Membrane Transport: An Interdisciplinary Approach

342 Pages·1977·13.387 MB·English

by  Arnošt Kotyk, Karel Janáček (auth.)

BIOMEMBRANES
Volume 9
BIOMEMBRANES
A series edited by Lionel A. Manson
The Wistar Institute, Philadelphia, Pennsylvania
Recent Volumes in this Series
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Edited by D. H. Smyth
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Edited by Henryk Eisenberg, Ephraim Katchalski-Katzir, and Lionel A. Manson
1976. Biomembranes • Volume 8
Articles by Robert W. Baldwin, William C. Davis, Paul H. DeFoor,
Carl G. Gahmberg, Sen-itiroh Hakomori, Reinhard Kurth, Lionel A. Manson,
Michael R. Price, and Howard E. Sandberg
1977 • Biomembranes . Volume 9
Membrane Transport-An Interdisciplinary Approach
By Amost Kotyk and Karel Janacek
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BIOMEMBRANES,
Volume 9
MEMBRANE TRANSPORT
An Interdisciplinary Approach
Arnost Kotyk and Karel J amicek
Institute of Microbiology
Czechoslovak Academy of Sciences, Prague
PLENUM PRESS • NEW YORK AND LONDON
ISBN-13: 978-1-4684-3335-7 e-ISBN-13: 978-1-4684-3333-3
DOl: 10.1007/978-1-4684-3333-3
Library of Congress Catalog Card Number: 75-18942
Scientific Editor: Prof. Dr. J. Kostli', DrSc.
Scientific Advisers: Prof. Dr. J. Koryta, DrSc.
Doc. Dr. L Drobnica, CSc.
Published in coedition by ACADEMIA
Publishing House of the Czechoslovak
Academy of Sciences, Prague
and outside of the Socialist countries by
PLENUM PRESS
A Division of Plenum Publishing Corporation
227 West 17th Street, New York, N.Y. 10011
© A. Kotyk, K. Janacek, Prague 1977
Softcover reprint of the 1 st edition 1977
AI! 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 Publishers
PREFACE
Not many years ago, problems of membranes and transport
attracted the attention of but a few dozen enthusiasts, mainly physiolo
gists who recognize~ the significance of membranes for the stabilization
of the general steady state of organisms. The first symposium organ
ized some fifteen years ago could boast of the attendance of perhaps
fifty scientists (the remaining fifty were not yet sure that membranes
was the topic of their choice), ranging in specialization from physical
chemistry to bacterial genetics, who clairvoyantly decided to study
what now has become the number one subject at most congresses of
biophysics, physiology, and even biochemistry and microbiology.
As is the case with many rapidly developing fields, the interest
in membranes and transport seems to be growing out of bounds and
the whole field of membra no logy, interdisciplinary as it is, has
penetrated into the realms of a number of branches of physics,
chemistry, and biology. Its subject is primarily biological and, although
much has been done in the world to increase the "exactness" of
biology over the past thirty years, one cannot strive for a rigorous
mathematical description of biological phenomena since, as M. H.
Jacobs wrote appropriately back in 1935, "In the first place, it is
utterly hopeless for the biologist with the means at present at his
disposal, to reduce the variables that enter into his problem to the
small number usually encountered in physical investigations. He is
compelled, therefore, regretfully but of necessity, to be content with
a lesser degree of precision in his results than that attainable in the
so-called 'exact sciences'. It follows that in dealing with most
biological problems it is not only useless, but actually unscientific,
6
to carry mathematical refinements beyond a certain point, just as it
would be both useless and unscientific to employ an analytical balance
of the highest precision for obtaining the growth-curve of a rat."
The present book is rather an attempt at bringing together data
on membranes and their primary function, viz. transport of substances,
in a relatively small volume. Recent literature abounds with compendia,
highly specialized monographs and conference proceedings dealing
with various aspects of membranes, but it is our feeling that there is
a need for an introductory text that would integrate at least most of the
available information about membranes and transport.
Several years ago, it was possible to include in a single volume
devoted exclusively to transport studies (Kotyk and Janacek, Plenum
Press, New York, 1970; 2nd edition 1975) a large section of compara
tive aspects, including various microorganisms, plants and animal
tissues. At present, there are so many pieces of information available
from various biological objects that they deserve specialized reviews.
For this reason, only generally valid and most important observations
are included in the present book while, on the other hand, due
attention is being paid to membrane functions other than transport,
including their physical chemistry and biochemistry.
We are aware of the fact that some chapters of the present book
are better than others because some of the authors' personal experience
and perhaps even hobbies are reflected in them. We hope that the
reader will not fail to peruse the more erudite monographic works
that we refer to at the appropriate places in the text if he wishes to
obtain more enlightenment.
In preparing the manuscript we were greatly aided, both
through consultations and through technical assistance, by some of
our co-workers, in particular Dr. K. Sigler, Dr. J. Horak, and Dr. R.
Rybova of this laboratory. The most competent photography of Dr.
A. Wolfofthis Institute is gratefully acknowledged. When mentioning
these collaborators, we do not overlook the general stimulating
atmosphere created by the whole staff of the department of membra
ne transport. Without it, this book could not have been written.
Arnost Kotyk
Karel Janacek
Laboratory for Cell Membrane Transport
Institute of Microbiology
Czechoslovak Academy of Sciences
CONTENTS
Preface
1. Introduction (K. J.) . 11
1.1. Important events in the history of membranology 12
1.2. Evolution and transport . 14
1.3. Transport in space and time 16
2. Membranes (A. K.) 21
2.1. Surfaces and interfaces 22
2.2. Chemical composition 36
2.2.1. Lipids 36
2.2.1.1. Chemistry . 36
2.2.1.2. Distribution 45
2.2.1.3. Fatty acids 49
2.2.1.4. Solubilization 51
2.2.2. Proteins. 53
2.2.2.1. Types and occurrence . 53
2.2.2.2. Solubilization 57
2.2.3. Carbohydrates . 60
2.3. Structure of membranes . 61
2.3.1. Physico-chemical techniques 61
2.3.1.1. X-ray diffraction 61
2.3.1.2. Infrared spectroscopy . 63
2.3.1.3. Ultraviolet spectroscopy 64
2.3.1.3.1. . Absorption spectra . 65
2.3.1.3.2. Optical rotatory dispersion 66
8
2.3.1.3.3. Circular dichroism . . . . 67
2.3.1.4. Nuclear magnetic resonance 70
2.3.1.5. Electron spin resonance. . 75
2.3.1.6. Fluorescence ...... 79
2.3.1.7. Calorimetry and related techniques 82
2.3.1.8. Other techniques. . . 85
2.3.2. Organization of lipids. 86
2.3.2.1. Biological membranes 86
2.3.2.2. Artificial membranes . 95
2.3.3. Organization of proteins 99
2.3.4. Lipid - protein interactions and membrane structure. 10 1
2.4. Assembly of membranes 107
2.5. Electron'microscopy. . . 117
2.6. Isolation of membranes . . 119
2.7. Morphology and function of different biological
membranes ... 126
2.7.1. Plasma membrane . 126
2.7.1.1. Morphology . . . . 126
2.7.1.2. Functional properties 130
2.7.1.2.1. Antigenicity .. 130
2.7.1.2.2. Enzyme content .. 132
2.7.1.2.3. Cell walls ..... 133
2.7.1.2.4. Binding and receptor properties 145
2.7.2. Mitochondrion 147
2.7.3. Chloroplast . . . . . 152
2.7.4. Mesosome ..... 156
2.7.5. Endoplasmic reticulum 156
2.7.6. Golgi apparatus 158
2.7.7. Lysosome 161
2.7.8. Tonoplast .. . 163
2.7.9. Nucleus ... . 163
4-
2.7.10. Other membranes 165
Synopsis 165
3. Thermodynamics of transport (K. J.) . 169
3.1. Thermodynamic equilibrium, passive and active
transport processes . . . . . . . . 169
3.2. Thermodynamics of the steady state 172
3.3. Network thermodynamics 177
Synopsis 181
9
4. Transport of nonelectrolytes (A. K.) . . . . . 183
4.1. Principles of diffusion CK. J.) ....... 184
4.2. Diffusion across membranes CA. K. + K. J.) . 192
4.3. Kinetics of mediated transport . 207
4.3.1. Mediated or facilitated diffusion 207
4.3.1.1. Steady-state approach 208
4.3.1.2. Equilibrium approach. . 211
4.3.1.3. Two-site carrier . . . . 220
4.3.2. Primary active transport 222
4.3.2.1. Kinetics. . . . . 222
4.3.2.2. Combined systems 226
4.3.2.3. Energetics . . . . 228
4.3.3. Coupled transport 229
4.4. Chemical nature of nonelectrolyte transport systems 233
4.4.1. Group-translocation systems. 233
4.4.2. Oxidoreductive systems . . . . . . . . . . .. 235
4.4.3. Binding proteins . . . . . '. . . . . . . . .. 238
4.5. Distribution and role of nonelectrolyte transport. 239
Synopsis 240
5. Transport of ions (K. J.). . . . . . . . . . . . . 243
5.1. Equilibria of ions . . . . , . . . . . . . 243
5.1.1. A simple membrane equilibrium and membrane
potentials . . . . . . . . . 243
5.1.2. Gibbs-Donnan equilibrium 247
5.1.3. Diffuse electrical double layer 250
5.2. Electrodiffusion and membrane potentials 256
5.2.1. Introduction............. 256
5.2.2. The electrodiffusion equation-general considerations 259
5.2.3. Schlogl's (1964) derivation of the general differential
equation of electrodiffusion . . . . . . . . . . . 262
5.2.4. Henderson's equation-potential difference across
a continuous layer with constant concentration
gradients of individual ions . . . . . . . . . . . 269
5.2.5. Planck's procedure-potential difference across a
microscopically electro neutral continuous layer 271
5.2.6. Goldman's procedure-potential difference across
a continuous layer with constant field. . . . . . . 279
5.2.7. Constant-field equation for potential difference across
the whole membrane . . . . . . . . . . . . . . 282
10
5.2.8. Constant-field equation for steady-state membrane
potential in the presence of an electrogenic sodium
pump ..................... 284
5.2.9. The Hodgkin-Horowicz equation. . . . . . . . . 286
5.3. Chemical nature of ion-translocating systems (A. K.) 288
5.3.1. Na,K-Adenosinetriphosphatase 288
5.3.2. Ca-Adenosinetriphosphatase.. 293
5.3.3. Other adenosinetriphosphatases 294
5.3.4. Ion-binding proteins . 294
5.3.5. Transport of ferric ions . . . . 295
5.3.6. Ionophores......... 296
5.4 Distribution and role of ion transport. 301
Synopsis 304
6. Transport of water (K. J.) 307
6.1. Steady-state thermodynamics of water permeation 307
6.2. The state of water in cells 314
Synopsis 316
7. Transport by special mechanisms (A. K.) 317
7.1. Oligopeptide permeases . . . . . . . 317
7.2. Pinocytosis . . . . . . . . . . . . 318
7.3. Uptake of nucleic acids and special proteins 320
Synopsis 323
References 325
Subject index 339