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Discovering Cell Mechanisms
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  • Page extent: 334 pages
  • Size: 228 x 152 mm
  • Weight: 0.575 kg

Library of Congress

  • Dewey number: 571.6
  • Dewey version: 22
  • LC Classification: QH581.2 .B42 2006
  • LC Subject headings:
    • Cytology
    • Cytology--history
    • Cell Physiology
    • Cytological Techniques

Library of Congress Record


 (ISBN-13: 9780521812474 | ISBN-10: 052181247X)

Discovering Cell Mechanisms

The Creation of Modern Cell Biology

Between 1940 and 1970, pioneers in the new field of cell biology discovered the operative parts of cells and their contributions to cell life. They offered mechanistic accounts that explained cellular phenomena by identifying the relevant parts of cells, the biochemical operations they performed, and the way in which these parts and operations were organized to accomplish important functions. Cell biology was a revolutionary science in its own right, but in this book, it also provides fuel for yet another revolution, one that focuses on the very conception of science itself. Laws have traditionally been regarded as the primary vehicle of explanation, but in the emerging philosophy of science it is mechanisms that do the explanatory work. William Bechtel emphasizes how mechanisms were discovered by cell biologists, focusing especially on the way in which new instruments – the ultracentrifuge and the electron microscope – made these inquiries possible. He also describes how scientists organized new journals and professional societies to provide an institutional structure to the new enterprise.

William Bechtel is professor of philosophy and science studies at the University of California, San Diego. He is the author and editor of many books, including Discovering Complexity (with Robert C. Richardson, 1993) and Connectionism and the Mind (with Adele Abrahamsen, 2002), and is editor of the journal Philosophical Psychology. He is past president of the Society for Philosophy and Psychology and the Southern Society for Philosophy and Psychology.


General Editor
Michael Ruse Florida State University

Advisory Board
Michael Donoghue Yale University
Jean Gayon University of Paris
Jonathan Hodge University of Leeds
Jane Maienschein Arizona State University
Jesús Mosterín Instituto de Filosofía (Spanish Research Council)
Elliott Sober University of Wisconsin

Alfred I. Tauber The Immune Self: Theory or Metaphor?
Elliott Sober From a Biological Point of View
Robert Brandon Concepts and Methods in Evolutionary Biology
Peter Godfrey-Smith Complexity and the Function of Mind in Nature
William A. Rottschaefer The Biology and Psychology of Moral Agency
Sahotra Sarkar Genetics and Reductionism
Jean Gayon Darwinism’s Struggle for Survival
Jane Maienschein and Michael Ruse (eds.) Biology and the Foundation
of Ethics
Jack Wilson Biological Individuality
Richard Creath and Jane Maienschein (eds.) Biology and Epistemology
Alexander Rosenberg Darwinism in Philosophy, Social Science, and Policy
Peter Beurton, Raphael Falk, and Hans-Jörg Rheinberger (eds.)
The Concept of the Gene in Development and Evolution
David Hull Science and Selection
James G. Lennox Aristotle’s Philosophy of Biology
Marc Ereshefsky The Poverty of the Linnaean Hierarchy
Kim Sterelny The Evolution of Agency and Other Essays
William S. Cooper The Evolution of Reason
Peter McLaughlin What Functions Explain
Steven Hecht Orzack and Elliott Sober (eds.) Adaptationism and Optimality
Bryan G. Norton Searching for Sustainability
Sandra D. Mitchell Biological Complexity and Integrative Pluralism
Joseph LaPorte Natural Kinds and Conceptual Change
Greg Cooper The Science of the Struggle for Existence
Jason Scott Robert Embryology, Epigenesis, and Evolution
William F. Harms Information and Meaning in Evolutionary Processes

Discovering Cell

The Creation of Modern Cell Biology

University of California, San Diego

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press
40 West 20th Street, New York, NY 10011-4211, USA
Information on this title:

© William Bechtel 2006

This publication is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without
the written permission of Cambridge University Press.

First published 2006

Printed in the United States of America

A catalog record for this publication is available from the British Library.

Library of Congress Cataloging in Publication Data

Bechtel, William.
Discovering cell mechanisms : the creation of modern cell biology / William Bechtel.
p. cm. – (Cambridge studies in philosophy and biology)
Includes bibliographical references and index.
ISBN-13: 978-0-521-81247-4 (hardback)
ISBN-10: 0-521-81247-X (hardback)
1. Cytology. I. Title. II. Series.
[DNLM: 1. Cytology–history. 2. Cell Physiology. 3. Cytological Techniques. QU11.1B391d 2006]
QH581.2.B42    2006
571.6 – dc22    2005017960

ISBN-13 978-0-521-81247-4 hardback
ISBN-10 0-521-81247-X hardback

Cambridge University Press has no responsibility for
the persistence or accuracy of URLs for external or
third-party Internet Web sites referred to in this publication
and does not guarantee that any content on such
Web sites is, or will remain, accurate or appropriate.


Preface page xi
1.   Introduction: Cell Mechanisms and Cell Biology 1
    1. A Different Kind of Science 1
    2. The Organization of Science into Disciplines 5
    3. The New Discipline of Cell Biology 13
2.   Explaining Cellular Phenomena through Mechanisms 19
    1. Historical Conceptions of Mechanism 20
    2. Twentieth-Century Conceptions of Mechanism 24
    3. Current Conceptions of Mechanism 26
      Mechanisms Explain Phenomena 27
      Component Parts and Component Operations 30
      Organization and Orchestration 32
    4. Representing and Reasoning about Mechanisms 33
    5. Levels of Organization and Reduction 40
    6. Organization: From Cartesian to Biological Mechanisms 44
    7. Discovering and Testing Models of Mechanisms 54
      Identifying Working Parts 55
      Identifying Component Operations 57
      Localizing Operations in Parts 60
      Testing Models of Mechanisms 61
    8. Conclusion 62
3.   The Locus of Cell Mechanisms: Terra Incognita between Cytology and Biochemistry 64
    1. Cytological Contributions to Discovering Cell Mechanisms up to 1940 65
      Cytology in the Nineteenth Century 68
      Cell Membranes (1825–1935) 77
      Mitochondria (1890–1925) 80
      Ergastoplasm or Basophilia (1900–1930) 83
      The Golgi Apparatus (1900–1940) 84
      The State of Cytology circa 1940 88
    2. Biochemical Contributions to Discovering Cell Mechanisms up to 1940 89
      Foundations for Biochemistry in the Nineteenth Century 89
      The Emergence of Biochemistry in the Twentieth Century 94
      Alcoholic and Lactic Acid Fermentation (1895–1940) 97
      Aerobic Cellular Respiration (1910–1940) 105
      The State of Biochemistry circa 1940 116
    3. The Need to Enter the Terra Incognita between Cytology and Biochemistry 117
4.   Creating New Instruments and Research Techniques for Discovering Cell Mechanisms 118
    1. The Epistemology of Evidence: Judging Artifacts 121
    2. The Ultracentrifuge and Cell Fractionation 128
      Breaking Cell Membranes 131
      Choice of Media 133
      Centrifugation Regimes 135
      Interpreting Fractionation Results 137
    3. The Electron Microscope and Electron Microsopy 140
      Obtaining Sufficiently Thin Specimens 143
      Altering the Specimen to Survive Microscopy and Generate an Image 149
    4. A Case Study of an Artifact Charge 157
    5. Equipped with New Instruments and Techniques to Enter Terra Incognita 160
5.   Entering the Terra Incognita between Biochemistry and Cytology: Putting New Research Tools to Work in the 1940s 162
    1. First Steps toward Cell Biology at the Rockefeller Institute: Claude’s Introduction of Cell Fractionation 163
    2. Robert Bensley: An Alternative Approach to Fractionation 166
    3. Competing Interpretations of Fractions from Normal Cells 168
    4. Linking Claude’s Microsomes to Protein Synthesis 171
      Brachet: Selective Staining of RNA and Correlation with Protein Synthesis 171
      Caspersson: Spectrographic Analysis, RNA, and Protein Synthesis 173
    5. Adding a Biochemical Perspective to the Rockefeller Laboratory 177
    6. Adding Electron Microscopy as a Tool 182
    7. The State of Cell Studies at the End of the 1940s 188
6.   New Knowledge: The Mechanisms of the Cytoplasm 190
    1. The Mitochondrion 192
      Biochemists Confront Particulate Structure: Mitochondrial Enzyme Systems 192
      More Structure: The Discovery of the Cristae of the Mitochondrion 199
      A Competing Perspective on Mitochondrial Morphology 201
      Biochemists Further Fractionate Mitochondria 209
      One More Piece of Structure and a Proposal as to Its Function 215
      Radical Reconceptualization of Oxidative Metabolism 219
    2. Microsomes, the Endoplasmic Reticulum, and Ribosomes 222
      From Lace–like Reticulum to Endoplasmic Reticulum 222
      Dissenting Voices 228
      Securing the Connection to Protein Synthesis 231
      Integrating Morphology and Biochemistry 233
      Naming the Ribosome 236
      Going to a Lower Level: Decomposing the RNA Machinery 237
      Transporting Newly Sequenced Polypeptides 243
    3. Two Additional Organelles 243
      The Golgi Apparatus 244
      The Lysosome 250
    4. Conclusion 257
7.   Giving Cell Biology an Institutional Identity 258
    1. Creation of the Journal of Biophysical and Biochemical Cytology 260
    2. Creation of the American Society for Cell Biology 268
    3. Conclusion 276
Afterword 279
References 281
Index 313


This book is the product of research spanning two decades. In the 1980s I had been investigating the history of cytology in the nineteenth century and biochemistry in the early twentieth century when I responded to an announcement from the American Society for Cell Biology of a fellowship for support of research on the history of cell biology. With their financial support in 1986 and 1987 I began to examine the creation of modern cell biology in the decades after World War II. I am enormously grateful not only for the fellowship funding but also for the invaluable assistance of individuals associated with ASCB. In particular, I thank Robert Trelstad, then Secretary of ASCB, who invited me to society and executive council meetings, gave me encouragement, and provided entrée to senior members of the society. A number of the founders of modern cell biology were still active in the society and I had the opportunity to meet and interview them regarding their own contributions to cell biology and their recollections of the early days of this field. I also had access to the archives of the society, which were then housed at the Society offices. (They have since been transferred to the University of Maryland, Baltimore County.) I have relied heavily on this material in analyzing in Chapter 7 the history of the American Society for Cell Biology.

   In the early 1990s I received additional support from the National Endowment for the Humanities. I am most appreciative of the support from NEH as well as from the National Science Foundation, which funded my earlier work on the history of biochemistry. Among other activities, this support enabled me to interview many additional pioneer cell biologists.

   I particularly wish to thank the following scientists for taking time to meet with me and provide their reflections and insights on the history of cell biology: Max Alfert, Vincent Allfrey, Helmut Beinert, Britton Chance, Christian de Duve, Morgan Harris, Daniel Mazia, Montrose Moses, George Palade, George Pappas, Keith Porter, Van Potter, Hans Ris, Birgit Satir, Peter Satir, Philip Siekevitz, Paul Stumpf, and Hewson Swift. I am also deeply appreciative for the guidance Pamela Henson provided me on the techniques for doing aural history with scientists.

   The NEH grant also enabled me to carry out research at the Rockefeller Archive Center (RAC). This center holds records of laboratories at the Rockefeller Institute for Medical Research, as well as the archives of the Rockefeller Foundation, which provided funding for many of the early cell biologists. Especially useful for Chapters 5 and 6 were the Annual Reports submitted each April by every laboratory at the Rockefeller Institute. These were incorporated into the Scientific Reports of the Laboratories to the Board of Scientific Directors (RG 439, Rockefeller University Archives, RAC). I will refer to these reports throughout the text as simply the Annual Report for a laboratory.

   While working on this project I have been associated with the philosophy departments of three universities: Georgia State University, Washington University in St. Louis, and the University of California, San Diego. Each has provided invaluable support for which I am most grateful. I particularly appreciate the contributions of several of my graduate students. In particular, at Washington University Jennifer Mundale transcribed many of the oral interviews and engaged in numerous fruitful discussions with me about this and related projects. More recently, at UCSD I have benefited from the ideas and insights of Andrew Hamilton, who provided detailed comments on Chapter 2, and of Cory Wright. I have also benefited from interactions with graduate students who participated in my seminars on mechanism at both Washington University (presented jointly with Carl Craver) and UCSD.

   My spouse and frequent collaborator, Adele Abrahamsen, has made an enormous contribution to this project. I thank her for extremely valuable comments on the entire manuscript. I have benefited from our many productive discussions of mechanism and the history of research on cells. As well, various parts of the text draw upon papers we coauthored.

   A key component of my analysis is that the knowledge developed in cell biology consisted of the discovery of various cell mechanisms. My understanding of what a mechanism is and how scientists investigate them has benefited from many discussions with Robert Richardson, my coauthor on Discovering Complexity, as well as Carl Craver, Lindley Darden, Stuart Glennan, and William Wimsatt.

   Finally, I thank Michael Ruse, who initially invited me to submit a proposal for this book to Cambridge University Press and then showed admirable patience while I produced it.

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