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THE EVOLUTIONARY BIOLOGY PAPERS OF EUE METCHNIKOFF BOSTON STUDIES IN THE PHILOSOPHY OF SCIENCE Editors ROBERT S. COHEN, Boston University JURGEN RENN, Max-Planck-Institutefor the History of Science KOSTAS GAVROGLU, University ofA thens Editorial Advisory Board THOMAS F. GLICK, Boston University ADOLF GRUNBAUM, University of Pittsburgh SYLVAN S. SCHWEBER, Brandeis University JOHN 1. STACHEL, Boston University MARX W. WARTOFSKY t (Editor 1960-1997) VOLUME 212 THE EVOLUTIONARY BIOLOGY PAPERS OF ELIE METCHNIKOFF Edited by HELENA GOURKO Belorussian University & Boston University, U.S.A. DONALD I. WILLIAMSON Liverpool University, United Kingdom and ALFRED I. TAUBER Boston University, U.S.A. A C.I.P Catalogue record for this book is available from the Library of Congress. ISBN 978-90-481-5356-5 ISBN 978-94-015-9381-6 (eBook) DOI 10.1007/978-94-015-9381-6 Printed on acidjree paper AII Rights Reserved © 2000 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2000 Softcover reprint ofthe hardcover lst edition 2000 No part of this publication may be reproduced or utilized in any form or by any means, electronic, mechanical, including photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner. Dedicated in admiration and friendship to Robert S. Cohen and Annie Kuipers TABLE OF CONTENTS PREFACE IX INTRODUCTION I. ON THE DEVELOPMENTAL LIFE-HISTORY OF MYZOSTOMUM (1866) 22 Zur Entwicklungsgeschichte von Myzostomum. - Z. wiss. Zool., V.xVI, SS.236-243. 2. ON THE EMBRYONIC DEVELOPMENT OF LOWER CRUSTACEA (1866) 28 o pa3BlfTlUi HH3WHX paKoo6pa3HhIX B HHue. - HaTypaJlllCT, T 5, cTp.65-72. 3. ANTHROPOLOGY AND DARWINISM (1875) 36 Embryologisches uber Geophylis. - Z.wiss. Zoo!., v. XXV, SS. 313-322. AHTpOnOJIOflfH H .uapBHHlf3M - BecTH. EBponhI, KH. 1, CTp. 159-195. 4. ESSAY ON QUESTIONS ABOUT THE ORIGIN OF SPECIES (1876) 60 OqepK Bonpoca 0 npoHcxo)J(.ueHlflf BH.uOB. - BecTH. EBPOllhI, KH. 3,CTp. 68-134; KH.4,CTp. 715-747; KH. 5,CTp. 117-149; KH. 7, CTp. 158-197; KH. 8, CTp. 567-606. 5. COMPARATIVE EMBRYOLOGICAL STUDIES (1881-1885) 92 Vergleichhend-embryologische Studien. 1. Entodermbildung bei Geryoniden. 2. Ueber einige Stadien der Cunina. - Z. wiss. Zool., v. XXXVI, SS. 434-444. 3. Ueber die Gastrula einiger Metazoen. Z. wiss. Zoo!., - v. XXXVII, SS.286-313. 4. Ueber die Gastrulation und Mesodermbildung der Ctenophoren. Z. wiss. Zool., XLII, SS.648-55.5. Ueber die Bildung der Wanderzellen bei Asterien und Echinidien. - Z. wiss. Zoo!., v. XLII, SS.656-673. 6. EMBRYOLOGICAL STUDIES ON MEDUSAE: ON THE EVOLUTIONARY 144 ORIGINS OF THE PRIMARY TISSUES (1886) Embryologische Studien an Medusen. Ein Beitrag zur Genealogie der Primitiv-Organe, Wien, IV. 7. THE STRUGGLE FOR EXISTENCE BETWEEN PARTS OF THE ANIMAL ORGANISM (1892) 207 boph6a 3a CYIUeCTBaHHe qaCTeH )J(HBOTHOfO opaHH3Ma. - «llOMOIUhfOJIo.ualOIUHM. HayqHO-JlllTepaTYPHhIH C60PHHK». M., CTp. 321-326. [In French: Revue scient, v. L, pp. 321-326]. INDEX OF AUTHORS 217 INDEX OF ORGANISMS 221 vii PREFACE We are pleased to offer the English-speaking world the key evolutionary biology papers of Elie Metchnikoff (1845-1916), a work demanded by the rekindled interest in this key nineteenth century biologist. While hardly a household name today, Metchnikoff was a celebrated and well-known biologist, whose life has been well documented. He was a founding father of immunology, and became a popular figure in France in the first decade of this century through his phagocytosis theory - an immune theory applied broadly to issues of health and disease. With the Nobel Prize for Physiology or Medicine co-awarded to him and Paul Ehrlich (1908), Metchnikoff s prominence was assured. The publication by his wife, Olga, of The Life of Elie Metchnikoff (1919) shortly after his death has long served as the official rendition of his career. The account, at one level, is a simple offering with a minimum of psychological and scientific complexity. Written as a personal retrospective, the logical development of the research that led to Metchnikoff s immunological theory is defined, and the thread of its subsequent defense is given with self-righteous zeal. In this sense, the biography presents an insightful glimpse as how Metchnikoff viewed himself: a figure whose logical approach, masterful defense, and honest intelligence attempted to devour, very much like the phagocyte he championed, his scientific opponents and establish the primacy of his research. As such, the evolutionary biology in which he was immersed during the two decades preceding his studies in pathology and host immunity were relatively minimized. But the significance of this earlier research can hardly be over-emphasized in terms of its importance in formulating Metchnikoff s better known studies in natural immunity. With the recent publication of Daniel Todes's account of the historical context of Metchnikoffs research, Darwin without Malthus, The Struggle for Existence in Russian Evolutionary Thought (Oxford, 1989), and two intellectual biographies - the monograph, Metchnikoff and the Origins of Immunology. From Metaphor to Theory (Oxford 1991) by Alfred I. Tauber and Leon Chemyak, and the doctoral dissertation, "The wisdom of cells: The integrity of Elie Metchnikoffs ideas in biology and pathology" (Notre Dame University, 1991) by Craig Stillwell-students of nineteenth century Darwinism, developmental biology, immunology, pathology, and "socio biological" thinking now have ready access to critical introductions to a corpus of research that clearly articulated the biology of that era and helped formulate its agenda. Here we offer the most important primary sources in their first English translation. This translation project is the synergenic result between a translator/philosopher (Helena Gourko), a marine biologist (Don Williamson), and a historian/philosopher of science (Alfred Tauber). In our respective efforts, we have each profitted from the expertise of the other collaborators, and in framing this book, much has been appreciated heretofore ignored or unappreciated in Metchnikoffs opus. We can only hope that the reader also will profit from these diverse perspectives. Our translation is based on Metchnikoff s Russian edition (Academic Collection of Works / Akademicheskoe sobranie sochinenii / vols. I-XVII. Moscow, Academy of ix x PREFACE Medical Sciences of the USSR). As listed in the Table of Contents, some of these papers appear in the German and French literatures. Two editorial points: (1) Although Metchnikoffs style is complex -lucid, terse, and literary - the very density of the problems analyzed, as well as the significant difference between the expressions of sophisticated scientific and philosophical ideas in English and Russian, and more, the somewhat "archaic" nineteenth century Russian used by Metchnikoff make his writings difficult to translate. It is important to note that when Metchnikoff himself translated his works from Russian into other languages (he made a good many translations of this sort into German and French), he often transformed and changed them significantly. These changes did not alter the conceptual or descriptive details, but were obviously made for linguistic reasons. This immediately raises a major question concerning the degree of the translator's freedom, which, on the one hand, should be capable of keeping the content and context true to the original, but, on the other hand, must bring into consideration the significant inter-linguistic differences, as well as Metchnikoffs own approach in dealing with the problem. To resolve this difficulty fully is probably impossible, but it can be diminished by an understanding of Metchnikoff s major ideas and general orientation, which guided us in adjudicating any inconsistencies. (2) Significant difficulties were created by extracting sections from the extended text of the "Essay on the Question about the Origin of Species" (238 pp. in printed form). Because of the limited space for this translation, this text has been reduced by a careful selection of the most relevant passages that emphasize Metchnikoff s relationship to the Darwinian debate. Since Metchnikoffs views are not widely known, we have provided a concise introduction (based on Tauber and Chernyak, 1991), which presents the major ideas of Metchnikoff s views on evolutionary biology, as well as the results of his studies. The perspectives offered here concerning the development of Metchnikoff career and his underlying philosophy are, to be sure, still open to scholarly debate and represent a summary of a previously presented interpretation. Finally, without the encouragement of Robert S. Cohen and Annie Kuipers, this book would, in all likelihood, not have been completed. We thank them for their steadfast commitment to publish this translation. We, and the community of scholars at large, are in their debt. In appreciation of their friendship and support, we dedicate this volume. Helena Gourko Donald I. Williamson Alfred I. Tauber Boston University and The Isle of Man June, 1999 INTRODUCTION Why read Elie Metchnikoffs evolutionary biology papers? First, this research is of historical importance in its own right, reflecting a highly creative and critical opus of the latter half of nineteenth-century biology. Any consideration of this topic requires acquaintance with his thought. Second, although best known for his immunological theories, not his earlier work in embryology, it was Metchnikoff s foundational research in the problematics of Darwinian theory that drove his studies in pathology and natural host defense. Metchnikoff (1845-1916) was a celebrated scientist, known best as a key founder of modern immunology, but he played a crucial role during the formative period of evolutionary biology by first helping to establish comparative embryology as important support for Darwinism, and then applying his notions of the organism to the new field of infectious diseases. His ideas concerning development were extended to adult animals in what he called the "phagocytosis theory," which became a critical component of a new discipline, immunology. As a system of defense in the struggle of competing species, Metchnikoffs "phagocytosis theory" extended the particular clinical concerns with infection to a grand biological theory. Thus his contributions to zoology, comparative embryology, evolutionary biology, comparative pathology, medicine, immunology, anthropology, and social theory should be regarded as of one piece, each aspect of his work contributing to the expression of a wider philosophical understanding of biology. Therefore, Metchnikoff can only be understood by integrating his thought, and thus his immune theories must be appreciated in the context of his broader contributions to evolutionary biology, which explicitly or implicitly served to frame many critical areas of twentieth century medicine (Tauber and Chern yak 1989) and biology (Tauber and Chernyak, 1991; Tauber, 1991). Metchnikoff s primary scientific reports have remained largely untranslated into English. Of the many volumes of his published papers (seventeen in a definitive Russian edition), only a small fraction of these are available to the English-speaking reader. The best known translated works are his great summary texts, which are obviously valuable in serving as vehicles for his grand theories of biology and more general philosophical precepts, but these were primarily written as weapons in his polemics with other scientists. For instance, Immunity in Infectious Diseases (1901) is an elaborate argument for the centrality of Metchnikoff s phagocytosis theory against the competing views of German immunochemists. Such retrospective narratives of his work are inadequate records of Metchnikoff s research, the basis of his wider thinking, and the intimacy of revealing how a great nineteenth century zoologist regarded the major research issues of his time. In this regard, Metchnikoff s primary research is invaluable. We have assembled and translated Metchnikoffs key evolutionary biology papers dating from his earliest writings (1865) to those of his mature period of the 1890s,

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