Foundations of the Neuron Doctrine: 25th Anniversary Edition

385 Pages·2015·31.08 MB·English

by  Gordon M Shepherd

Foundations of the
Neuron Doctrine

Foundations of the
Neuron Doctrine
25th Anniversary Edition
Gordon M. Shepherd
Professor of Neurobiology
Yale University School of Medicine
1
1
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Library of Congress Cataloging-in-Publication Data
Shepherd, Gordon M., 1933–, author.
Foundations of the neuron doctrine / Gordon M. Shepherd.—25th anniversary edition.
p. ; cm.
Includes bibliographical references and index.
ISBN 978–0–19–025938–9 (alk. paper)
I. Title.
[DNLM: 1. Ramón y Cajal, Santiago, 1852–1934. 2. Neurons. 3. History, 19th
Century. 4. Neuroanatomy—history. WL 102.5]
QM451
611ʹ.809—dc23
2015016485
9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid-free paper
For Grethe

Contents
Preface to the 25th Anniversary Edition  ix
Preface to the Original Publication  xiii
Commentaries on the “Neuron Doctrine”  xvii
Cajal, Golgi, and Ariadne’s Thread—Marina Bentivoglio  xvii
Reflections on the Neuron Doctrine—Javier DeFelipe  xx
The Neuron Doctrine Revisited: A Personal Account—Sten Grillner  xxiv
Camillo Golgi, Foundations of the Neuron Doctrine, and the History
of Neuroscience—Paolo Mazzarello  xxviii
Some Reflections on the Neuron Doctrine—Larry Swanson  xxxi
Back to Golgi? Neural Networks as a New Paradigm for Brain
Circuits—Rafael Yuste  xxxiii
1. Introduction  1
2. From the Beginnings to the Cell Theory  9
3. Do Nerve Cells Belong in the Cell Theory?  19
4. Nerve Cells or Nerve Nets?  33
5. KÖlliker Gives In  43
6. Support Builds for Networks  51
7. The Nerve Cell Studies of Freud  65
8. The Revolutionary Method of Golgi  77
9. A Neuron Theory Begins to Take Form: His, Forel, Nansen  101
10. Ramón y Cajal: The Shock of Recognition  127
11. The Early Discoveries of Cajal  139
12. The Laws of Cajal  155
13. Joining the Mainstream  167
14. The Neuron Doctrine  179
vii
15. The Law of Dynamic Polarization  199
16. Controversy  217
17. The Synapse and the Growth Cone  231
18. Forging a Consensus  243
19. Confrontation in Stockholm  267
20. Modern Revisions of the Neuron Doctrine  279
Comment on Sources  297
References  299
Index  311
viii Contents
Preface to the 25th Anniversary Edition
This account of the work that established the neuron doctrine appears to have stood
the test of time over the past quarter century. In addition to emphasizing the key role
of Santiago Ramon y Cajal, and providing translations of a number of papers by the
classical histologists, I was particularly interested in rehabilitating the reputation of
Camillo Golgi, from being a loser in the fight to establish the neuron doctrine to being
a winner in not only inventing the tool to do it but also a pioneer in conceiving of the
network function of the nervous system. This is both with regard to the gap junctions
that provide continuity between cells, but also the way that workers in neural systems
and artificial intelligence think of the nervous system as a network of interconnected
nodes. I also hoped that this in-depth account would place the work on the neuron
doctrine at the highest level of intellectual history and in its rightful place among the
greatest scientific achievements of all time. The book has helped make progress toward
these goals.
In the 25 years since the first edition, several important events for the neuron doc-
trine have stood out for me. First was the English translation in 1994 by Neely and
Larry Swanson of Cajal’s great work, Histologie du système nerveux de l’homme et des
vertébrés. Now every neuroscientist can appreciate the studies that founded modern
cellular neuroscience and that remain relevant today.
Second was the biography of Golgi by Paolo Mazzarello, published in Italian as
La Struttura Nascosta in 1996 and in English as The Hidden Structure. A Scientific
Biography of Camillo Golgi in 1999, and Golgi. A Biography of the Founder of Modern
Neuroscience in 2010. I was gratified to be asked to write a foreword in which I could
emphasize that this elegant study of Golgi’s life and works enabled him for the first
time to be appreciated for his attractive personal qualities, the great scientific tradition
in which he worked, his deep commitment to the experimental method in biology, and
his other key discoveries. As noted by Mazzarello, the association of Golgi’s name with
the Golgi apparatus alone makes him “probably the most cited biologist in scientific
literature” (see Fabene and Bentivoglio, 1998, for the numbers). These insights into
Golgi can now be compared with Cajal’s fascinating accounts of his own life to give a
much richer understanding of both men.
A third event was the centenary in 2006 of the awarding of the Nobel Prize in 1906 to
Cajal and Golgi for their discoveries of nerve cells. The Cajal Club organized a meeting
ix