Table Of ContentDevelopmental
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
