HANS KREBS MONOGRAPHS ON THE HISTORY AND PHILOSOPHY OF BIOLOGY Editors RICHARD BURIAN, RICHARD BURKHARDT, JR. RICHARD LEWONTIN, JOHN MAYNARD SMITH THE CUVIER-GEOFFREY DEBATE French Biology in the Decades Before Darwin Toby A. Appel CONTROLLING LIFE Jacques Loeb and the Engineering Ideal in Biology Philip J.Pauly BEYOND THE GENE Cytoplasmic Inheritance and the Struggle for Authority in Genetics Jan Sapp THE HERITAGE OF EXPERIMENTAL EMBRYOLOGY Hans Spemann and the Organizer Viktor Hamburger THE EVOLUTIONARY DYNAMICS OF COMPLEX SYSTEMS A Study in Biosocial Complexity C. Dyke THK WELLBORN SCIENCE Eugenics in Germany, France, Brazil, and Russia Edited by Mark B. Adams DARWIN WITHOUT MALTHUS The Struggle for Existence in Russian Evolutionary Thought Daniel P. Todcs MHTCHNIKOFF AND THE ORIGINS OF IMMUNOLOGY From Metaphor to Theory Alfred I. Tauber and Leon Chernyak THEORY CHANGE IN SCIENCE Strategies from Mendelian Genetics Ltndley Dardcn THE MOLECULAR VISION OF LIFE Caltech, the Rockefeller Foundation, and the Rise of the New Biology Lily E. Kay HANS KRRBS The Formation of a Scientific Life, 1900-1933 Frederic; L. Holmes HANS KREBS: Architect of Intermediary Metabolism, 1933-1937 Frederic L. Holmes HANS KREBS Architect of Intermediary Metabolism 1933-1937 Volume I! FREDERIC LAWRENCE HOLMES New York Oxford OXFORD UNIVERSITY PRESS 1993 Oxford University Press Oxford New York Toronto Delhi Bombay Calcutta Madras Karachi Kuala Lumpur Singapore Hong Kong Tokyo Nairobi Dar es Salaam Cape Town Melbourne Auckland Madrid and associated companies in Berlin Ibadan Copyright © 1993 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 200 Madison Avenue, New York, New York 10016 Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data (Revised for vol. 2) Holmes, Frederic Lawrence. Hans Krebs. (Monographs on the history and philosophy of biology) Includes bibliographical references and indexes. Contents: v. 1. The formation of a scientific life, 1900-1933— v. 2. Architect of intermediary metabolism, 1933-1937. 1. Krebs, Hans Adolf, Sir. 2. Biochemists—Germany—Biography. I. Title. II. Series. QP511.8.K73H65 1991 574.19T092 [B] 91-2201 ISBN 0-19-507072-0 (v. 1) ISBN 0-19-507657-5 (v. 2) 135798642 Printed in the United States of America on acid-free paper For Catherine, Susan, and Rebecca This page intentionally left blank NOTES ON SOURCES Documents designated "KC" (Krebs Collection), were, at the time I used them, in the personal possession of Sir Hans Krebs. They were grouped by categories in boxes, but were not formally organized. They have since then been catalogued and deposited in the Sheffield University Library. I have not had opportunity to visit this archive in order to identify the catalogue entries under which the individual documents can now be found in that collection. Readers who wish to examine these materials should be able to locate them with facility by means of the comprehensive Catalogue of the Papers and Correspondence of Sir Hans Adolf Krebs FRS (1900-1981), compiled by Jeannine Alton and Peter Harper. Inquiries concerning the catalogue and the collec- tion should be addressed to The Librarian, Sheffield University, Sheffield S10 2TN, UK. For documents that I consulted after the collection had been catalogued in Oxford, 1 have included the entry numbers. I have referred to the laboratory notebooks of Hans Krebs by numbers that he and I affixed to their covers during the course of my visits to Oxford so that he could easily retrieve those that I wished to examine further. These numbers do not coincide with the numbering system adopted in the Catalogue and Correspondence. They can most easily be identified with the Catalogue entries through the dates covered by each note- book. References to transcribed conversations are given according to the participants, in the form "person interviewed-FLH," the date of the conversation, the number of the tape, and the page number of the taped transcription. Because there are some irregu- larities in the latter numbers some ambiguities result, and these references are given only as an informal guide to those who may wish to consult the transcripts. Micro- films of most, but not all, of the transcripts are on deposit in the Sheffield archive. The complete set of transcripts is presently in my possession. All translations from published and unpublished sources originally written in German, including Hans Krebs's laboratory notebooks, are my own. During the period covered by this work, the notations for organic compounds were less standardized than they have since become. I have, in general reproduced the rep- resentations of structural formulas originally used by the authors whose writings I cite, rather than to modernize them. In the footnotes the following abbreviations are used for titles of journals repeated- ly cited: VI11 NOTES ON SOURCES Ann. Chem. Annalen der Chemie und Pharmacie Arch. exp. Path. Archiv fur experimentelle Pathologic und Pharm- akologie Arch. ges. Physiol. Pfliiger' s Archiv fur die gesammte Physiologie des Menschen und der Thiere Arch, mikros. Anat. Archiv fur mikroskopische Anatomie Arch. Pharm. Naunyn-Schmiedeberg's Archiv fur Pharmakologie und experimentelle Pathologic Beitr. chem. Physiol. Beitrage zur chemischen Physiologie und Pathologic Ber. chem. Ges. Berichte der deutschen chemischen Gesellschaft Biochem. J. Biochemical Journal Biochem. Z. Biochemische Zeitschrift C. r. Acad. Sci. Comptes rendus hebdomadaires des seances de 1'Academic des Sciences C. r. Soc. Biol. Comptes rendus de la Societe de Biologie Erg. Physiol. Ergebnisse der Physiologie J. Biol. Chem. Journal of Biological Chemistry Journ. de physiol. Journal de physiologic et de pathologic generale J. Gen. Physiol. The Journal of General Physiology J. Physiol. The Journal of Physiology J. prakt. Chem. Journal fur praktische Chemie Klin. Woch. Berliner Klinische Wochenschrift Physiol. Rev. Physiological Reviews Proc. Roy. Soc. Land. Proceedings of the Royal Society of London Skand. Arch. Physiol. Skandinavisches Archiv fiir Physiologie Z. Biol. Zeitschrift fiir Biologie Z. exp. Med. Zeitschrift fiir die gesamte experimentelle Medizin Z. physiol. Chem. Hoppe-Seyler's Zeitschrift fur physiologischen Chemie ACKNOWLEDGMENTS Much of the content of the acknowledgments in volume I applies also to this volume. Here, therefore, I add only my special thanks to those whose help was particularly associated with volume II. Among the friends and colleagues of Hans Krebs whom I interviewed, those whose recollections have been incorporated into volume II include, Hermann Blaschko, Vernon Booth, Lady Margaret Krebs, E. Wolfgang Krebs, David Nachmansohn, Norman W. Pirie, Antoinette Pirie, Juda H. Quastel, Sir Edward and Lady Nan Wayne, and Hans Weil-Malherbe. Mrs. David Nachmansohn kindly gave me a group of letters written by Hans Krebs to her late husband. Mr. W. J. Hitchins, director of the Hans Krebs archive in Sheffield, was graciously hospitable during my visit there during the final preparation of volume II. It has been a special pleasure to work with two successive editors at Oxford Uni- versity Press: William F. Curtis, who encouraged me to submit a very long manu- script, and who astonished me by agreeing with enthusiasm to my tentative suggestion that it might be better to publish it in two volumes; and Kirk Jensen, who has effec- tively and graciously followed both volumes to completion. Careful management by Susan Hannan, and meticulous copyediting by Randi Laisi have contributed greatly to the effective production of volume II. Robert Berger, owner of the Write Way, capably produced the camera-ready copy for both volumes. He and I shared the frustrations of encountering many unanticipated problems, and the satisfaction of resolving most of them. Selda Lippa, who typed all of volume I and most of volume II, retired before the latter was finished. Her successor, Joanna Gorman, completed the work with great care and efficiency. In the last stages of preparation of the manuscript, when I was preoccupied with other matters and could barely look back at what I already viewed as finished, she and Pat Johnson took great care to ensure that all of the final details were accurately completed. In January 1992, my wife Harriet and I celebrated the appearance of volume I by traveling to England to visit again Hans Krebs's wife Margaret, and his brother Wolf. The joy of these reunions was tinged only by my regret that I could not share the occasion with Hans Krebs himself. I have wondered often how he would have regard- ed the outcome of a project to which he had contributed so much. I feel certain that he would have questioned the length of the volumes: in part because he valued succinct- ness in all forms of writing, in part because he would have doubted that the fine details of his scientific and personal life would be of interest to others. But I like to believe that he would otherwise have approved the way in which I have portrayed
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