A Conceptual History of Modern Embryology PDF
Preview A Conceptual History of Modern Embryology
Developmental Biology A COMPREHENSIVE SYNTHESIS Volume 7 A Conceptual History of Modern Embryology Developmental Biology A COMPREHENSIVE SYNTHESIS Editor LEON W. BROWDER University of Calgary Calgary, Alberta, Canada Editorial Board EVERETT ANDERSON ELIZABETH D. HAY Harvard Medical School Harvard Medical School JOSEPH T. BAGNARA RALPH S. QUATRANO The University of Arizona The DuPont Company SAMUEL H. BARONDES RUDOLF A. RAFF University of California at San Francisco Indiana University ANTONIE W. BLACKLER L. DENNIS SMITH Cornell University University of California at Irvine MARIE A. DiBERARDlNO IAN M. SUSSEX The Medical College of Pennsylvania Yale University RALPH B. L. GWATKIN The Cleveland Clinic Foundation Volume 1 OOGENESIS Edited by Leon W. Browder Volume 2 THE CELLULAR BASIS OF MORPHOGENESIS Edited by Leon W. Browder Volume 3 THE CELL SURFACE IN DEVELOPMENT AND CANCER Edited by Malcolm S. Steinberg Volume 4 MANIPULATION OF MAMMALIAN DEVELOPMENT Edited by Ralph B. 1. Gwatkin Volume 5 THE MOLECULAR BIOLOGY OF CELL DETERMINATION AND CELL DIFFERENTIATION Edited by Leon W. Browder Volume 6 GENOMIC ADAPTABILITY IN SOMATIC CELL SPECIALIZATION Edited by Marie A. DiBerardino and Laurence D. Etkin Volume 7 A CONCEPTUAL HISTORY OF MODERN EMBRYOLOGY Edited by Scott F. Gilbert Developmental Biology A COMPREHENSIVE SYNTHESIS Volume 7 A Conceptual History of Modern Embryology Edited by SCOTT F. GILBERT Swarthmore College Swarthmore, Pennsylvania PLENUM PRESS • NEW YORK AND LONDON Library of Congress Cataloging in Publication Data (Revised for volume 7) Developmental biology. Vol. 4 edited by Ralph B. L. Gwatkin. Vol. 6 edited by Marie A. DiBerardino and Laurence D. Etkin. Vol. 7 edited by Scott F. Gilbert. Includes bibliographies and indexes. Contents: v. 1. Oogenesis-[etc.]-v. 6. Genomic adaptability in somatic cell specialization - v. 7. A conceptual history of modern embryology. 1. Developmental biology. I. Browder, Leon W. QH491.D426 1985 574.3 85-3406 Cover illustration: Spemann's hairloop experiment. From Experimentelle Beitriige zu einer Theorie der Entwicklung (1936). ISBN-13: 978-1-4615-6825-4 e-ISBN-13: 978-1-4615-6823-0 DOl: 10.1007/978-1-4615-6823-0 © 1991 Plenum Press, New York Softcover reprint ofthe hardcover 1st edition 1991 A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013 All rights reserved- No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Contributors P. G. Abir-Am Department of History of Science, Johns Hopkins University, Baltimore, Maryland 21218 Richard M. Burian Department of Philosophy, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061-0126 Frederick B. Churchill Department of the History and Philosophy of Science, Indiana University, Bloomington, Indiana 47405 Jean-Louis Fischer Centre de Recherche d'Histoire des Sciences, Universite de Paris, Paris, France Jean Gayon Faculte de Lettres et Philosophie, Universite de Bourgogne, 2100 Dijon, France Scott F. Gilbert Department of Biology, Swarthmore College, Swarthmore, Pennsylvania 19081 Gerald B. Grunwald Department of Anatomy, Jefferson Medical College, Thomas Jefferson University, Philadelphia, Pennsylvania 19107 Johannes Holtfreter Department of Biology, University of Rochester, Rochester, New York 14627 Jane Maienschein Department of Philosophy, Arizona State University, Tempe, Arizona 85287 Jane M. Oppenheimer Department of Biology, Bryn Mawr College, Bryn Mawr, Pennsylvania, 19010-2899; present address: One Independence Place, 6th Street and Locust Walk, Philadelphia, Pennsylvania 19106 Margaret Saha Department of Biology, University of Virginia, Charlottesville, Virginia 22901 Jan Sapp Department of the History and Philosophy of Science, University of Melbourne, Parkville, Victoria, Australia Doris T. Zallen Center for the Study of Science in Society and Center for Programs in the Humanities, Virginia Polytechnic Institute and State Univer sity, Blacksburg, Virginia 24061-0227 v Preface "Glory to the science of embryology!" So Johannes Holtfreter closed his letter to this editor when he granted permission to publish his article in this volume. And glory there is: glory in the phenomenon of animals developing their complex morphologies from fertilized eggs, and glory in the efforts of a relatively small group of scientists to understand these wonderful events. Embryology is unique among the biological disciplines, for it denies the hegemony of the adult and sees value (indeed, more value) in the stages that lead up to the fully developed organism. It seeks the origin, and not merely the maintenance, of the body. And if embryology is the study of the embryo as seen over time, the history of embryology is a second-order derivative, seeing how the study of embryos changes over time. As Jane Oppenheimer pointed out, "Sci ence, like life itself, indeed like history, itself, is a historical phenomenon. It can build itself only out of its past." Thus, there are several ways in which embryology and the history of embryology are similar. Each takes a current stage of a developing entity and seeks to explain the paths that brought it to its present condition. Indeed, embryology used to be called Entwicklungsgeschichte, the developmental history of the organism. Both embryology and its history interpret the interplay between internal factors and external agents in the causation of new processes and events. The embryologist, of course, has the advantage of seeing this "history" repeating itself every time a new organism is generated. So it is not surprising that historians, embryologists, embryologists-turned historians, and historians-turned-embryologists can collaborate on a history of embryology. This is, of course, a history of embryology, and there is no pretense that it is the history of the field (any more than the excellent volumes on oogenesis and morphogenesis in this series can be considered the complete texts in these areas). The predominant theme in this volume is the concept of induc tion. This was not the way the book was originally planned, but rather, a fortu itous accident of those who were able to write their chapters during this time. Other themes could have predominated, and I hope that there will be more historical volumes in this series that will address these areas. However, induc tion has certainly been a pivotal principle in the history of embryology, and it remains an extremely active field of contemporary research. Given the current interest in the molecular mechanisms of neural and "secondary" induction, this history becomes all the more timely. One of the values of this book for contemporary developmental biologists vii viii Preface should be to clarify some of the concepts that embryologists have bequeathed to developmental biology. The volume begins with a chapter by Frederick Churchill that details the bases for the tradition of comparative embryology. The work of Baer, Pander, and Rathke is central to this endeavor, and it is here that the first observations of mutually interactive tissues were made. When Christian Pander discovered the primary germ layers of the chick embryo, he concluded that a unique metamorphosis begins in each of these three [germ layers] and hurries toward its goal; although each is not yet independent enough to indicate what it truly is; it still needs the help of its sister travellers, and therefore although already designated for different ends, all three influence each other collectively until each has reached an appropriate level. This quotation shows that both internal and inductive features of embryo genesis were recognized very early. Jean-Louis Fischer then provides the context of French comparative anatomy and teratology in which Laurent Chabry per formed his experiments showing the mosaic development in tunic ate embryos. Fischer also shows how personal and national ideologies can influence the interpretation of experimental results. Next, Jane Maienschein provides a history of the early work in Entwicklungsmechanik, belying the textbook notion of a few "founding fathers" and demonstrating the rich tapestry provided by the work of numerous investigators. One of these investigators, Curt Herbst, is given special attention in the chapter by Jane Oppenheimer. Her study delineates how calcium and lithium ions were first used to study morphogenesis, how the first concepts of induction entered into experimental embryology, and how physiological theories of tropisms influenced research into embryonic cell movements. The work of Herbst was extremely important in the intellectual development of Hans Spemann. Margaret Saha documents' the evolution of Spemann's concep tualization of induction by discussing his experiments on lens formation. These studies were critical for the design of Spemann's later experiments and provide insight into how he came to interpret them as he did. The next chapter is an autobiographical essay by Johannes Holtfreter in which he discusses his science and his art. He describes how he came to Spemann's and Mangold's laboratories, how his concept of induction changed over the years, and how he attempted to find the molecules that were responsible for the induction of the neural tube. Several research programs came out of the studies on induction and mor phogenesis that were performed by Spemann and Holtfreter. One of these was the study of the cell surface in development. The search for the processes and molecules responsible for intercellular adhesion is shown by Gerald Grunwald to have been full of surprising results and serendipitous observations. Another research program starting from the Freiburg group was Joseph Needham's at tempt to produce a biochemistry of the embryo. P. G. Abir-Am documents Need ham's attempts to place the biochemistry of the embryo on a solid theoretical and philosophical foundation. Here we see the complexities of forming a new inter disciplinary science and the need to free biological thinking from the paradigms of nineteenth-century physics. Needham's colleague in these endeavors, Conrad Waddington, attempted to weld Entwicklungsmechanik with genetics and evolutionary theory. Together with Salome Gluecksohn-Waelsch and Boris Ephrussi, he helped create the basic tenets of developmental genetics. Richard Burian, Jean Gayon, and Doris ix Preface Zallen discuss the ways in which Ephrussi attempted to synthesize genetics and development, while I relate the attempts of Gluecksohn-Schoenheimer and Waddington to use the concepts of induction and competence to reunite these disciplines. One of the most hotly debated topics in those early days of develop mental genetics was whether all the genotype resided in the nucleus. Both Ephrussi and Waddington were initially partial to the presence of plasmagenes in the cytoplasm. Jan Sapp details the debates on this issue, especially as they pertained to protozoans, organisms that some scientists saw as models for meta zoan development, but which other scientists saw as interesting exceptions to the general rule of nuclear inheritance. Developmental biology is the anagenetic descendant of embryology. N. J. Berrill relates that Paul Weiss wrote to him asking Berrill to suggest a name for the science that included embryology and also gene activity, regeneration, cell movement studies, and other areas of developmental biology. Berrill sent the letter back to him with the last two words capitalized. We who study develop mental biology are the inheritors of embryology'S concepts, organisms, and sense of wonder. From other sources, we have received a new set of tools with a resolving power far greater than what was available a generation ago. Frogs, chicks, and sea urchins (along with nematodes, flies, and leeches) are now being dissected with monoclonal antibodies, antisense mRNAs, and confocal micro scopes. We are presently seeing a return to those old embryological enigmas that were abandoned for lack of such specific tools. The morphogenesis of the disci ple continues. I thank all the authors for their contributions, and Marie DiBerardino, Leon Browder, and Mary Born for their roles in the conception and birth of this volume. "Glory to the science of embryology!" Scott F. Gilbert Contents Chapter 1 • The Rise of Classical Descriptive Embryology Frederick B. Churchill 1. Introduction ................................... '" . . .. . . .. . . . . 1 2. Descriptive Embryology in the Baltic Periphery. . . . .. . . .. . ... . . . . 2 3. Christian Pander .............................................. 3 4. Karl Ernst von Baer ........................................... 5 5. Heinrich Rathke .............................................. 12 6. A New Theoretical Framework at Mid-Century .................. 16 7. Evolution and Development.................................... 18 8. Specificity of the Germ Layers ................................. 20 9. Conclusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Notes and References ................... :. . . . . . . . . . . . . . . . . . . . . . 23 Chapter 2 • Laurent Chabry and the Beginnings of Experimental Embryology in France Jean-Louis Fischer 1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 2. E. Geoffroy Saint-Hilaire ....................................... 31 3. C. Dareste .................................................... 33 4. Stanislas Warynski and Hermann Fol ........................... 33 5. Laurent Chabry ............................................... 34 6. Edwin Grant Conklin, Albert Dalcq, and Oscar Hertwig .......... 39 7. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Notes and References. . . . . . . . . .. . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . 40 Chapter 3 • The Origins of Entwicklungsmechanik Jane Maienschein 1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 2. Foundations of Entwicklungsmechanik ......................... 43 3. Experimental Embryology...................................... 55 xi
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