Table Of ContentTopics in Environmental Physiology and Medicine
Edited by Karl E. Schaefer
Oxygen and
Living Processes
An Interdisciplinary Approach
Edited by
Daniel L. Gilbert
With 105 Figures
Springer-Verlag
N ew York Heidelberg Berlin
Karl E. Schaefer, MD.
Submarine Medical Research Laboratory
Groton, Connecticut 06340, U. S.A.
Daniel L. Gilbert, Ph.D.
Marine Biological Laboratory
Woods Hole, Massachusetts 02543, U.S.A.
and
Laboratory of Biophysics, NINCDS
National Institutes of Health
Bethesda, Maryland 20205, U.S.A.
Sponsoring Editor: Larry W. Carter
Production: William J. Gabello
Library of Congress Cataloging in Publication Data
Oxygen and living processes.
(Topics in environmental physiology and medicine)
Bibliography: p.
Includes index.
1. Oxygen in the body. 2. Oxygen-Physiological
effect. 3. Oxygen. I. Gilbert, Daniel L.
II. Series. [DNLM: 1. Oxygen. 2. Physiology.
QV312 0955]
QP535.0lO9 574.19'214 81-8899
AACR2
© 1981 by Springer-Verlag New York, Inc.
Softcover reprint ofthe hardcover 1st edition 1981
All rights reserved. No part of this book may be translated or reproduced in any form without
written permission from Springer-Verlag, 175 Fifth Avenue, New York, New York lOOlO,
U.S.A.
The use of general descriptive names, trade names, trademarks, etc. in this publication, even if
the former are not especially identified, is not to be taken as a sign that such names, as
understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely
by anyone.
Printed in the United States of America
9 8 7 6 5 432 1
ISBN-13: 978-1-4612-5892-6 e-ISBN -13: 978-1-4612-5890-2
DOl: 10.1007/978-1-4612-5890-2
Contents
Part I: General Aspects of Oxygen
1 Perspective on the History of Oxygen and Life 1
Daniel L. Gilbert
I. Ideas Preceding the Discovery of Oxygen 2
II. Events Leading up to the Discovery of Oxygen 4
III. The Discovery of Oxygen and Its Significance 11
IV. Oxygen Terminology 14
V. Early Biological and Medical Research with Oxygen 16
VI. Measurements of Atmospheric Oxygen and High
Altitude Effects 23
VII. Developments in the Nineteenth Century 25
VIII. Pasteur and Later Developments 28
IX. Intracellular Oxidation 29
X. Conclusion 30
2 Historical Introduction to the "Free Radical Theory"
of Oxygen Toxicity 44
Rebeca Gerschman
3 Reactive Forms of Oxygen 47
Sheldon Kanfer and Nicholas J. Turro
I. Atomic Oxygen 48
II. Dioxygen Species (Singlet Molecular Oxygen) 55
III. Trioxygen (Ozone) 59
IV. Superoxide 60
V. Summary 61
4 The Origin and Astronomical History of Terrestrial Oxygen 6S
George H Herbig
I. The Astronomical Scenario 65
II. The History of Terrestrial Oxygen 68
III. Summary 72
vi Contents
5 Significance of Oxygen on Earth 73
Daniel L. Gilbert
I. Geochemical Oxygen Cycle 73
II. Ozone 80
III. Beginning of Evolution on Earth 81
IV. The First Cells 84
V. Oxygen Production 86
VI. Development of Eukaryotes and Multicellular
Organisms 90
VII. Possible Future Events 93
VIII. Summary 94
Part II: Biology of Oxygen
6 Photosynthetic Oxygen Production 102
David C. Mauzerall and Richard G. Piccioni
I. Photosynthesis and Evolution 102
II. Overview of Modem Photosynthetic Mechanisms 105
III. The Formation of Oxygen 108
IV. Conclusion 119
7 Oxygen Toxicity in Unicellular Organisms 124
Sheldon F. Gottlieb
I. Introduction 124
II. Factors Affecting Responses to Increased Oxygen
Tensions 125
III. Kinetics 129
IV. Mechanisms of Oxygen Toxicity 131
V. Some Miscellaneous Effects of Oxygen 134
VI. Anaerobiosis and Mechanisms of Resistance to Oxygen
Toxicity 135
VII. The Use of Increased Oxygen Tensions as an
Antimicrobial 136
VIII. Conclusions 140
8 Oxygen Exchange in the Metazoa ISO
Johannes Piiper and Peter Scheid
I. Transport of Oxygen by Diffusion 150
II. Gas Exchange Organs: Structural Features 154
III. Gas Exchange Organs: Functional Organization 155
IV. External Medium: Air Versus Water 157
V. Internal Medium: Carriage of Oxygen by Blood 160
VI. Respiratory Organs of Vertebrates and Their Models 162
VII. Models for Gas Exchange: Qualitative Description 165
VIII. Models for Gas Exchange: Quantitative Analysis 167
IX. Application ofM odel Analysis in Animal Experiments 170
X. Real Systems and Model Analysis 172
XI. Summary 174
Contents vii
9 Facilitated Oxygen Difjusion by Oxygen Carriers 177
Jonathan B. Wittenberg and Beatrice A. Wittenberg
I. Steady-State Partial Deoxygenation of Myoglobin in
Active Tissue 178
II. Myoglobin in Tissue Oxygen Delivery 179
III. Leghemoglobin-Mediated Oxygen Delivery in Plants 180
IV. Myoglobin-Facilitated Oxygen Diffusion 182
V. Oxygen Delivery in Saline-Perfused Muscle 187
VI. Oxygen Gradients in Isolated Cardiac Cells 187
VII. Oxygen Gradients in Isolated Hepatic Cells 189
VIII. Oxygen Transfer to Mitochondria and Bacteroids 191
IX. Oxymyoglobin as an Electron Acceptor 191
X. Myoglobin-Associated Iron Protein 192
XI. The Myoglobin Molecule 192
10 The Reaction of Oxygen with Cytochrome Oxidase: The Role
of Sequestered Intermediates 200
Britton Chance
I. Introduction 200
II. A Possible Mechanism for Oxygen Reduction 200
III. Large-Scale Structural Aspects 205
IV. Short-Range Structural Features 206
V. Summary 208
11 Biochemical Aspects of Oxygen Toxicity in the Metazoa 210
Gyula B. Kovachich and Niels Haugaard
I. Molecular Mechanisms of Oxygen Toxicity 211
II. Oxygen Toxicity in Brain Tissue 213
III. Factors Influencing Susceptibility to Oxygen Poisoning
in Vivo 221
IV. Concluding Remarks 225
12 Antioxidant Defenses 235
Henry 1. Forman and Aron B. Fisher
I. Introduction 235
II. Mechanisms of Protection 237
III. Evidence for the Role of Antioxidant Defense
Mechanisms 243
IV. Summary 246
13 Superoxide Radical and Superoxide Dismutases 250
Irwin Fridovich
I. Some Biological Sources of Superoxide 250
II. The Potential Consequences of Superoxide Production 251
III. Assays for Superoxide Dismutases 253
IV. Varieties of Superoxide Dismutases 257
V. Mechanisms of Enzymatic Superoxide Dismutation 259
VI. The Physiological Role of Superoxide Dismutases 261
viii Contents
Part III: Human Aspects of Oxygen
14 Pulmonary Oxygen Toxicity 273
Gary L. Huber and David B. Drath
I. Terminology and Relative Concepts 273
II. The Lung as a Target Organ of Oxygen Toxicity 277
III. Clinical Manifestations of Pulmonary Oxygen
Toxicity 278
IV. Pulmonary Function in Oxygen Toxicity 279
V. The Pulmonary Pathology of Oxygen Toxicity 284
VI. Oxygen Toxicity and the Pulmonary Surfactant
System 290
VII. Oxygen Toxicity and Pulmonary Infection 297
VIII. Biochemical Mechanisms of Pulmonary Oxygen
Toxicity 302
IX. Sequence of Injury in Pulmonary Oxygen Toxicity 305
X. Pathogenesis of Oxygen Toxicity 310
XI. Oxygen Tolerance 312
XII. Modification of Oxygen Toxicity 321
XIII. Summary 323
15 Oxygen in Closed Environmental Systems 343
Karl E. Schaefer
I. Clinical Manifestation of Central Nervous System Oxygen
Toxicity 346
II. Etiology 347
III. Prevention of CNS Oxygen Toxicity 348
IV. Pulmonary Oxygen Toxicity 349
V. Clinical Manifestations and Indices of Pulmonary Oxygen
Toxicity 350
VI. Pulmonary Oxygen Toxicity during Spacecraft and
Aircraft Operations 350
VII. Pathology of Pulmonary Oxygen Toxicity 351
VIII. Etiology for Pulmonary Oxygen Toxicity 352
IX. Effects of Hyperoxia on Other Organ Systems 352
X. Oxygen Paradox 352
XI. Hypoxia 353
XII. Summary 354
16 Oxygen Tension in the Clinical Situation 358
LA. Silver
I. The Historical Background 358
II. Oxygen Tension and Radiotherapy 359
III. Tissue P02 in Surgical Conditions 359
IV. Oxygen Toxicity 360
V. Hyperbaric Oxygen 361
VI. Pediatric Application 361
VII. Current Practice and Problems 361
Contents ix
VIII. Future Developments 363
IX. Conclusion 365
17 Retinopathy ofP rematurity and the Role of Oxygen 368
Robert W. Flower and Arnall Patz
I. Pathogenesis of the Retinopathy of Prematurity 368
II. Newborn Animal Model of RLF 370
III. Monitoring of Blood Oxygen Levels 371
IV. Summary 374
Part IV: Concluding Remarks
18 Oxygen: An Overall Biological View 376
Daniel L. Gilbert
I. Hypoxia 378
II. Hyperoxia 380
III. Normoxia 382
IV. Practical and Clinical Considerations 384
V. Summary 386
Index 393
Preface
The field of oxygen study is immense. No single work on the subject can be
comprehensive, and this volume makes no such claim. Indeed, coverage here is
selective and the selection is somewhat personal. However, the choice of topics is
vast There are chapters on the history of oxygen, oxygen in the universe, the
biochemistry of oxygen, and clinical uses of oxygen. An alternate title could have
been, "Some things you always wanted to know about oxygen, but didn't know where
to find them easily." Some information in this wide-ranging work can not be found
elsewhere.
This book is intended not only for specialists, but also for nonspecialists engaged in
or curious about any field of oxygen study, particularly if they wish to know more
about other fields of oxygen. Thus, those who are interested in oxygen and are
historians, astronomers, chemists, geochemists, evolutionists, biochemists, physiolo-
gists, pathologists, or clinicians will find here much of extreme value. It is intended to
be read and understood at the graduate or advanced undergraduate level.
This volume is divided into four parts. The first constitutes the background for
Parts II and III, and the last integrates the preceding material with an overall
perspective on oxygen in living organisms.
Part I begins by putting oxygen into its historical context, including quotes from
original sources; this leads up to the presentation of the Gerschman "free radical
theory" of oxygen action in biological systems. The chemistry of the different reactive
forms of oxygen and the history of this reactive substance in the entire universe and on
earth are then presented. The present oxygen cycle in nature, with its photosynthetic
production and its removal by metazoan life, is described.
Part II begins a description of the biology of oxygen, with the mechanism of
photosynthetic oxygen production. The reader is introduced first to the harmful
effects of oxygen in unicellular organisms and then to the mechanisms of oxygen
transport to the tissues of multicellular organisms. The reaction of oxygen with
cytochrome oxidase in the cell is the beginning of the biochemical events that
eventually lead to the utilization of energy released by oxygen reduction in the cell.
The uncontrolled release of oxidative energy produces harmful effects in the
multicellular organisms, and description of these effects and the mechanisms of
control in living organisms, by such naturally occurring antioxidants as superoxide
dismutase, complete this part of the book.
Part III deals with the clinical and practical uses of oxygen. Since the lung is
normally exposed to higher oxygen tensions than other organs, the detrimental effects
of oxygen in the lung are discussed first. With the advent of manned exploration of
xii Preface
undersea, high-altitude, and space environments, the optimal supply of oxygen to
humans is of paramount importance, and this receives extensive treatment. The last
chapters in this section deal with the clinical use of oxygen; of special mention is
retrolental fibroplasia, a disease causing blindness in premature babies who have been
treated by excess oxygen-inhalation therapy.
Part IV integrates the entire subject of oxygen in living organisms. It becomes
apparent here that a multidisciplinary approach is vital to grasp a topic of such vast
proportions and consequences. The contributors to this volume are recognized
authorities in their various fields, ranging from microcosm to macrocosm. Their
approaches are blended here to provide a rich contribution to our understanding of
this gas in living organisms.
The reader will encounter, running through the chapters, the theme of the dualism
of oxygen. Oxygen provides the energy that nourishes most of life on earth, but at the
same time it destroys life. Aerobic life requires a proper balance between these
opposing forces in order to maintain the optimal functioning of a biological system.
Many individuals have contributed to my understanding of oxygen over the years.
Dr. R Gerschman's fascinating seminar on oxygen initiated my interest, which led to
an active collaboration between the two of us. Dr. W. O. Fenn, my Ph.D. advisor,
supported this collaboration and was a source of inspiration to me. Dr. H. Rahn
broadened my understanding of respiration physiology. Dr. K. Schaefer, editor of this
series, invited me to be the editor of this volume and encouraged me throughout its
preparation. He also invited me to present publicly for the first time in 1958 my
thoughts on oxygen and evolution.
Besides these individuals, I wish to thank my wife, Dr. Claire Gilbert, for acting as
my editorial assistant in the preparation of this volume, and my son, Raymond
Gilbert, for being extremely patient with me while this book was in preparation.
Finally, I wish to thank both my parents without whose support in my formative years
this volume would never have been undertaken.
I would also like to take this opportunity to thank the staff of Springer-Verlag,
whose endeavors have made this volume possible.
Daniel L. Gilbert
