Mechanistic Approaches to Interactions of Electric and Electromagnetic Fields with Living Systems PDF
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Mechanistic Approaches to In teractions of Electric and Electromagnetic Fields with Living Systems Mechanistic Approaches to In teractions of Electric and Electromagnetic Fields with Living Systems Edited by Martin Blank Columbia University New York, New York and Eugene Findl Technical Consultants Group Encino, California Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data Mechanistic approaches to interactions of electric and electromagnetic fields with living systems. Includes bibliographical references and index. 1. Electromagnetism—Physiological effect. I. Blank, Martin, date. II. Findl, Eugene. QP82.2.E43M4 1987 574.19'17 87-7170 ISBN 978-1-4899-1970-0 ISBN 978-1-4899-1970-0 ISBN 978-1-4899-1968-7 (eBook) DOI 10.1007/978-1-4899-1968-7 © Springer Science+Business Media New York 1987 Originally published by Plenum Press, New York in 1987 Softcover reprint of the hardcover 1st edition 1987 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 PREFACE Although there is general agreement that exogenous electric and electromagnetic fields influence and modulate the properties of biological systems. there is no concensus regarding the mechanisms by which such fields operate. It is the purpose of this volume to bring together and examine critically the mechanistic models and concepts that have been proposed. We have chosen to arrange the papers in terms of the level of biological organization emphasized by the contributors. Some papers overlap categories. but the progression from ions and membrane surfaces. through macromolecules and the membrane matrix to integrated systems. establishes a mechanistic chain of causality that links the basic interactions in the relatively well understood simple systems to the complex living systems. where all effects occur simultaneously. The backgrounds of the invited contributors include biochemistry. biophysics. cell biology. electrical engineering. electrochemistry. electrophysiology. medicine and physical chemistry. As a result of this diversity. the mechanistic models reflect the differing approaches used by these disciplines to explain the same phenomena. Areas of agreement define the common ground. while the areas of divergence provide opportunities for refining our ideas through further experimentation. To facilitate the interaction between the different points of view, the authors have clearly indicated those published observations that they are trying to explain. i.e. the experiments that have been critical in their thinking. This should establish a concensus regarding important observations. In the discussion of theories. authors have emphasized the assumptions made. the published data incorporated. and the tests that have been done to evaluate the predictions. Wherever possible, quantitative estimates and illustrations have been given. We trust that this volume has provided a discussion of mechanism in the broadest sense. by giving an up-to-date summary of the ideas in the field. together with a critical evaluation that can guide us into the future. Martin Blank. Eugene Findl v CONTENTS IONS AND MEMBRANE SURFACES Ionic Processes at Membrane Surfaces: The Role of Electrical Double Layers in Electrically Stimulated Ion Transport ••••••••••• l M. Blank +t Membrane Transduction of Low Energy Level Fields and the Ca Hypothesis •••••••••••••••••••••••••••••••••••••••••••••••••••••• 15 E. Findl Electrochemical Kinetics at the Cell Membrane: A Physicochemical Link for Electromagnetic Bioeffects ••••••••••••••••••••••••••••• 39 A. Pilla, J.J. Kaufman and J.T. Ryaby Modification of Charge Distribution at Boundaries between Electrically Dissimilar Media ••••••••••••••••••••••••••• 63 C. Polk The Role of the Magnetic Field in the EM Interaction with Ligand Binding ••••.•••••••••••••••••••••••••••••••••••••••• 79 A. Chiabrera and B. Bianco Cyclotron Resonance in Cell Membranes: The Theory of the Mechanism ••••• 97 B.R. McLeod and A.R. Liboff Experimental Evidence for Ion Cyclotron Resonance Mediation of Membrane Transport •••••••••••••••••••••••••••••••••••••••••• 109 A. R. Liboff, S.D. Smith and B.R. McLeod Frequency and Amplitude Dependence of Electric Field Interactions: Electrokinetics and Biosynthesis •••••••••••••••••••••••••••••••• 133 L. A. MacGinitie, A.J. Grodzinsky, E.H. Frank and Y.A. Gluzband MACROMOLECULES The Influence of Surface Charge on Oligomeric Reactions as a Basis for Channel Dynamics ••••••••••••••••••••••••••••.••• 151 M. Blank vii Internal Electric Fields Generated by Surface Charges and Induced by Visible Light in Bacteriorhodopsin Membranes ••••••••••••••••• 161 F.T. Hong Interaction of Membrane Proteins with Static and Dynamic Electric Fields via Electroconformational Coupling ••••••••••••••••••••••• 187 T.Y. Tsong, F. Chauvin and R.D. Astumian Interactions Between Enzyme Catalysis and Non Stationary Electric Fields •••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 203 H.V. Westerhoff, F. Kamp, T.Y. Tsong and R.D. Astumian Patterns of Transcription and Translation in Cells Exposed to EM Fields: A Review ••••••••••••••••••••••••••••••••• 217 R. Goodman and A.S. Henderson Interaction of Electromagnetic Fields with Genetic Information •••••••• 231 P. Czerski and C.C. Davis MEMBRANE MATRIX Transient Aqueous Pores: A Mechanism for Coupling Electric Fields to Bilayer and Cell Membranes •••••••••••••••••••••••••••• 249 J.C. Weaver Electrorotation - The Spin of Cells in Rotating High Frequency Electric Fields ••••••••••••••••••••••••••••••••••••••••••••••••• 271 R. Glaser and G. Fuhr Membranes, Electromagnetic Fields and Critical Phenomena •••••••••••••• 291 J.D. Bond and N.C. Wyeth Field Effects in Experimental Bilayer Lipid Membranes and Biomembr anes .................................................... 301 H.T. Tien and J.R. Zon Fusogenic Membrane Alterations Induced by Electric Field Pulses ••••••• 325 A.E. Sowers and V. Kapoor INTEGRATED SYSTEMS Some Possible Limits on the Minimum Electrical Signals of Biological Significance •••••••••••••••••••••••••••••••••••••• 339 F.S. Barnes and M. Seyed-Madani Electrostatic Fields and their Influence on Surface Structure, Shape and Deformation of Red Blood Cells •••••••••••••••••••••••• 349 D. Lerche Cell Surface Ionic Phenomena in Transmembrane Signaling to Intracellular Enzyme Systems ••••••••••••••••••••••••••••••••• 365 W.R. Adey and A.R. Sheppard Low Energy Time Varying Electromagnetic Field Interactions with Cellular Control Mechanisms •••••••••••••••••••••••••••••••• 389 D.B. Jones and J.T. Ryaby viii The Mechanism of Faradic Stimulation of Osteogenesis •••••••••••••••••• 399 T.J. Baranowski, Jr. and J. Black The Role of Calcium Ions in the Electrically Stimulated Neurite Formation in Vitro •••••••••••••••••••••••••••••••••••••••••••••• 417 B.F. Sisken On the Responsiveness of Elasmobranch Fishes to Weak Electric Fields •• 431 H.M. Fishman Contributors •••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 437 Index ••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 441 IONIC PROCESSES AT MEMBRANE SURFACES: THE ROLE OF ELECTRICAL DOUBLE LAYERS IN ELECTRICALLY STIMULATED ION TRANSPORT Martin Blank Dept. of Physiology and Cellular Biophysics Columbia University, College of Physicians & Surgeons 630 W. 168 St., New York, NY 10032 INTRODUCTION Surface properties differ significantly from bulk properties. At charged membrane (or channel) surfaces the surface concentrations and surface potentials of ions differ from the bulk values, but the combined electrochemical potentials are the same. Any increase in surface concentration is exactly balanced by the decrease in electrical potential, so ions at the surface are in equilibrium with those in the bulk. Since ion transport is driven by electrochemical potentials, it is clear that the driving forces for the ions are the same at the surface as in the bulk solution. While this analysis justifies using the same electrochemical potential for an ion at the surface as in the bulk, it is nevertheless necessary to introduce surface concentrations when considering fluxes, even in steady state processes. Ionic fluxes depend upon the absolute concentrations as well as the electrochemical potential differences, so in calculating permeabilities (or conductances) from flux data one must use the appropriate concentration, i.e., the surface concentration (1). In non-steady state systems, the surface concentrations of ions can become quite different from bulk concentrations, particularly during current flow. Nernst and Riesenfeld (2) were the first to show ion concentration changes at liquid/liquid interfaces. Nernst (3) even developed a theory of excitation thresholds based on the accumulation of ions at a membrane surface. The actual concentration of ions at a surface was later measured with the aid of surface active ions (4), and the concentration changes during current flow were found to be large and long-lived. It was later shown that the effects of charged surfaces on interfacial transference (5) could be explained by the concentration changes in the electrical double layer region (6). These studies reenforced the idea that the properties of ions in the electrical double layers at membrane surfaces are important for an understanding of transport. In transient or non-steady state membrane processes, the two driving forces for ionic movement, the chemical potential for diffusion and the electrical potential for migration, change at different rates. A membrane can be depolarized quite rapidly, with time constants on the order of 1-10 microseconds, while chemical potentials readjust at much slower rates characteristic of diffusion processes over distances on the order of cell diameters, i.e., 1 millisecond; It is therefore possible to generate unbalanced chemical gradients for short periods of time by manipulating membrane (electrical) potentials. This disparity in the response times of the two forces that drive ions across membranes can lead to unusual transient ionic fluxes. An analysis based on these processes can account for the ionic fluxes seen in excitable membranes and also for the different apparent selectivities of channels that open at different rates (7). Finally, it is important to bear in mind that natural membranes normally separate solutions having very different compositions and concentrations, so large ionic gradients exist. Because of this asymmetry, small currents can cause large changes in the ionic concentrations in the surface regions. Also, the effects of alternating currents are additive rather than self-canceling. All of these changes are greatest in the surface regions and are best understood in terms of changes in surface concentration. Let us consider Table 1 to illustrate the effects of current flow on ion concentrations in the surface layers of a cation selective cell membrane. We have arbitrarily chosen layers that contain 220 ions in the steady state, showing only the layers at the membrane surface and the two layers adjacent to them. (Other layers, and those next to the two electrodes are omitted for simplicity.) The steady state concentrations are shown in part A. If we pass a current pulse of 22 charges, the ions·carrying the current are shown with arrows in part B, along with the resulting instantaneous surface concentrations (shown in the boxes). If the current is reversed, as in part C, the arrows change direction and different surface concentration·s result. This example ignores the effects due to charged membrane surfaces, diffusion effects due to the concentration differences that are generated, etc., but it does illustrate that large transient concentration changes can result. The effect of these changes can be seen in the ~E column. If the membrane potential is set primarily by the potassium ion, the resting Table 1. Concentration Changes at Membrane Surfaces during Transference Assumptions: Membrane Permeability K Na, CI=O Solution Mobilities K = Na = Cl A. Steady State Concentrations Inside Outside Inside Membrane Outside A E 100 K 10K 58 mV 10 Na 100 Na 110 CI 110 CI B. Cathodal pulse of 22 charges 20K+_ _- ++ 1K 1INa0 K$ 2Na -+----++o 10Na llC! lle! New Surface Concentrations: 90 K 29 K 29 mV 9 Na 92 Na 99 C! 121 CI C. Anodal pulse of 22 charges 2 K 1 K 20 Na1-f----+- 10 Na ::~~$ 11 CI New Surface 9 K 58.6 mV 90 Na 99 Cl 2
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