Structure and Function in Excitable Cells PDF
Preview Structure and Function in Excitable Cells
Structure and Function in Excitable Cells Structure and Function in Excitable Cells Edited by Donald C. Chang 11aylor College of Medicine and Rice University Houston, Texas Ichiji Tasaki National Institute of Mental Health National Institutes of Health Bethesda, Maryland, and The Marine Biological Laboratory Woods Hole, Massachusetts William J. Adelman, Jr. National Institute of Neurological and Communication Disorders and Stroke National Institutes of Health at the Marine Biological Laboratory Woods Hole, Massachusetts and H. Richard Leuchtag Texas Southern University Houston, Texas, and University of Texas Medical Branch Galveston, Texas PLENUM PRESS • NEW YORK AND LONDON Library of Congress Cataloging in Publication Data Main entry under title: Structure and function in excitable cells. Bibliography: p. Includes index. 1. Excitation (Physiology). 2. Cell membranes. 3. Electrophysiology. I. Chang, Donald C. II. Title: Excitable cells. QP363.S8 1983 599'.087 83-13475 ISBN-13: 978-1-4615-9339-3 e-ISBN-13: 978-1-4615-9337-9 DOl: 10.1007/978-1-4615-9337-9 © 1983 Plenum Press, New York Softcover reprint of the hardcover 1s t edition 1983 A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013 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 William J. Adelman, Jr. Louis DeFelice Laboratory of Biophysics Department of Anatomy Intramural Research Program Emory University School of Medicine NINCDS Atlanta, Georgia 30322 National Institutes of Health at the Marine Biological Laboratory Woods Hole, Massachusetts 02543 George Eisenman Department of Physiology Gilbert Baumann UCLA Medical School Los Angeles, California 90024 Department of Physiology Duke University Medical Center Durham, North Carolina 277 10 Mark H. EUisman Martin Blank Department of Neurosciences University of California, San Diego Department of Physiology School of Medicine College of Physicians and Surgeons La Jolla, California 92093 Columbia University New York, New York 10032 Sachiko Endo Donald C. Chang Department of Biophysics and Biochemistry Faculty of Science Department of Physiology University of Tokyo Baylor College of Medicine Bunkyo·ku, Tokyo 113, Japan Houston, Texas 77030 and Department of Physics Rice University Houston, Texas 77251 Harvey M. Fishman Department of Physiology and Biophysics University of Texas Medical Branch D.F. Clapin Galveston, Texas 77550 Electron Microscopy Unit Department of Anatomy Faculty of Health Sciences Jarl Hagglund University of Ottawa of Ottawa, Ontario Canada KIN 9A9 Department Neurology and Marine Biological Laboratory University Hospital Woods Hole, Massachusetts 02543 S-751 23 Uppsala, Sweden v vi Contributors Nobutaka Hirokawa H. Richard Leuchtag Department of Anatomy and Neurobiology Department of Biology Washington University School of Medicine Texas Southern University St. Louis, Missouri 63110 Houston, Texas 77004 and Department of Physiology and Biophysics University of Texas Medical Branch Alan Hodge Galveston, Texas 77550 Laboratory of Biophysics NINCDS S. Rock Levinson National Institutes of Health at the Marine Biological Laboratory Department of Physiology Woods Hole, Massachusetts 02543 University of Colorado Health Sciences Center School of Medicine Denver, Colorado 80262 Kunihiko Iwasa Laboratory of Neurobiology James D. Lindsey National Institutes of Mental Health Bethesda, Maryland 20205 Department of Neurosciences and Marine Biological Laboratory University of California, San Diego Woods Hole, Massachusetts 02543 School of Medicine La Jolla, California 92093 Michael J. Kell Gilbert N. Ling Department of Anatomy Emory University School of Medicine Department of Molecular Biology Atlanta, Georgia 30322 Pennsylvania Hospital Philadelphia, Pennsylvania 19107 B.I. Khodorov Gen Matsumoto Vishnevsky Surgery Institute Electrotechnical Laboratory Moscow 113093, U.S.S.R. Tsukuba Science City, Ibaraki 305, Japan Takaaki Kobayashi Patrick Meares Department of Biochemistry Chemistry Department Jikei University University of Aberdeen Minato-ku, Tokyo 105, Japan Old Aberdeen AB9 2UE, Scotland J. Metuzals Harold Lecar Electron Microscopy Unit Laboratory of Biophysics Department of Anatomy IRP Faculty of Health Sciences National Institute of Neurological and University of Ottawa Communicative Disorders and Stroke Ottawa, Ontario, Canada National Institutes of Health Bethesda, Maryland 20205 and Marine Biological Laboratory Woods Hole, Massachusetts 02543 Graham V. Lees Hans Meves Departement de Biophysique I. Physiologisches Institut der Universitat des Saarlandes Laboratoire de Neurobiologie Cellulaire du CNRS 6650 Homburg, Saar F 91190 Gif sur Yvette, France West Gennany Contributors vii L. E. Moore Charles L. Schauf Department of Physiology and Biophysics Department of Physiology University of Texas Medical Branch Rush University Galveston, Texas 77550 Chicago, Illinois 60612 Catherine Morris Zadila Suarez-Mata Department of Biology Centro de Biofisica y Bioquimica University of Ottawa Instituto Venezolano de Investigaciones Cientificas Ottawa, Ontario, Canada KIN 9A9 Caracas 10IOA, Venezuela and Instituto International de Estudios Av anzados Caracas 1015A, Venezuela Hiromu Murofushi Department of Biophysics and Biochemistry Ichiji Tasaki Faculty of Science University of Tokyo Laboratory of Neurobiology Bunkyo-ku, Tokyo 113, Japan National Institutes of Health Bethesda, Maryland 20205 and Marine Biological Laboratory Yves Pichon Woods Hole, Massachusetts 02543 Departement de Biophysique Laboratoire de Neurobiologie Cellulaire du CNRS F 91190 Gif sur Yvette, France Torsten TeoreU Department of Physiology and Medical Biophysics Denis Poussart Biomedical Center Uppsala University Departement de Genie electrique S-751 23 Uppsala, Sweden Universite Laval Quebec GIK 7P4, Canada Gloria Villegas Centro de Biofisica y Bioquimica Francisco Rodriguez Instituto Venezolano de Investigaciones Cientificas Centro de Biofisica y Bioquimica Caracas IOIOA, Venezuela Instituto Venezolano de Investigaciones Cientificas and Instituto International de Estudios A vanzados Caracas IOIOA, Venezuela Caracas IOI5A, Venezuela and Instituto International de Estudios A vanzados Caracas 1015A, Venezuela Raimondo Villegas Centro de Biofisica y Bioquimica Jack Rosenbluth Instituto Venezolano de Investigaciones Cientificas Department of Physiology Caracas 10IOA, Venezuela New York University School of Medicine and Instituto International de Estudios Av anzados New York, New York 10016 Caracas IOI5A, Venezuela Hikoichi Sakai Clayton Wiley-Livingston Department of Biophysics and Biochemistry Department of Pathology Faculty of Science University of California, San Diego University of Tokyo School of Medicine Bunkyo-ku, Tokyo 113, Japan La Jolla, California 92093 John Sandblom Brendan S. Wong Department of Physiology and Medical Biophysics Department of Physiology University of Uppsala Baylor College of Dentistry S-751 23 Uppsala, Sweden Dallas, Texas 75246 Preface This book is a collection of up-to-date research reviews dealing with various aspects of the structure and function of excitable cells. Its overall objective is to further the search for a better understanding of the mechanism of excitation on a structural and physicochemical basis. The chapters are written by active investigators from a variety of disciplines, repre senting many different points of view. Their complementary fields of expertise give this book the rare feature of extraordinary breadth. Excitability is a fundamental property of many biological systems. The mechanisms by which nerve impulses are initiated and propagated, and by which rhythmical activities are produced in nerve, muscle, and cardiac cells, can be fully elucidated only when the process of excitation is derived from fundamental principles applied to known structural forms, at both the macroscopic and the molecular level. The problems of excitation are complex, requiring knowledge of many aspects of cells, including their morphology, elec trobiology, chemical physics, and biochemistry. From recent studies of excitable cells a number of new facts about their structure and function have emerged, so that it became imperative for physiologists interested in studies of cellular excitation processes to take these findings into consideration. Therefore, several investigators working at the Marine Biological Laboratory in Woods Hole decided to bring together for a discussion of these problems a group of electron microscopists, electrophy siologists, physical chemists and biochemists who have made significant contributions to our present-day knowledge of biological membranes. Thanks to financial aid from three U.S. government agencies-the Office of Naval Research, National Institute of Mental Health, and National Institute of Neurological and Communicative Disorders and Stroke and the administrative support of the Marine Biological Laboratory, this hope became a reality: At the International Conference on Structure and Function in Excitable Cells, from August 31 to September 3, 1981, some 130 researchers from 12 countries gathered in Woods Hole and exchanged their views on various problems related directly or indirectly to the cellular excitation mechanism. The present volume is based on the papers given by the invited lecturers at this Conference, expanded and brought up to date to keep pace with current developments. This book consists of four more or less distinct but interrelated parts. Part I deals with the results of recent studies of the ultrastructure of excitable cells. Among the many new tools that have become available to provide a clearer view of the structures of these cells is the increasing application of electron microscopy, in conjunction with rapid freezing, ix x Preface freeze-fracture and new fixation techniques. Ellisman, Lindsey, Wiley-Livingston and Levinson discuss the differentiation and maintenance of the membrane cytoskeletal struc ture in nerve. Rosenbluth studies the structure of the node of Ranvier of myelinated nerve fibers. Metuzals, Clapin and Tasaki present evidence for the existence of a highly organized structure consisting of different types of filamentous elements intimately connected to the plasmalemma and for the importance of this structure for the maintenance of excita bility. The chapter by Hodge and Adelman shows that neurofilaments and neurotubules, together with cross bridges, form a three-dimensional lattice. Hirokawa's application of the method of rapid freezing and deep etching provides new views of synaptic and axonal structures. Part II, introduced by Adelman, deals mainly with recent electrophysiological findings. Lecar, Morris and Wong review the work on the minute current pulses, known as "single channel currents," that have recently been observed in many membrane structures, and DeFelice and Kell discuss the relation of these to noise and impedance studies. Moore studies the complex admittance of muscle to observe the linear manifestations of its ion conductances, while the results of analyses of axon membrane noise are discussed by Pichon, Poussart and Lees. Chang presents a view of the ion selectivity of the conduction pathway based on a membrane cortex model. Baumann applies an aggregation model to stochastic processes. The use of specific neurotoxins to probe membrane behavior is discussed by Meves, who studies scorpion toxin, and by Khodorov, whose findings relate to batrachotoxin. In Part III, a physicochemical approach to studies of the cellular excitation mechanisms is adopted. Tasaki and Iwasa relate their findings on the mechanical responses of the axon membrane to structural studies of the axolemma-ectoplasm complex. Torsten Teorell, the founder of the Teorell-Meyer-Sievers theory for ionic membranes, gives an authoritative historical review of the development of physical chemistry of the membrane. Meares analyzes nonlinear membrane phenomena that arise from flux coupling. The universal biological solvent, water, has been replaced with heavy water in the studies of Schauf. Ling studies the effects of intracellular potassium-ion sorption on the production of cell potentials, and Eisenman, Sandblom and Hagglund analyze the behavior of narrow ion-conducting channels. A newly found connection between electrodiffusion and nonlinear waves is explored by Leuchtag and Fishman, and charged layers at the membrane surface are modeled by Blank. The fourth part describes various observations emphasizing the importance of protein molecules in the nerve membrane. Villegas, Villegas, Suarez-Mata and Rodriguez examine the behavior of reconstituted vesicles containing specific proteins from excitable cells. The last chapter, by Matsumoto, Murofushi, Endo, Kobayashi and Sakai, points to the necessity for the presence of microtubules for the maintenance of excitability in squid axon. This book is founded on the premise that to achieve a clear understanding of the mechanism of the excitation process, experts from many fields must join forces to attack the problem. Only by developing improved morphological techniques can we hope to learn about the structure of the "functional membrane"; only after we gain enough knowledge about the way ions and solvent molecules interact with protein molecules can we hope to know the mechanisms that control the flow of ions. We will need a better understanding of the chemical physics and physical chemistry of polyelectrolyte systems and of the physics of permeable membranes before we can explain how excitable cells work. Certainly we need further biochemical analyses to work out the chemical structure of the protein and phospholipid molecules in the membrane, so that we will know the kind of molecular apparatus with which we are dealing. The reader will see that the authors emphasize different aspects of the problem as they see them to be the most important ones. As a result, this Preface xi book explores a vast territory. There is very little doubt that some of these explorations will ultimately prove to be fruitful. We have organized this book in the belief and hope that such a multidisciplinary approach will help unlock the mystery of the excitation process. The help of Dr. Arthur B. Callahan of the Office of Naval Research, which has been particularly valuable both in organizing the conference and preparing this book, is gratefully acknowledged. Donald C. Chang Ichiji Tasaki William J. Adelman, Jr. H. Richard Leuchtag
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