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Searching for a Mechanism: A History of Cell Bioenergetics PDF

297 Pages·2019·23.334 MB·English
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Searching for a Mechanism Searching for a Mechanism A History of Cell Bioenergetics John N. Prebble 3 1 Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America. © Oxford University Press 2019 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, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. Library of Congress Cataloging- in- Publication Data Names: Prebble, J. N. (John N.), author. Title: Searching for a mechanism : a history of cell bioenergetics / by John N. Prebble. Description: New York, NY : Oxford University Press, 2019. | Includes bibliographical references and index. Identifiers: LCCN 2018023975 | ISBN 9780190866143 Subjects: LCSH: Bioenergetics—History. Classification: LCC QH510.P74 2018 | DDC 572/.43—dc23 LC record available at https://lccn.loc.gov/2018023975 1 3 5 7 9 8 6 4 2 Printed by Sheridan Books, Inc., United States of America In memory of Peter Mitchell (1920– 1992), whose genius laid the foundation for the revolution in our understanding of cell bioenergetics. To Pat Contents Preface xi Acknowledgments xv List of Abbreviations xvii 1. Introduction: Respiration, phosphorylation, and mechanism 1 1.1 Respiration, photosynthesis, and bioenergetics 2 1.2 Vitalism 3 1.3 Historical questions 5 1.4 Phosphorylation 13 1.5 Mechanisms 14 1.6 The relevance of cell bioenergetics to the question of mechanism 15 2. From physiology to biochemistry: Respiration and oxidation from 1600 to 1900 17 2.1 Initiation of the experimental study of respiration 18 2.2 John Mayow’s Tractatus Quinque 20 2.3 Stephen Hales’s Vegetable Staticks 22 2.4 Respiration and combustion 23 2.5 The location of respiration 26 2.6 Thermodynamic questions 28 2.7 Bernard’s criticism of slow combustion 31 2.8 Hofmeister’s integration of cell biology 32 2.9 O and oxidation 33 2 2.10 Spectroscopy, hemoglobin, and animal pigments 35 2.11 Cell- free systems 38 3. Relating phosphorylation, respiration, and oxidation: 1900– 1945 41 3.1 Resolving nineteenth- century questions 41 3.2 Achievements of the first half of the twentieth century 42 3.3 Yeast and animal juices: The importance of phosphate 43 3.4 Thunberg, Wieland, and the nature of biological oxidation 47 3.5 Warburg’s Atmungsferment 51 3.6 Keilin’s cytochrome 54 3.7 DPN (NAD) and its oxidation 59 3.8 Muscle, lactic acid, and energy 61 3.9 Adenosine triphosphate and muscle phosphates 62 3.10 Aerobic ATP synthesis: Engelhardt and Kalckar 65 vii 3.11 Phosphorylation linked to respiration: Belitzer and Ochoa 68 3.12 Lipmann: The significance of phosphorylation 69 4. Emergence of the field of cell bioenergetics: 1945– 1960 73 4.1 Emergence of a new field 73 4.2 The mitochondrion as the location of respiratory activity 74 4.3 Further elucidation of the respiratory chain 76 4.4 Phosphorylation 82 4.5 Sites for phosphorylation 85 4.6 Seeking to understand the mechanism of phosphorylation 87 4.7 The phosphorylating enzyme 91 4.8 Fragmenting mitochondria 91 4.9 Physiological aspects of mitochondria 94 5. Defining the mechanism: 1960– 1977 97 5.1 What mechanism? 97 5.2 The first proton theory: Robert J. P. Williams 99 5.3 The chemiosmotic hypothesis of Peter Mitchell 100 5.4 Revising the respiratory chain 104 5.5 Exploring the ATP synthase 108 5.6 Reconstituting oxidative phosphorylation 111 5.7 Bacteriorhodopsin 112 5.8 Conformational theories 113 5.9 Mitochondrial membranes 117 5.10 Ion movements across the mitochondrial membrane 119 5.11 Resolving the mechanism 122 6. Discovering photosynthesis 128 6.1 The development of ideas on photosynthesis 128 6.2 Initial studies of photosynthesis 129 6.3 The importance of water and CO 131 2 6.4 Energy 132 6.5 Discovering chlorophyll and chloroplasts 133 6.6 Understanding the nature of photosynthesis 137 6.7 Photosynthetic bacteria and an oxidation– reduction mechanism 139 6.8 Light and dark reactions 141 6.9 O evolution and the Hill reaction 143 2 6.10 CO assimilation 146 2 6.11 Discovering photophosphorylation 149 7. Elucidating the photosynthetic light reaction 154 7.1 The fourth period of photosynthetic history 154 7.2 Seeking a coherent model of photosynthesis 155 7.3 The photosynthetic unit 158 viii Contents 7.4 Two light reactions and a reaction center 161 7.5 The Z- scheme and two photosystems 162 7.6 The contribution from bacterial photosynthesis 167 7.7 The chloroplast electron- transport chain 171 7.8 Chloroplast photophosphorylation 175 8. The impact of protein technology: 1977– 1997 181 8.1 The fifth period of investigation 181 8.2 The chemiosmotic mechanism for oxidative and photosynthetic phosphorylation 182 8.3 The Q cycle 183 8.4 Stoichiometric problems in mitochondria 186 8.5 Uncoupling and uncoupling proteins in mitochondria 188 8.6 Bacteriorhodopsin, a bioenergetic protein 190 8.7 Understanding the respiratory chain 194 8.8 Elucidating the mechanism of respiratory chain complexes 195 8.9 Photosynthetic complexes 200 8.10 Adenine nucleotide transport 205 8.11 The ATP synthase 207 9. The search for mechanism 214 9.1 Photosynthetic and oxidative biochemistry— the relationship 214 9.2 The course of bioenergetics history 216 9.3 Does methodology drive bioenergetics? 218 9.4 The significance of the membrane 223 9.5 Membrane proteins 224 9.6 The concept of mechanism 225 9.7 Revolution, hypothesis, and crisis 229 9.8 Resolving the crisis: Accepting the chemiosmotic theory 233 References 237 Index 267 Contents ix

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