Table Of ContentMechanistic 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
