MEMBRANE PHYSIOLOGY Second Edition MEMBRANE PHYSIOLOGY Second Edition Edited by Thomas E. Andreoli, M.D. University of Texas Medical School Houston, Texas Joseph F. Hoffman, Ph.D. Yale University School of Medicine New Haven, Connecticut Darrell D. Fanestil, M. D. University of California, San Diego La Jolla, California and Stanley G. Schultz, M.D. University of Texas Medical School Houston Texas PLENUM MEDICAL BOOK COMPANY New York and London Library of Congress Cataloging in Publication Data Physiology of membrane disorders. Selections. Membrane physiology. "This volume is a reprint with minor modifications of parts I, II, and III of Physiology of membrane disorders, second edition, published by Plenum Medical Book Company in 1986"-T.p. verso. Includes bibliographies and index. 1. Membranes (Biology) 2. Biological transport. I. Andreoli, Thomas E., 1935- . II. Title. [DNLM: 1. Biological Transport. 2. Membranes-physiology. 3. Membranes-physiopathology. QS 532.5.M3 P5782m] QH601.P4825 1987 574.87'5 87-20229 ISBN-13: 978-0-306-42697-1 e-ISBN-13: 978-1-4613-1943-6 001: 10.1007/978-1-4613-1943-6 This volume is a reprint with minor modifications of Parts I, II, and III of Physiology of Membrane Disorders, Second Edition, published by Plenum Medical Book Company in 1986. © 1986, 1987 Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013 Plenum Medical Book Company is an imprint of Plenum Publishing Corporation All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Contributors EDWARD A. ADELBERG, M.D. DARRELL D. FANESTIL, M.D. Professor of Human Genetics Professor of Medicine Department of Human Genetics Head, Division of Nephrology Yale University School of Medicine University of California, San Diego New Haven. Connecticut 06510 La Jolla, California 92093 THOMAS E. ANDREOLI, M.D. GUIDO GUIDOTTI, PH.D. Edward Randall III Professor and Chairman Professor Department of Internal Medicine Department of Biochemistry and Molecular Biology Professor Harvard University Department of Physiology and Cell Biology Cambridge, Massachusetts 02138 University of Texas Medical School Houston, Texas 77225 JOSEPH F. HOFFMAN, PH.D. Eugene Higgins Professor of Physiology Department of Physiology AMIR ASKARI, PH.D. Yale University School of Medicine Professor and Chairman New Haven, Connecticut 06510 Department of Pharmacology and Therapeutics Medical College of Ohio PAUL HOROWICZ, PH.D. Toledo, Ohio 43699 Professor and Chairman Department of Physiology TED BEGENISICH, PH.D. University of Rochester School of Associate Professor of Physiology Medicine and Dentistry Department of Physiology Rochester, New York 14642 University of Rochester School of Medicine and Dentistry C. HUANG, PH.D. Rochester, New York 14642 Professor Department of Biochemistry MICHAEL D. CAHALAN, PH.D. University of Virginia School of Medicine Assistant Professor Charlottesville, Virginia 22908 Department of Physiology and Biophysics University of California MICHAEL J. JACKSON, PH.D. Irvine, California 92717 Professor of Physiology Department of Physiology MICHAEL J. CAPLAN George Washington University School of Medicine Graduate Assistant Washington, D.C. 20037 Department of Cell Biology Yale University School of Medicine JOHN A. JACQUEZ, M.D. New Haven, Connecticut 06510 Professor Department of Physiology TED H. CHIU, PH.D. The Medical School Associate Professor of Pharmacology Department of Biostatistics Department of Pharmacology and Therapeutics The School of Public Health Medical College of Ohio The University of Michigan Toledo, Ohio 43699 Ann Arbor, Michigan 48109 WILLIAM P. DUBINSKY, PH.D. JAMES D. JAMIESON, M.D., PH.D. Assistant Professor Professor of Cell Biology Department of Physiology and Cell Biology Department of Cell Biology University of Texas Medical School Yale University School of Medicine Houston. Texas 77225 New Haven, Connecticut 06510 v vi CONTRIBUTORS ROLF KINNE, M.D. J. DAVID ROBERTSON, M.D., PH.D. Max-Planck-Institut for Systemphysiology James B. Duke Professor and Chairman Dortmund, West Gennany Department of Anatomy Duke University Medical Center Durham, North Carolina 27710 PHILIP A. KNAUF, PH.D. Associate Professor HOWARD C. ROSENBERG, M.D., PH.D. Department of Radiation Biology and Associate Professor of Pharmacology Biophysics Department of Pharmacology and Therapeutics University of Rochester School of Medical College of Ohio Medicine and Dentistry Toledo, Ohio 43699 Rochester, New York 14642 STEVEN A. ROSENZWEIG ALEXANDER LEAF, M.D. Assistant Professor of Cell Biology Professor of Medicine Department of Cell Biology Ridley Watts Professor of Preventive Medicine Yale University School of Medicine Department of Preventive Medicine and Clinical Epidemiology New Haven, Connecticut 06510 Massachusetts' General Hospital Department of Medicine GEORGE SACHS, M.B., CH. B Harvard Medical School Professor of Medicine and Physiology Boston, Massa,chusetts 02114 Department of Membrane Biology University of California Los Angeles, California 90073 WERNER R. LOEWENSTEIN, PH.D. Professor Department of Physiology and Biophysics JAMES A. SCHAFER, PH.D. University of Miami School of Medicine Professor of Physiology and Biophysics Miami, Florida 33101 Nephrology Research and Training Center Departments of Physiology and Biophysics, and Medicine MARIANNA M. LONG, PH.D. University of Alabama School of Medicine Associate Professor Binningham, Alabama 35294 Laboratory of Molecular Biophysics University of Alabama School of Medicine MARTIN F. SCHNEIDER, PH.D. Binningham, Alabama 35294 Associate Professor of Physiology Department of Physiology ROBERT I. MACEY, PH.D. University of Rochester School of Professor of Physiology Medicine and Dentistry Department of Physiology-Anatomy Rochester, New York 14642 University of California Berkeley, California 94720 STANLEY G. SCHULTZ, M.D. Professor and Chainnan Department of Physiology and Cell Biology ANTHONY D. C. MACKNIGHT, M.D., PH.D. University of Texas Medical School Department of Physiology Houston, Texas 77225 University of Otago Medical School Dunedin, New Zealand CAROLYN W. SLAYMAN, PH.D. Professor of Human Genetics and Physiology LAZARO J. MANDEL, PH.D. Departments of Human Genetics and Physiology Associate Professor Yale University School of Medicine Department of Physiology New Haven, Connecticut 06510 Duke University Medical Center Durham, North Carolina 27710 T. E. THOMPSON, PH.D. Professor of Biochemistry Department of Biochemistry PAMELA A. PAPPONE, PH.D University of Virginia School Postgraduate Researcher of Medicine Department of Animal Physiology Charlottesville, Virginia 22908 University of California Davis, California 95616 DAN W. URRY, PH.D. Professor of Physiology and Biophysics JAMES W. PUTNEY, JR., PH.D. and Biochemistry Associate Professor Director, Laboratory of Molecular Biophysics Department of Pharmacology University of Alabama Medical College of Virginia School of Medicine Richmond, Virginia 23298 Birmingham, Alabama 35294 Preface Membrane Physiology (Second Edition) is a soft-cover book containing portions of Physiology of Membrane Disorders (Second Edition). The parent volume contains six major sections. This text encompasses the first three sections: The Nature of Biological Membranes, Methods for Studying Membranes, and General Problems in Membrane Biology. We hope that this smaller volume will be helpful to individuals interested in general physiology and the methods for studying general physiology. THOMAS E. ANDREOLI JOSEPH F. HOFFMAN DARRELL D. FANESTIL STANLEY G. SCHULTZ vii Preface to the Second Edition The second edition of Physiology of Membrane Disorders represents an extensive revision and a considerable expansion of the first edition. Yet the purpose of the second edition is identical to that of its predecessor, namely, to provide a rational analysis of membrane transport processes in individual membranes, cells, tissues, and organs, which in tum serves as a frame of reference for rationalizing disorders in which derangements of membrane transport processes playa cardinal role in the clinical expression of disease. As in the first edition, this book is divided into a number of individual, but closely related, sections. Part V represents a new section where the problem of transport across epithelia is treated in some detail. Finally, Part VI, which analyzes clinical derangements, has been enlarged appreciably. THE EDITORS IX Contents PART I: The Nature of Biological Membranes CHAPTER 1: The Anatomy of Biological Interfaces J. DAVID ROBERTSON 1. Introduction ....................................................... . 3 2. Models of Membrane Structure ........................................ 4 3. Membrane Junctions ................................................. 6 4. Certain Specialized Membranes ........................................ 10 5. Lipid-Protein Interactions in Model Membranes .......................... 14 6. Membrane Fluidity .................................................. 15 7. Mechanisms of Protein-Lipid Interactions ............................... 16 8. Mueller-Rudin Bilayer Membranes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 9. Freeze-Fracture-Etch Studies of Membranes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 10. Summary .......................................................... 17 References ......................................................... 17 CHAPTER 2: Composition and Dynamics of Lipids in Biomembranes T. E. THOMPSON and C. HUANG 1. Molecular Organization of Lipids in Biomembranes . . . . . . . . . . . . . . . . . . . . . . . . 25 2. Lipid Composition of Mammalian Cell Membranes ........................ 26 3. Molecular Structure of Membrane Lipids ................................ 26 4. Molecular Motions in Bilayers ....................................... . . 28 5. Interactions between Lipids in Bilayers .................................. 34 6. Summary .......................................................... 39 References ......................................................... 39 CHAPTER 3: Membrane Proteins: Structure, Arrangement, and Disposition in the Membrane GUIDO GUIDOTTI 1. Introduction ..................................................... . . . 45 2. Protein Composition of Membranes . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . 45 3. Types of Membrane Proteins .......................................... 45 4. Extrinsic Proteins ................................................... 46 5. Intrinsic Proteins .................................................. . . 47 6. Types of Intrinsic Membrane Proteins ................................... 48 7. The Structure of the Intramembrane Portion .............................. 52 xi xii CONTENTS 8. Disposition of Proteins in the Membrane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 9. Summary .......................................................... 53 References ......................................................... 54 PART II: Methods for Studying Membranes CHAPTER 4: The Nature and Limitations of Electron Microscopic Methods in Biology J. DAVID ROBERTSON 1. Introduction ........................................................ 59 2. The Conventional Transmission Electron Microscope ...................... 59 3. Specimen Preparation ................................................ 64 4. Embedding ........................................................ 66 5. Sectioning ......................................................... 72 6. Positive Staining .................................................... 72 7. Negative Staining ................................................... 75 8. Low-Dose Electron Microscopy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 9. The Freeze-Fracture-Etch (FFE) Technique ............................. 76 10. Summary .......................................................... 80 References ......................................................... 80 CHAPTER 5: Isolation and Characterization of Biological Membranes ROLF KINNE and GEORGE SACHS 1. Introduction ........................................................ 83 2. Methods for Dissociating and Separating Cells ............................ 83 3. Isolation of Plasma Membranes ........................................ 86 4. Assessment of Purification ............................................ 86 5. Expression of Data .................................................. 87 6. Functional Properties of Plasma Membranes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 7. Pathological Considerations ........................................... 88 8. Summary .......................................................... 90 References ......................................................... 91 CHAPTER 6: Absorption and Optical Rotation Spectra of Biological Membranes: Distortions and Their Corrections MARIANNA M. LONG and DAN W. URRY 1. Introduction ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 2. Distortions and Corrections for Suspensions .............................. 96 3. Purple Membrane as a Sample Calculation ............................... 10 1 4. Summary .......................................................... 106 References ......................................................... 106 CHAPTER 7: Mathematical Models of Membrane Transport Processes ROBERT I. MACEY 1. Introduction ........................................................ III 2. Diffusion .......................................................... III 3. Osmosis ........................................................... 118 4. Ionic Diffusion ..................................................... 119 5. Facilitated Diffusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . 125 6. Single-File Diffusion ................................................ 128 7. Summary .......................................................... 130 References ......................................................... 130 CONTENTS xiii CHAPTER 8: Application of Tracers to the Study of Membrane Transport Processes JOHN A. JACQUEZ 1. Introduction ........................................................ 133 2. Tracers and Problems in the Use of Tracers .............................. 133 3. Isotope Effects ..................................................... 135 4. Rate Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 5. Compartmental Systems and Compartmental Analysis ...................... 138 6. Theoretical Foundations for the Use of Tracers to Measure Rates ............. 141 7. Applications to the Study of Membrane Transports ........................ 143 8. Summary .......................................................... 150 References .......................................... . . . . . . . . . . . . . . . 150 CHAPTER 9: Principles of Electrical Methods for Studying Membrane Movements of Ions PAUL HOROWICZ, MARTIN F. SCHNEIDER, and TED BEGENISICH 1. Introduction ........................................................ 151 2. Steady-State Characteristics of Biological Membranes ...................... 151 3. Non-Steady-State Electrical Properties of Membranes ...................... 155 4. Fluctuation Analysis of Electrical Properties of Membranes. . . . . . . . . . . . . . . . . . 160 5. Single-Channel Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 6. Summary .......................................................... 163 7. Appendix .................................................. . . . . . . . . 163 References ........................................ . . . . . . . . . . . . . . . . . 164 CHAPTER 10: The Study of Transport and Enzymatic Processes in Reconstituted Biological Systems WILLIAM P. DUBINSKY 1. Introduction ........................................................ 167 2. Energy-Linked Processes ............................................. 168 3. Receptors and Channels .............................................. 170 4. Passive Transport System .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171 5. Summary .......................................................... 171 References ......................................................... 171 PART III: General Problems in Membrane Biology CHAPTER 11: Principles of Water and Nonelectrolyte Transport across Membranes JAMES A. SCHAFER and THOMAS E. ANDREOLI 1. Introduction ........................................................ 177 2. Diffusion as a Permeation Mechanism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 3. Activation Energy for Diffusion: Measurement and Significance. . . . . . . . . . . . . . 178 4. Convection ........................................................ 181 5. Osmosis. . . ... . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 6. Water and Nonelectrolyte Membrane Permeation Mechanisms ............... 183 7. Summary .......................................................... 188 References .................................................. . . . . . . . 188 CHAPTER 12: Anion Transport in Erythrocytes PHILIP A. KNAUF 1. Introduction .................................................. . . . . . . . 191 2. Modes of Anion Transport. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191