louis, 17 Broussonet, Pierre-Marie-Auguste, 15, 21, 61, 92 Bruguiere, Jean-Guillaume, 15, 16, 19, 21 Buffon, Georges-Louis Leclerc, comte de, 47, 127, 183; on classification, 12, 14, 15, 23; Histoire naturelle, 14, 16, 22-26 passim, 38, 49; on environment and modification of species, 14, 23, 34, 187, 211-12; as intendant at Jardin du Roi, 14, 16-17; work as stimulus to Geoffroy, 22-27, 2829, 90, 187; unity of plan in the work of, 23-24, 28-29, 211 reputation of, 15, 17, 26, 201; Cuvier on, 22, 25, 49; Geoffroy on, 134, 179, 186-87; Goethe on, 160; Flourens on, 211-12 Burnet, Thomas, 49

Cahn, Theophile, 246n4 Camper, Peter, 159 Candolle, Augustin-Pyramus de, 93, 96, 103 Carpenter, William B., 207, 223, 224, 225 Career paths of zoologists, 62, 63-64 Catastrophism: Cuvier and, 44, 58, 253nl5; Geoffroy and, 134 Chain of being: Daubenton on, 25, 26; Geoffroy and, 27, 28, 89; Cuvier and, 32, 34, 48, 49, 50-51, 137, 250n59; Lamarck and, 50-51; Blainville and, 58, 119, 138; Sand and, 189-90 Chambers, Robert, 68 Charles X, king of France, 1, 108, 143, 158, 166 Chevreul, Eugene, 192 Classification: Buffon on, 12, 14, 15, 23; Daubenton on, 12, 22, 23, 24-26, 32; Linnaean system of taxonomy, 14, 15, 21, 23, 25, 33; continuity (nature makes no leaps), 25, 28, 32, 34, 50; doctrine of multiple affinities, 25, 28; Blainville and animal series, 119-20, 212-14; Duges's system, 163; satires of, 194-95, 200 Geoffroy on: divergent attitudes of Geoffroy and Cuvier, 12, 34, 45, 83, 209; contributions to, 25-26, 27-29, 33-34, 75, 83, 96, 261n49; skepticism toward, 23, 28, 38, 83; desire to supersede, 69, 83, 180, 181; of malformations, 126-27; of man and primates, 185, 187

INDEX natural system of classification: Cuvier and, 12, 32-34, 41, 43-46, 49, 50, 51; and French botany, 15; philosophical anatomy as an aid in revealing, 96, 204, 219, 223; application to teratology, 126; Flourens on, 210-11; embryology as an aid in revealing, 216, 218, 219, 225; Milne Edwards on, 218, 219; Barry on, 225 four embranchements: Cuvier and, 41, 4445, 51; different from abstract types, 3, 45; challenged in 1820s, 106; challenged by Blainville, 119-20, 213, 215; embryological support for, 205, 219; acceptance of in generation after debate, 203, 211-12, 215, 219-21; as bulwark against evolution, 211, 221; upheld in Britain, 225, 228. See also Homology College d'Etampes, 19 College de France: before 1793, 14, 20, 41, 61; chair of natural history, 41, 61, 66, 119; Cuvier's course in history of science, 48, 145, 157, 165, 169-70; list of holders of chairs related to zoology, 241; and experimental physiology, 257n83; College de Navarre, 19, 20 College du Cardinal Lemoine, 20 Comparative anatomy: Daubenton and, 14, 24-25, 30, 67; Vicq d'Azyr and, 14, 25; Cuvier and renaissance of, 4, 30, 42-43, 47; Cuvier and introduction into zoology, 4, 33, 39, 42, 252n81; as replacement for experiment (Cuvier), 47; difference from philosophical anatomy, 70, 85-86, 98-99, 120, 149-50, 153, 171-72; incorporation of German embryology into, 207; introduced into England by Owen, 225 Comte, Auguste, 48, 254n35 Conde, prince of, 18 Condillac, Etienne, 47 "Conditions of existence," 224, 231; Cuvier on, 41, 43, 46, 48, 58, 137, 148 Connections, principle of, 115, 146, 231; in work of Geoffroy, 85, 89, 98-99, 100, 115, 126, 153 Conte, Jacques-Nicholas, 76 Controversies in science, 1-2 Cordier, Pierre-Louis-Antoine, 179 "Correlation of parts," 33, 41, 46, 47, 58 Coste, Victor, 180 Coulomb, Charles Augustin, 32 Creations, multiple: rejected by Geoffroy and others, 133, 134, 184, 212; Cuvier's ambiguity concerning, 44, 133 Crocodiles. See Paleontology

295

Cumul, 61; and Cuvier, 53 Cuvier, Clementine, 142 Cuvier, Frederic: career, 36, 60, 63, 66, 186; scientific work of, 36; disciple of his brother, 65, 66, 172, 202; review of Philosophie anatomique, 103, 265n 119; defeated by Blainville for seat in Academie, 67, 119, 121; dispute with Geoffroy on his brother and religion, 281n48 Cuvier, Georges: training and early work, 2930; rapid rise to success, 38-39, 250n50; in public education and administration, 53, 54-55, 84; personality, 66, 67, 72, 142; decides not to go to Egypt, 84; death of, 170; bibliography and manuscripts of, 249n44, 252n4, 254n38 at Museum: arrival at, 11-12, 29, 30-32; collaboration with Geoffroy, 12, 29-34, 38; professor of comparative anatomy, 34-37, 60, 143; preface to Nouvelles annales, 169 at Academie des Sciences: election to, 12, 38-39; permanent secretary of, 42, 53, 6465, 84, 160-61; exploits eloges of, 42, 48. See also individual subjects of eloges. at College de France: professor of natural history, 41, 61, 119; course in history of science, 48, 145, 157, 165, 169-70 as France's leading naturalist: great authority of, 40-41, 42, 48, 52, 55; disciples of, 65-68, 95, 172, 202-3; authority under attack in 1820s, 106, 1078, 119-21, 124-25; strains patience of colleagues in 1830s, 165-66; as a public scientist, 254n53. See also Patronage political attitudes, 30, 53-55, 59, 108, 120, 143, 255n57, 255n60, 256n78; Lyell on, 255n56, 266n9 religious attitudes, 41, 46, 53, 54, 55-59, 108, 137-38, 151-52, 154, 170, 256n78 criticism of: as orthodox in religion, 58-59, 188, 189, 190, 194, 266n9, 281n48, 289n83; as a champion of analytical science, 175; as a manipulator of patronage, 194-95, 196, 197, 199; as an uninventive fact collector, 197, 199 works: Lecons d'anatomie comparee, 34, 36, 41, 42-43, 44, 65, 85-86, 168; Recherches sur les ossemens fossiles, 34, 42, 43-44, 49, 57, 58, 82, 131; Le regne animal, 34, 42, 43, 51, 59; Tableau elementaire, 44; Histoire naturelle des poissons, 65, 105, 136, 138, 139-40, 272nl21; article "Nature," 136-38, 151, 153.

296

INDEX

Cuvier-Geoffroy debate: as subject for historical study, 1-2; summary of issues, 2-9, 141-42, 172-73, 235-37; as a particularly French controversy, 6; many meanings of, 9, 10, 178, 207, 236-37; role of evolution in the conflict, 82, 130, 135, 153, 154; other naturalists avoid involvement in, 108-9, 142, 147, 161, 165, 171-72; precipitating factors, 139, 142, 143-44, 145-47; and Revolution of 1830, 143-45, 157, 173; publicized in France and England, 144-45, 154-56, 173-74, 177, 223; course of, in 1830, 14555; continuation of controversy to 1832, 145, 165-70; Duges's intervention in, 161-65; need for reconciliation, 171, 2023; of interest to nonacademicians, 173-74, 175-78; synthesis by the next generation in France and Britain, 202-230; and Darwin, 230-32; controversy reawakened in France, 232-33 discussion of outcome of the contest: Russell on, 3, 235, 246n6; historiographical discussion, 3, 5, 144, 170-71, 204, 235-36; perceptions of contemporaries that Geoffroy won or would win in the end, 157, 190, 191-92, 193, 209-10; Flourens on, 211, 233 interpretations of: as a matter of evolution, 2-3, 5, 154, 156, 171, 207, 233-35, 236; teleology versus morphology, 3, 5, 205, 226, 235, 236; as progress versus conservatism, 145, 155-58 passim, 171; as a matter of anatomy, 156-58, 161, 172, 191-92; as a matter of philosophy and religion, 158, 176, 188-93, 194, 204, 236; as analysis versus synthesis, 159-60, 161, 168; and politics of science, 160-61, 173, 176, 177, 193-201, 204, 236, 289n83; as facts versus ideas, 161, 202, 209-10, 233, 236; professional scientist versus philosopher of nature, 177, 204; interpretations by scientists versus outsiders, 201, 204; as one plan versus four, 205, 211, 212, 233-34, 236, 288n71 commentators on: Goethe, 1, 143-45 passim, 156, 158-61, 210; Russell, 3, 5, 235, 237, 246n6; newspapers and journals, 144, 155-61; Isidore Geoffroy, 161, 172, 177, 202, 208-11, 215, 236; naturalists in immediate aftermath, 171-72; Cuvier, 172; Blainville, 172, 177, 208, 212-15, 287n38; Flourens, 172, 177, 202, 204-5, 208, 210-12, 215, 233-34, 236; Balzac, 175, 191-92, 193, 194-95; Sand, 190;

Quinet, 193, 236; Raspail, 197, 236; Milne Edwards, 172, 222; Owen, 226, 289n83; Darwin, 230; Quatrefages, 234, 236; Perrier, 234-35, 237; Michelet, 283n82; Whewell, 288n71 Dagognet, F., 90 D'Alembert, Jean Le Rond, 57 d'Angers, David, 188 Danielo, Mien, 279n5 Darwin, Charles: reception of in France, 130, 233-35; and the Cuvier-Geoffroy debate, 144, 230, 290nl05; teleology, morphology and the development of his theory, 206, 207, 223, 225, 230-32, 290nl07 Darwin, Erasmus, 201 Daubenton, Louis-Jean-Marie: as Geoffroy's mentor, 11, 12, 19, 21-22, 31, 90; encourages specialization and professionalization, 24-27, 37; Cuvier on, 22, 25, 49; Goethe on, 160 career, 14, 20, 41, 61, 84; at Jardin du Roi and Museum, 14, 16-18, 17, 19, 20, 22; collaboration with Buffon, 14, 16, 24-25, 49 scientific doctrine: definition of natural history, 26; on state of zoology in 1793, 11, 12, 39; on classification, 12, 22, 23, 24-26, 32; and comparative anatomy, 14, 24-25, 30, 67 Daudin, Henri, 41, 42, 51 Decade egyptienne, La, 73, 74 Decade philosophique, 27, 36 De Gosse, Isidore (pseud.). See Salles, B.-I. de Delalande, Pierre-Antoine, 98 Desfontaines, Rene, 17, 21 Desgenettes, Rene-Nicholas, 76 Desmarest, Anselme-Gaetan, 61 Dictionnaire classique des sciences naturelles, 63 Dictionnaire classique d'histoire naturelle, 63 Diderot, Denis, 14 Division of physiological labor, 218, 222 Dolomieu, Deodat de, 73 Dombey, Joseph, 74 Donne, Alfred, 157-58, 197 Duges, Antoine, 157, 161-65, 171 Dulong, Pierre-Louis, 178-79 Dumas, Jean-Baptiste, 63, 179; appreciation of Geoffroy, 93, 188, 192; supports Milne Edwards, 188, 216 Dumeril, Constant: career, 38, 60, 61, 65-66, 84, 92, 114, 122; as disciple of Cuvier, 6566, 115-16, 172, 202; and philosophical anatomy, 72, 95, 105, 115-16, 264n99;

INDEX reports on Audouin and Milne Edwards, 115-16, 117, 217; in controversy over openness of Academie, 161; at Geoffroy's funeral, 192; competition for Faculte des Sciences, 264n99 Dunal, Michel-Felix, 96 Dupetit-Thouars, Marie Aubert, 96 Dutrochet, Henri, 72, 95, 172, 180 Duverney, Joseph-Guichard, 87 Duvernoy, Georges Louis, 61, 62, 200; as disciple of Cuvier, 40, 65-66, 172, 202, 204; and philosophical anatomy, 202, 204 Ecole Centrale du Pantheon, 32, 61 Ecoles centrales. See Secondary schools Ecole de Medecine (Paris), 31, 61 Ecole Normale (1795), 25, 26, 38 Ecole Normale (1808), 62, 66, 93 Ecole Veterinaire d'Alfort, 14, 61 Egypt, French expedition to, 72-81, 82 Electric fishes (Geoffroy on), 78, 83, 182 Embranchements. See Classification Embryology, 117, 159, 224-25; Geoffroy's use of fetal anatomy in determining homologies, 83, 85, 87-88, 88-89, 91, 150; in work of Serres, 122-124, 128, 171-72; in Cuvier's studies (1832), 167; insufficiently understood by Cuvier and Geoffroy, 205, 224, 206-7, 285nlO; importance of German embryology for synthesis of morphology and teleology, 205-7, 222, 223, 285n8, 285nlO; branching conception of development, 206, 218, 219, 222, 228, 231; in work of Milne Edwards, 207, 216-20 passim; embryological parallels explained by Darwin, 231, 232. See also Generation; Arrests of development Empiricism. See Methodology, scientific Encyclopedie methodique, 14, 23, 168, 247n9 Epigenesis. See Generation Evolution: significance of debate for, 5, 207; importance of morphological approach to, 5; and Cuvier-Geoffroy debate, 5, 82, 130, 135, 153, 154, 156, 173; and philosophical anatomy, 135, 136; and debate in France after Darwin, 207, 232-35 Geoffroy and: early interest in role of environment in modifying animal structure, 23, 24, 34, 75-77, 81; collections from Egypt and evolutionary timescale, 52, 74, 82, 132, 212; papers in 1820s, 130-34, 178; in early 1830s, 13436, 153, 166, 167; post-1832, 179, 181,

297

184; Flourens disassociates from rest of Geoffroy's doctrine, 212 Cuvier and: to 1820, 44, 48, 49, 51-52, 58, 82; links to unity of composition, 130, 137-38, 154; opposes in 1830s, 166, 16869, 170 in work of others: Buffon, 23, 33-34; Lamarck, 37-38, 50-52, 82, 130, 132, 134, 153; Lacepede, 38; reception of Darwin in France, 130, 233-35; Serres, 136; development of Darwin's theory, 207, 230-32; Isidore Geoffroy, 209; Flourens, 210-11, 233-34; Blainville, 212; Milne Edwards, 221, 234; Owen, 226, 232, 289n84; Quatrefages, 233, 234 Experimentation: Cuvier on comparative anatomy as replacement for, 47; in work of Geoffroy, 76-77, 128-29, 132, 271nl03 Extinction, 43, 44, 58 Fabricius, Johann Christian, 93 Faculte de Medecine de Paris (ancien regime), 16, 20 Faculte de Medecine, 65 Faculte des Sciences (Paris): Cuvier's influence on appointments to, 53, 62, 66; initial organization and professors, 62, 257n88, 257n89; becomes locus of zoological teaching, 216; list of holders of chairs of zoology, 242. See also individual professors Faujas de Saint-Fond, Barthelemy, 17 Final causes, 24, 158; Geoffroy on, 24, 69, 71, 153; Cuvier on, 41, 46, 47; Isidore Geoffroy on, 210; Flourens on, 212, 223, 265nl34; Blainville on, 213, 223; Milne Edwards on, 221, 223; understanding of modified in Britain, 223-24, 227, 231 Fishes. See Homology Fishes, electric. See Electric fishes Flourens, Pierre: as disciple of Cuvier, 47, 65, 66, 119, 125, 210; career, 60, 61, 103, 125, 210; as disciple of Geoffroy, 66, 104, 11819; and Academie des Sciences, 66, 119, 178-79, 210, 269n59 scientific doctrine: eloge of Cuvier, 40, 45, 210-11; philosophical anatomy, 103-4, 154, 211, 265nl34; combines elements from Cuvier and Geoffroy, 171, 203, 210; on the debate, 172, 177, 202, 204-5, 208, 210-12, 215, 234-35, 236; eloge of Geoffroy, 202, 211; reinterprets Cuvier's views, 204-5, 211; and evolution, 211-12;

298

INDEX

Flourens, Pierre (continued) on Blainville, 212, 215; and final causes, 212, 223, 265nl34 Fourcroy, Antoine-Francois, 17, 20, 21-22, 98 Fourier, Joseph, 73, 80, 120 Fraysinnous, Denis-Antoine Luc, 58, 108 French Revolution: and Cuvier, 12, 30, 53-54, 255n60; disruption of science during, 15, 16, 18-19, 20, 21, 39; and founding of Museum, 15-19, 21, 35; and Geoffrey, 20-21 French science: institutional transformation of and role in the debate, 7-8, 60, 63-64; and importance of patronage politics, 8-9, 59, 64, 258n94; critique of by outsiders, 193-201; becomes more bureaucratized and insulated from the public in generation after debate, 204, 220; issue of the decline of, 258n94. See also Specialization; Professionalization Functionalism: Russell and, 3, 235; Latreille and, 113; Geoffroy fails to banish from his writings, 203; strict functionalism rejected in 1830s and 1840s, 205; Milne Edwards's synthesis of with philosophical anatomy, 217-22 in the work of Cuvier: before 1820, 4, 33, 41, 43, 44-46, 47, 51; in 1820s, 105, 137, 139-40; arguments during debate, 148, 150-51, 154; shortcomings of doctrine, 203. See also Teleology Furcula (wishbone), 70, 74-75, 86-87 Gall, Franz Joseph, 28, 104, 118, 119, 190 Gazette medicale de Paris, 156-57, 163, 166 Generation: Cuvier on, 49-50, 91, 167; Geoffroy on, 75-77, 79; homologies of organs of, 95 epigenesis and preformation: Geoffroy on, 24, 75, 127, 132, 205; Cuvier on, 48, 4950, 127, 205; Serres on, 122, 167, 171; Milne Edwards on, 220 spontaneous generation, 120, 169, 221; Cuvier on, 48, 49; Geoffroy on, 276n49 See also Embryology Geoffroy, Etienne-Francois, 19 Geoffroy, Jean-Gerard, 19 Geoffroy Saint-Hilaire, Anai's, 96 Geoffroy Saint-Hilaire, Etienne: mentors of, 11, 19, 20-29, 31; early work (to 1799), 12, 22-26 passim, 27-29, 38; education, 19-20, 29; support of Cuvier, 12, 29, 3032, 39, 40, 178; collaboration with Cuvier, 12, 29-34, 38, 279n5; expedition to

Portugal, 92; bibliography and manuscripts of, 248n22 and Academie des Sciences, 119; election to, 38-39, 84, 90, 92; as president of (1833), 165, 179, 197-98, 277n83, 281n45; candidacy for permanent secretary, 17879. See also Cuvier-Geoffroy debate and Museum d'Histoire Naturelle: obtains first chair of vertebrate zoology, 11, 19, 20-22; as professor, 22, 34-38 passim, 62, 80, 81, 83; report on collections from Egypt, 82; difficulties with leadership of in 1830s, 180, 186 and Faculte des Sciences, 62, 90, 92-93, 114, 121, 188 expedition to Egypt: experiences in, 72-81, 84, 125-26, 128; significance for later career, 71, 72-73, 80-81; relations with Cuvier during, 73-80 passim, 261n30; animal mummies brought back from, 74, 82, 132, 212; subtle fluid theories, 78-80, 100-101; papers after return to Paris, 83, 85, 86; 1830s theories extending work in, 178, 182 personal: marriage and family, 96; personality, 72, 142; view of himself as engaged in a philosophical mission, 149, 152, 153, 176, 177, 186; death and funeral, 188, 192-93 politics and religion: political views and activities, 20, 96-97, 277n81; belief in a deistic universe, 24, 79, 81, 133, 176, 18385; response to charges of irreligion, 14041, 155, 179, 183; dispute with F. Cuvier on G. Cuvier and religion, 281n48 and professional colleagues: concern with professional recognition, 27, 74, 77, 78, 79-80, 81; strains patience of colleagues in 1830s, 165, 176, 179-80, 280nl4; disciples of, 121-22, 202 others' perceptions of: as founder of philosophical anatomy, 93, 211, 220, 290nl09; as leader of a progressive school, 155, 157, 177; appeal of to nonacademicians, 175-78, 184, 188-201. See also Cuvier-Geoffroy debate; Philosophical anatomy activities in the 1830s: philosophical writings, 173, 178, 179-87; seeks out popular support, 176, 187, 189, 190, 19798; salon, 176, 187, 279n5; lack of response on part of academicians, 179, 183 works: Philosophie anatomique (1818), 34, 69, 71, 85, 93, 97-104; Catalogue des

INDEX

mammiferes (unpublished), 83, 261n49; Cours de l'histoire naturelle des mammiferes, 105, 136, 140; Philosophie anatomique, vol. 2 (1822), 119, 126; Principes de philosophie zoologique, 144, 151, 155, 157, 171, 223; Etudes progressives, 176, 179, 180-81, 182, 184; Notions synthetiques, 179, 181, 182, 183, 187, 189. See also individual people and concepts Geoffroy Saint-Hilaire, Isidore: career, 60, 63, 96, 188, 198, 199, 208; teratology, 126, 127; doctrine combines elements from Cuvier and Geoffroy, 171, 203, 208; philosophical anatomy, 208-9, 264nl07, 264nl08 and his father, 83, 89, 190; disciple of his father, 122, 202, 209; on the debate, 161, 172, 177, 202, 208-11, 215, 236; embarrassed by his father, 180, 280nl3; retreats from positions taken by his father, 183, 202, 209, 280nl3; reinterprets his father's views, 205, 209; contrast in personality to his father, 208-9 Geoffroy Saint-Hilaire, Pauline, 96 Geoffroy Saint-Hilaire, Stephanie, 96 Gerard, Frederic, 177, 198, 199 German biology: compared to French in 1790s, 39; philosophical anatomy compared to in France, 90, 106-7, 270n85; defended by Goethe, 159; significance of German embryology for synthesis of teleology and morphology, 205-7. See also Naturphilosophie Globe, Le, 157, 170 Gmelin, Johann Georg, 27 Goethe, Johann Wolfgang von, 22; on the debate, 1, 2, 5, 143-45 passim, 156, 15861, 210; morphological work of, 70, 90, 158-59; role in debate noted by others, 156, 161, 192, 193, 202, 287n38; Geoffroy's appreciation of, 159, 166-67, 178, 187 Gouan, Antoine, 15, 86 Grandville, J.-J., 194 Grant, Robert Edmund, 206, 225 Guerin, Jules, 156-57 Guizot, Francois, 55 Guyton de Morveau, Louis Bernard, 20 Haeckel, Ernst, 124 Hahn, Roger, 57 Hartmann, E.G.F., 30 Hauy, Rene-Just, 16, 20-21, 31, 32, 90 Herman, Jean, 39

299

Hetzel, Pierre Jules, 194 Homology: as key concept of philosophical anatomy, 4, 69, 85; Geoffroy and the emergence of the concept of, 28-29, 6971, 84-85, 89-90, 149; as ideal pattern, 69, 88, 173, 232; origin of term, 70-71, 106-7, 225, 227, 266n4, 289n87; method of intermediary forms in determining, 8788, 98-99, 117, 126, 217, 218; Geoffroy's physiological explanations for, 100, 11011; search for becomes an integral part of French biology, 203; embryological criterion for, 206; in Britain, 224-29, 23132; significance of to various areas of biology, 236 in skeleton of vertebrates, 122-23, 228-29; Geoffroy on hyoid bone, 149-53 passim, 169; Cuvier on hyoid bone, 150-51, 15354 between fishes and other vertebrates, 94, 206-7, 229; Geoffroy on, 83-89, 97-103, 109, 141, 150, 152; Cuvier on, 85-86, 87, 101-2, 140, 141, 153-54 in the vertebrate skull, 95; Geoffroy on, 8889, 109, 132, 141, 271nl07; Cuvier on, 96, 97, 140, 141 in other vertebrate systems, 95, 123; Geoffroy on, 83, 109 in exoskeleton of articulates: in work of French naturalists, 93-95, 106, 110, 11218, 217-19; Cuvier's critique, 94, 113, 115-16 between vertebrates and invertebrates, 17172, 264n99 between articulates and vertebrates: 117, 163, 217, 267nl8, 287n46; Geoffroy on, 93, 105, 106, 109-12, 115; Latreille on, 106, 110, 113-14, 267n25; Cuvier's critique, 111-12, 113-14, 164 between mollusks and vertebrates, 192; Latreille on, 114, 147; Meyranx and Laurenceton, 145-46, 274n10; Geoffroy on, 145-46, 152, 274nl5; Cuvier's denial of, 146-48, 167 See also Homology, serial; Philosophical anatomy Homology, serial, 93; in France, 70, 94, 162, 218; in German science, 70, 106-7; Owen and Darwin and, 227, 231, 232, 290nl07. See also Vertebral theory of the skull Howler monkey, 28, 150-51 Hugo, Victor, 188 Humboldt, Alexander von, 170, 255n56, 255n64 Hyoid, homologies of, 100, 149-51, 152-54, 169

300

INDEX

Imperial University. See University of France Institut d'Egypte, 73, 74-80 passim Institut de France, First Class of. See Academie des Sciences Jardin du Roi (Jardin des Plantes), 11, 14, 15, 16-18 Joly, Nicholas, 208 Jourdan, Antoine-Jacques-Louis, 180, 197, 264nl07, 265n119 Journal des debats, 157-58, 197 Journal d'histoire naturelle, 16, 31 Jussieu, Antoine-Laurent de, 15, 17, 30, 34, 67, 186, 201 Kant, Immanuel, 106, 107 Karlsschule, 29-30 Kepler, Johannes, 187 Kerner, Johann Simon von, 29 Kielmeyer, Carl Friedrich, 29, 39, 90, 107, 123, 158 Knox, Robert, 223, 290nl05 Kohlbrugge, J.H.F., 90, 246n5 Lacepede, Bernard-Germain-Etienne, 30, 115, 117, 121; at Jardin du Roi, 14, 17, 19, 21, 22; scientific work, 14, 38, 86; Geoffroy and, 22, 75, 82, 84, 91, 92; at Museum, 37-38, 60 Lachat (friend of Audouin), 114 Lacroix, Sylvestre Francois, 38 Lafontaine, Jean, 31 Lagrange, Joseph Louis, 20, 78, 80 Lakanal, Joseph, 18, 22, 192 Lamarck, Jean-Baptiste: as naturalistphilosopher, 37-38, 47 career, 91; at Museum d'Histoire Naturelle, 11, 19, 21, 60; supports Cuvier coming to Museum, 12, 30; in ancien regime, 16, 17; division of his chair at Museum, 67, 114, 118, 144, 147 scientific doctrine, 58, 100, 127, 133, 183; and invertebrates, 37, 45; theory of evolution and chain of being, 37-38, 5052; Philosophie zoologique, 37, 52, 82, 98, 132; on mummies from Egypt, 82; theory of evolution compared to Geoffroy's, 130, 132, 134, 136, 184 reputation: admired by Blainville, 38, 214; Cuvier on, 49, 52, 58, 168-69, 254n39; work ignored, 52, 68; Geoffroy's appreciation of, 132, 133, 153, 186, 199;

Cuvier's eloge of, 165, 168-69, 214, 278n92, 278n94; admired by critics of French science, 198, 199, 201, 284nl03; debate seen as conflict between Cuvier and, 207, 233, 235, 237 La Mettrie, Julien Offray de, 14, 201 Lamouroux, Felix, 131 Language of science: Daubenton on, 26; ambiguous language of philosophical anatomy, 88; Cuvier on, 138, 148, 149, 150; satire of esoteric terms in science, 200 Laplace, Pierre-Simon, 32, 201; and Cuvier, 38, 47, 50, 120; and science and religion, 57; as leader of physical scientists, 67, 80, 84, 120; Laplacian program for physics, 78-79, 120, 121 Larynx, homologies of, 100, 102 Latreille, Pierre-Andre, 61, 115; at Museum, 60, 112, 114, 118, 147; and philosophical anatomy, 94, 106, 110, 112-14, 147, 267n24, 267n25; and Cuvier, 112, 113-14, 118, 147, 274n12; and debate, 145, 147, 172 Laurencet, 145-49 passim, 155, 197, 234 Laurillard, Charles Leopold, 58-59, 60, 65-66, 148, 172 Lavater, Johann Kaspar, 190 Lavoisier, Antoine-Laurent, 20, 30, 32, 47, 90, 168 Laws of organization: Cuvier and, 43, 47; debate and, 5, 142, 171, 173, 203; Geoffroy's search for, 6, 38, 140, 142; reveal a larger purpose, 205, 223, 224, 225; Milne Edwards and, 221. See also Soi pour soi Leibniz, Gottfried Wilhelm, 49 Lhomond, Charles-Francois, 20 Linnaean system. See Classification Linnaean naturalists: in France, 15-16, 25, 37; Cuvier as, 29, 30, 67 Linnaeus, Carl, 15, 29, 30, 32, 44, 201. See also Classification Louis XVIII, king of France, 53, 55, 108 Louis-Philippe, king of France, 53, 143, 166, 176 Lubosch, Wilhelm, 246n5 Lycee, 21 Lycee des Arts, 21 Lycees. See Secondary schools Lyell, Charles, 52, 58, 82, 255n56, 255n64, 266n9 Macleay, William Sharp, 289n87 Magasin encyclopedique, 36, 103

INDEX Magendie, Francois, 47, 65, 122, 164, 179 Maillet, Benoit de, 14, 49 Makis, 24, 27, 28 Malformations. See Teratology Malus, Etienne Louis, 74 Man, classification of (Geoffroy), 185, 187 Marsupials, 27, 28, 184, 261n49 Martin Saint-Ange, Gaspard-Joseph, 158, 171 Maupertuis, Pierre Louis Moreau de, 14, 127 Meckel, Johann Friedrich, 106, 107, 123, 127, 158, 267nl8 Meckel-Serres Law, 123 Medical natural history, 31, 61 Medicine, and natural history, 63 Mertrud, Antoine-Louis-Francois, 12, 17, 31, 32, 41 Mesmer, Franz Anton, 54 Methodology, scientific: as issue in the debate, 6-7, 142, 173; next generation appreciates importance of ideas, 203 Cuvier on: early opposition to theorizing, 12, 30, 41, 46-53, 54; post-1820 insistence on primacy of facts, 92, 105, 138-39, 16770 Geoffroy on: penchant for generalization, 12, 27, 38, 73; defends theorizing, 134, 139, 141, 153, 275n37 in work of others: Blainville on, 120; Serres on, 125; Isidore Geoffroy on, 209-10; Milne Edwards on, 220-21; Flourens on, 233-34; Quatrefages on, 233, 234; Darwin rejected on basis of, 233-34; Dutrochet on, 172 Meyranx, Pierre-Stanislas: and the debate, 145-49 passim, 155, 197, 234, 274nlO; in novels of Balzac, 175, 191 Michelet, Jules, 192, 283n82 Millin, Aubin-Louis, 15, 16, 30-31, 37 Milne Edwards, Henri: career, 62, 63, 188, 216, 257n86; and philosophical anatomy, 94, 207, 216-19, 222, 274nl2, 287n46; and synthesis of ideas of Cuvier and Geoffroy, 171, 203, 215-22; and the debate, 172, 222, 225; on Geoffroy, 177, 220, 287n46; and embryology, 207, 21620 passim, 222, 288n57; becomes leader of French zoologists, 213, 215-16; and division of physiological labor, 218, 222; and development of Darwin's theory, 230, 231 Mollusks. See Homology Monge, Gaspard, 72, 73, 77, 78, 80, 182 Moniteur, 143, 158 Monotremes, 181, 184, 261n49 Monsters. See Teratology

301

Moquin-Tandon, Alfred, 96, 125, 162, 179, 180, 270n79, 281n50 Morphology: importance of in French biology, 3, 4-5; Cuvier's embranchements different from abstract types of, 3, 45; Goethe and, 70, 90, 158-59; use of the term, 159, 232; synthesis of teleology and, in Britain, 205, 222-30; and development of Darwin's theory, 207, 230-32; heyday of in late nineteenth century, 232; Geoffroy as a "pure morphologist," 235. See also Philosophical anatomy; Homology; Teleology Muscular contraction, Geoffroy's theory of, 77, 79 Museum d'Histoire Naturelle: founding of, 11, 16-21; as research center, 11, 34-37, 42, 248n21, 251n74; professorial chairs and subordinate positions, 60; elections to vacant chairs, 64, 258n96; satire of professors, 177, 193, 198-201; list of holders of chairs related to zoology, 23840. See also individual professors; Jardin du Roi. Napoleon I, emperor of France, 57, 61, 64, 84; in Egypt, 72-80 passim; and Geoffroy's subtle fluid theories, 78, 182; Geoffroy's admiration of, 96-97, 182 National, Le, 158, 171 Natural system. See Classification Natural theology: in France, 13-14, 56-59; and Cuvier, 41, 46, 56-59; in Britain, 41, 46, 56-58, 222, 227, 256n70; and the acceptance of philosophical anatomy in Britain, 205, 223. See also Teleology Naturphilosophie: Geoffroy associated with, 3, 90, 106, 155, 158, 177, 183-84, 193, 246n5; Cuvier's opposition to, 52, 107, 138, 154, 168; different foundation from philosophical anatomy, 90, 106, 107, 266n3; Geoffroy on, 141, 153, 184; Goethe's defense of against Cuvier, 159; Isidore Geoffroy on, 209, 210 Nepotism, 198, 199, 283n96, 284nl09 Newton, Isaac, 78, 201; Geoffroy and, 79, 140, 141, 181-82, 187 Nicard, Pol, 58, 121 Oken, Lorenz: vertebral theory of skull, 106, 107, 115, 159; Geoffroy and, 106, 107, 158, 183; repetition of cranium by rest of body, 107, 138; Cuvier on, 107, 138; Blainville on, 214

302

INDEX

Olivier, Guillaume-Antoine, 15, 16, 19, 21, 30, 61, 84 Operculum (gill-cover), homologies of in other vertebrates: Cuvier on, 70, 101, 140; Geoffroy on, 86, 88, 89, 97-99, 109, 111, 266nl 1; other naturalists on, 97, 103, 206, 229, 265nl 15; resolution through German embryology, 206-7 Orangutan (of the menagerie), 185, 186, 189 Ospovat, Dov, 5, 6, 205, 207, 229-30, 231 Owen, Richard: definition of homology and distinction from analogy, 70-71, 225, 227, 266n4; and the debate, 205, 223, 225-26, 289n83; synthesis of morphology (homology) and teleology, 205, 206, 207, 224, 225-30; career and Cuvier, 226; and evolution, 226, 232, 289n84; and the Archetype, 226, 228-29; and the development of Darwin's theory, 231 Pajou, Augustin, 186 Paleontology: Cuvier and, 38, 41, 43-44, 51, 58, 131, 133, 166; and Geoffroy's theory of evolution, 130-33, 166, 167, 184; Balzac admires Cuvier's discoveries, 190; Blainville and theory of a single creation, 212; and rejection of sufficiency of teleological explanation, 230 Paley, William, 56, 227, 231 Palissy, Bernard de, 201 Pallas, Peter-Simon, 22, 39, 214 Pantheism: German science identified with, by French, 107; Cuvier's opposition to, 138; Geoffroy seen as a pantheist, 175, 183, 194; Geoffroy on, 184. See also Naturphilosophie Pariset, Etienne, 102, 192, 265n119 Pasquier, Etienne, 55, 170 Pasteur-Pouchet debate, 289n83 Patronage: importance of in French science, 8, 59, 64, 258n94; Arago and, 67, 120-21; Laplace and, 67, 120-21; Geoffroy and, 108, 121-22; Brongniart and, 114 and Cuvier: before 1820, 39, 53, 59-60, 6568; use of in 1820s to keep followers in line, 107-8, 112-19, 125, 147; not always successful in managing, 119, 121, 125 Peale, Charles Willson, 34 Pectoral fin (compared to arm), 85-87, 98, 100 Pelletier, Bertrand, 16 Pencati, Joseph Marzari, 254n39 Perfection (in animal kingdom), 221-22 Perrier, Edmond, 162, 234-35, 237 Personalities, role in the debate, 6, 142

Pfaff, Christoph Heinrich, 30, 32, 50, 54, 143 Philosophes, 13, 14, 54, 67, 81, 127, 159 Philosophical anatomy: significance and aims of, 2, 3-5; homology as key concept of, 4, 69; and Cuvier's framework for natural history, 39, 42; origin and use of term, 69, 98, 264nl07; compared to German Naturphilosophie, 90, 106-7; and philosophical botany, 93, 96; difference from traditional comparative anatomy, 70, 85-86, 98-99, 120, 149-50, 153, 17172; enters mainstream of French zoology, 204-22, 223; in Britain, 222-30; persistence of after Darwin, 232; fruitfulness of, 236 in work of Geoffroy: role in the emergence of philosophical anatomy, 4, 23-24, 6972; unity of plan in his work before 1799, 12, 23-24, 28-29; in Egypt, 72-73, 74-75, 83; research program in memoirs of 1807, 84-90, 97, 99; Philosophie anatomique (1818), 85, 97-100; reception and spread of Geoffroy's program to 1818, 93-96, 101-4; perceived as founder of philosophical anatomy, 93, 211, 220, 290nl09; claim for originality, 98-99, 149-50, 153; originality supported by others, 101, 103, 202, 211; extension of in 1820s to articulates, 105-6, 109-12; and teratology, 106, 124, 125-30; in CuvierGeoffroy debate, 145-46, 149-50, 152-53; shortcomings of his doctrine, 203 Cuvier and: guarded approval of before 1820, 90-92, 96, 101-2, 104; contests originality of, 91, 118, 148; contributions to, 96, 97; growing criticism of in 1820s, 111-18, 137-38, 140; linkage to evolution, 137-38, 154, 168, 170; arguments during debate, 146-48, 150-51, 152, 153-54; post debate, 164, 167, 169; followers reinterpret Cuvier's views on, 204-5, 211 in work of other French naturalists: Blainville, 45, 72, 93, 94-95, 97-98, 120, 213-15; Savigny, 71-72, 93-94, 95, 106, 113, 115, 267n24, 285n8; Dumeril, 72, 95, 105, 115-16, 264n99; Dutrochet, 72, 95, 172; Latreille, 94, 106, 110, 112-14, 147, 267n24, 267n25; Audouin, 94, 106, 110, 114-18, 217; Milne Edwards, 94, 207, 216-19, 222, 274nl2, 287n46; Serres, 122-24, 171-72; Duges, 161-65; Isidore Geoffroy, 208-9, 264nl07, 264nl08. See also Homology; "Theory of analogues;" "Unity of organic composition" Philosophical botany, 93, 96, 159

INDEX

Phrenology, 48 "Physiological zoology." See Zoology Pinel, Philippe, 84, 92 Plato, 3, 170, 229, 230 Pluche, Noel-Antoine, 13, 56 Poisson, Simeon-Denis, 32 Politics of science: role in the debate, 8-9, 142, 173; and the debate, 173, 177, 193-201 Polyptems, 75, 83 Portal, Antoine, 17, 125 Positivism, 48, 254n35 Pouchet, Felix, 54 Preformation. See Generation Prevost, Constant, 58 Priestley, Joseph, 55-56 Principle of connections. See Connections, principle of Private enterprise in French science, 63 Professionalization: in French science, 37, 57, 204, 254n53, 256n71; Geoffroy's nonprofessional behavior in 1830s, 178; satire of esoteric terminology and, 200; historiographical debate over, 258n94 Progressivism, in Geoffroy's doctrine, 184, 185, 189-90 Publishing contracts as support for naturalists, 63 Quatrefages, Armand de, 222, 233, 234, 236 Quinet, Edgar, 176, 188, 192-93, 236 Quoy, Rene-Constant, 65, 67, 259nl04, 280nl4 Raffeneau-Delile, Alire, 73 Raspail, Francois-Vincent: and Geoffroy and the debate, 176, 177, 196, 197-98, 236; on politics of French science, 193, 196-98, 199, 236, 284nl03; career, 195-96 Rathke, Martin Heinrich, 206, 285n8, 285nlO Ray, John, 56 Reaumur, Rene-Antoine Ferchaultde, 13, 56 Recapitulation: Geoffroy and, 107, 205; Serres and, 107, 123-24; transcendental versus evolutionary form, 124; and branching conception of development, 205-6, 21920, 228. See also Arrests of development Redoute, Henri-Joseph, 73 Redoute, Pierre-Joseph, 73 Reductionism, 50, 183 Reichert, Karl Bogislaus, 206-7 Religion: and science in France and Britain, 56-58, 141; role in Cuvier-Geoffroy controversy, 7, 137-38, 140-41, 154, 155,

303

170, 173; and interpretation of the debate, 188-92, 194. See also Blainville, G. Cuvier; E. Geoffroy Repetition of parts, 162, 229 Respiration, Geoffroy on, 77, 128-29, 135-36 Revolution of 1830: and the debate, 1, 143-45, 156, 157, 158, 173; Cuvier and Geoffroy's activities during, 166 Revue encyclopedique, 103, 156, 158, 159 Reynaud, Jean, 186, 188 Reynier, J.-L.-E., 76 Richard, Claude, 31, 39 Robinet, Jean-Baptiste Rene, 14 Roulin, Francois-Desire, 133, 234 Rousseau, Jean Jacques, 54 Royer, Clemence, 233 Rudimentary organs: in work of Geoffroy, 74, 76, 83, 99, 101; Cuvier and, 101, 203; have meaning as showing plan, 225; Darwin reinterprets, 231, 232 Ruse, Michael, 224 Russell, Edward Stuart, 3, 5, 235, 237, 246n6 Saigey, J.-F., 196 Saint-Hilaire, Auguste de, 96, 179, 180 Salles, Bertrand Isidore de, 177, 198 Sand, George, 58, 176, 177, 188-90, 192 Savigny, Jules-Cesar: and philosophical anatomy, 71-72, 93-94, 95, 106, 113, 115, 267n24, 285n8; in Egypt, 72, 73, 74, 93, 263n90; career, 94; Geoffroy's relations with, 263n90 Schelling, Frederick Wilhelm Joseph, 90 Secondary schools (France), 44, 53, 61-62 Series, animal. See Classification Serres, Etienne Reynaud Augustin: career, 121-22, 125; and Cuvier, 121-22, 124-25, 168; as disciple of Geoffroy, 122, 124, 125, 171-72, 192, 202; tries to moderate Geoffroy, 180; Milne Edwards and, 217, 219, 220; personality, 270n79 scientific doctrine: embryology, 106, 110, 122-24, 128, 142, 153, 167, 171-72; arrests of development and recapitulation, 107, 123-24, 167, 171-72; philosophical anatomy, 122-24, 171-72; methodology, 125, 142; evolution, 133, 136 Silence, conspiracy of, as strategy, 68, 112, 179, 196, 233 Silurus, 75, 83 Societe d'Histoire Naturelle, 15-16, 21, 27, 30, 33, 38 Societe d'Histoire Naturelle (1822), 196 Societe Linneene, 15 Societe Philomatique, 21, 36, 38, 97, 98, 161

304

INDEX

Soi pour soi, Geoffroy's law of: and teratology, 128, 129; in writings of 1830s, 178, 179, 181-83; response to, 176, 179, 188-89, 192; Isidore Geoffroy restricts applicability of, 205, 209 Sonnerat, Pierre, 14, 27 Soret, Frederic, 1, 158-61 passim Sound, Geoffroy's theory of, 100-101, 265n119 Spaendonck, Gerard van, 17 Specialization in sciences: encouraged after founding of Museum, 24, 37-38; Geoffroy's rejection of, 24, 26, 78, 81, 181, 183, 191; protest of outsiders against, 177, 181, 191, 199 Spix, Johann Baptist von, 106, 107, 158 Spontaneous generation. See Generation Stadholder of Holland, 35 Steneosaums, 130, 132, 134, 166 Sternum, homologies of in vertebrates: Geoffroy on, 87, 100, 265nl 17; Cuvier on, 101-2, 153-54, 167 "Subordination of characters," 15, 33, 34, 219 Suppleants, 31 Swedenborg, Emanuel, 190 Temps, Le, 158 Tenon, Jacques Rene, 91, 92 Teleology: rejection of sufficiency of teleological explanation, 205, 207, 227-28; significance of transformation in understanding of purpose, 222-23, 22930; laws of organization and, 223, 224, 225; Darwin only gradually rejects, 23031 and morphology: synthesis of in Britain, 205, 222-30, 231; in development of Darwin's theory, 207, 230-32; morphology versus teleology derives from British context, 205, 223, 224; reconciliation more explicit in Britain than in France, 205, 214, 222-23 See also Functionalism; Morphology; Natural theology Teleosaurus, 130, 131, 132, 134-35 Teratology: Isidore Geoffroy and, 126, 208; Serres and, 124, 125, 142; as threat to Cuvier, 129-30 Geoffroy and: writings in the 1820s, 106, 124, 125-30, 139, 178, 182, 270n85; experiments, 128-29, 132, 271nl03; and mechanism of evolution, 131, 132, 179,

184; writings in the 1830s, 156, 178-82 passim Tessier, Henri-Alexandre, 29, 30 Tetraodon, 75, 86 Thenard, Louis Jacques, 114 "Theory of analogues," 157-58, 197; Geoffroy and, to 1818, 71, 85, 98, 103, 109; Cuvier opposes, 140, 154; Geoffroy and, in 1820s and 1830s, 149, 151, 153, 167, 187; Isidore Geoffroy distinguishes from unity of composition, 209, 264nl08. See also "Unity of organic composition" Thouin, Andre, 17 Tiedemann, Friedrich, 158 "Transcendental anatomy," 120, 121, 122, 123, 172 Tressan, abbe de, 19 Turpin, Pierre-Jean-Francois, 96, 180 "Unity of organic composition" Geoffroy and: early speculations on unity of plan, 12, 23-24, 28-29; doctrine of, 71, 85, 100, 103, 110, 126, 135, 146, 259n6; and evolution, 130, 131, 135, 136, 138, 153, 184; defense of, 140, 141, 149, 163; shortcomings of doctrine, 203 Cuvier and: rejection of, 137-38, 140, 148, 150-54 passim, 164, 168, 170; embryological arguments against, 167; sees as chief issue in debate, 172 in work of others: Buffon and Daubenton and unity of plan, 23-24; Audouin, 114, 117; Serres, 123; Guerin, 156-57; Duges, 162; Balzac, 191-92, 194-95; Isidore Geoffroy, 209, 264nl08; Flourens, 211, 233-34; Blainville, 213; Milne Edwards, 217, 221, 287n46; in Britain, 224, 225, 228, 230, 231; Darwin associated with, 233-34 See also Homology; Philosophical anatomy; "Theory of analogues" Unity of plan. See Philosophical anatomy; "Unity of organic composition" University of France, 53, 61-62. See also Faculte des Sciences (Paris); Ecole Normale(1808) Valenciennes, Achille, 60, 62, 65-66, 200, 202 Vauquelin, Nicolas Louis, 284nl03 Vernois, Maxime, 279n5 Vertebral theory of the skull: Dumeril on, 95, 105, 115-16; Audouin on (in articulates),

INDEX

106, 110, 115-16; in German biology, 106, 107, 115, 159; Geoffroy on, 107, 109, 110; Cuvier on, 115-16; in Britain, 229, 290nl07 Vicq d'Azyr, Felix, 14, 61, 213, 214; and comparative anatomy, 14, 31, 67; and homologies, 70, 87, 90 Vienot, John, 55 Virey, J.-J., 277n87 Vitalism, 103-4, 120, 221; Cuvier on, 50; Geoffroy on, 79, 81, 167, 182, 277n87 Volta, Alessandro, 168 Voltaire, Francois Marie Arouet de, 54

305

Weber, Ernst Heinrich, 109, 266nl 1 Whewell, William, 144, 223-24, 225, 230 Woodward, John, 49 Wyman, Jeffries, 232, 270n81 Zoology: in France before founding of the Museum, 11, 13-16, 21; need to make more philosophical, 12, 30, 32; institutions in France related to, 34-37, 59-65, 238-44; Milne Edwards and "physiological zoology," 215-22 Zoonite, 162