Membrane Structure and Mechanisms of Biological Energy Transduction

595 Pages·1974·15.165 MB·English

by  Albert Szent-Gyorgyi (auth.), J. Avery (eds.)

Membrane Structure and
Mechanisms of Biological
Energy Transduction
Metnbrane Structure
and Mechanistns
of Biological Energy
Transduction
Edited by
J.
Avery
Department 01 Chemistry, Imperial College 01 Science and Technology
London S. W.7, England
9?
PLENUM PRESS· London and New York . 1973
Plenum Publishing Company Ltd.
Davis House
8 Scrubs Lane
Harlesden
London NWIO 6SE
Tel. [phone]
U.S. Edition published by
Plenum Publishing Corporation
227 West 17th Street
NewYorkNewYork 10011
Copyright © 1973 by Plenum Publishing Company Ltd.
Softcover reprint of the hardcover 1st edition 1973
Reprinted fromJournal 01 Bioenergetics Volume 3 (Numbers 1 and 2)
and Volume 4 (Numbers 1 and 2)
All Rights Reserved
No part of this publication 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
ISBN-I3: 978-1-4684-2018-0 e-ISBN-J3: 978-1-4684-2016-6
DOI: 10.1007/978-1-4684-2016-6
Library ufCongress Catalog Card Number: 72-95064
Contents
Part A: Mechanistns of Biological Energy
Transduction
The Development ofBioenergetics, Albert Szent-Gyärgyi
Chemiosmotic Coupling in Energy Transduction : A Logical
Development of Biochemical Knowledge, P. Mitchell 5
Solution ofthe Problem ofEnergy Coupling in Terms ofChemi-
osmotic Theory, V. P. Skulachev . 25
A Model of Membrane Biogenesis, Alexander T zagoloJ! . 39
Energy Transduction in the Functional Membrane of Photo-
synthesis: Results by Pulse Spectroscopic Methods, H. T. Witt. 47
Toward a Theory of Muscle Contraction, Gustavo Viniegra-
Gonzalez and Manuel F. Morales . 55
Bioenergetics of Nerve Excitation, Ichiji Tasaki and Mark Hallet. 65
An Analytical Appraisal of Energy Transduction Mechanisms,
R. J. P. Williams 81
Oxidative Phosphorylation, A History of Unsuccessful At
tempts: Is I t Only An Experimental Problem? Giovanni Felice
Azzone . 95
On the Coupling of Electron Transport to Phosphorylation,
Jui H. Wang . 105
Functional Organization of Intramembrane Particles of Mito-
chondrial Inner Membranes, L. Packer 115
On Energy Conservation and Transfer in Mitochondria,
Y. Hateji and W. G. Hanstein 129
An Enzymological Approach to Mitochondrial Energy Trans-
duction, John H. Young 137
ATP Synthesis in Oxidative Phosphorylation: A Direct-Union
Stereochemical Reaction Mechanism, Ephraim F. Korman and
Jerome McLick 147
The Electromechanochemical Model of Mitochondrial Struc-
ture and Function, David E. Green and Sungchul Ji 159
Part B: Metnbrane Structure
The Relationship of the (Na+ + K+)-Activated Enzyme
System to Transport of Sodium and Potassium Across the Cell
Membrane, J. C. Skou 203
On the Meaning ofEffects ofSubstrate Structure on Biological
Transport, Halvor N. Christensen . 233
VI CONTENTS
Performance and Conservation of Osmotic Work by Proton-
Coupled Solute Porter Systems, Peter Mitchell 265
Molecular Basis for the Action of Macrocyclic Carriers on
Passive Ionic Translocation Across Lipid Bilayer Membranes,
G. Eisenman, G. Szabo, S. G. A. McLaughlin and S. M. Ciani 295
Mechanisms of Energy Conservation in the Mitochondrial
Membrane, Lars Ernster, Kerstin Juntti and Kouichi Asami . 351
Biogenesis of Mitochondria 23. The Biochemica1 and Genetic
Characteristics ofTwo Different Oligomycin Resistant Mutants
of Saccharomyces Cerevisiae U nder the Influence of Cytop1asmic
Genetic Modification, Carolyn H. Mitchell, C. L. Bunn, H. B.
Lukins and Anthony W. Linnane 363
A Physio-chemica1 Basis for Anion, Cation and Proton Distribu-
tions between Rat-liver Mitochondria and the Suspending
Medium, E. J. Harris 381
Microwave Hall Mobility Measurements on Heavy BeefHeart
Mitochondria, D. D. Eley, R. J. Mayer and R. Pethig 389
Possible Mechanisms for the Linkage of Membrane Potentials
to Metabolism by Electrogenic Transport Processes with Special
Reference to Ascaris Muscle, P. C. Caldwell . 403
The Effect ofRedox Potential on the Coupling Between Rapid
Hydrogen-Ion Binding and E1ectron Transport in Chromato
phores from Rhodopseudomonas Spheroides, Richard J. Cogdell,
J. Barry J ackson and Antony R. Crolts 413
A Note on Some 01d and Some Possible New Redox Indicators,
Robert Hitt 431
The Role of Lipid-Linked Activated Sugars in Glycosylation
Reactions, W. J. Lennarz and Malka G. Scher 441
Lipid-Protein Interactions in the Structure ofBiologica1 Mem-
branes, Giorgio Lenaz 455
Structure-Function Unitization Model of Bio10gica1 Mem-
branes, D. E. Green, S. Ji and R. F. Brucker 527
The Influence ofTemperature-Induced Phase Changes on the
Kinetics of Respiratory and Other Membrane-Associated
Enzyme Systems, John K. Raison 559
Interactions at the Surface of Plant Cell Protoplasts; An
Electrophoretic and Freeze-etch Study, B. W. W. Grout,
J. H. M. Willison and E. C. Cocking 585
Part A
Mechanisms of Biological Energy
Transduction
Reprinted from Journal of Bioenergetics Volume 3 (Numbers 1 and 2)
Journal of BIOENERGETICS
Volume 3 Numbers 1-2 February 1972
Special Issue on :
MECHANISMS OF BIOLOGICAL ENERGY
TRANSDUCTION
Contents
The Development of Bioenergetics, Albert Szent-Gyorgyi
Chemiosmotic Coupling in Energy Transduction: A Logical
Development ofBiochemical Knowledge, P. Mitchell 5
Solution of the Problem of Energy Coupling in Terms of
Chemiosmotic Theory, V. P. Skulachev. 25
A Model of Membrane Biogenesis, Alexander T zagoloJ! . 39
Energy Transduction in the Functional Membrane of Photo-
synthesis: Results by Pulse Spectroscopic Methods, H. T. Witt 47
Toward a Theory of Muscle Contraction, Gustavo Viniegra-
Gonzalez and Manuel F. Morales . 55
Bioenergetics ofNerve Excitation, Ichiji Tasaki and Mark Hallet 65
An Analytical Appraisal of Energy Transduction Mechanisms,
J.
R. P. Williams 81
Oxidative Phosphorylation, A History of Unsuccessful At
tempts: Is It Only An Experimental Problem? Giovanni Felice
Azzone 95
On the Coupling of Electron Transport to Phosphorylation,
Jui H. Wang . 105
Functional Organization of Intramembrane Particles of Mito-
chondrial Inner Membranes, L. Packer 115
On Energy Conservation and Transfer in Mitochondria,
Y. Hateji and W. G. Hanstein 129
An Enzymological Approach to Mitochondrial Energy Trans-
duction, John H. Young 137
ATP Synthesis in Oxidative Phosphorylation: A Direct-Union
Stereochemical Reaction Mechanism, Ephraim F. Korman and
Jerome McLick 147
The Electromechanochemical Model of Mitochondrial Struc-
ture and Function, David E. Green and Sungchul Ji . 159
This special issue will also be available in hard cover form. Inquiries
should be sent to Plenum Publishing Co. Ltd., Davis House, Scrubs
Lane, Harlesden, London NW10 6SE, England.
1
Bioenergetics (1972) 3, 1-4
The Developlllent of Bioenergetics
Albert Szent-Gyorgyi
Marine Biological Laboratory, Woods Hole, Massachusetts
Living systems, being built of matter, and being driven by energy,
their analysis can be approached from two sides: that of matter or
from the side of energy. This division is analogous to the division into
structure and function, or, anatomy and physiology. This division
may be very convenient but is artificial, and exists only in our mind,
not in nature. Structure and function, essentially, are identical, two
sides of the same coin. Structure generates function and function
generates structure. A deeper understanding can be achieved only by
a fusion ofthe two.
The basic blueprint of bioenergetics is laid down, and progress can
be expected rather in the development in details than in the discovery
of new principles. What I would like to see is a more philosophical
attitude. "Philosophy" means the search for deeper and wider
relations. Needless to say, a philosophical approach presupposes the
knowledge of the details, and so it is no wonder that we should have
lost ourselves in details. All the same, the loss of a philosophical
attitude is regrettable and is characteristic for the contemporary
biology. The word "Science", with its present meaning, is of recent
origin. In the last century "natural philosophy" was used instead,
which indicates a different attitude.
If I were a newcomer in Bioenergetics, I would start my research
with trying to clear my mind about the meaning ofthe word "energy".
Living systems consist of atoms, and atoms are, essentially, a cloud of
electrons held together by nuclei. So energy can be only electronic
energy. Kinetic energy plays no major role in biology except in
activation. The source of our energies are the foodstuffs, and their
energy can be only electronic. So my first question would be how an
electron can release energy? Strictly speaking, electrons have no
energy, so how can they release any?
In trying to answer this question I would take my refuge to the
table of atoms of Mendeleef. It contains all the atoms out of which
Copyright © 1972 Plenum Publishing Company Limited. No part ofthis publication may be reproducecl,
stored in a retrieval s)"Stem, Of transmitted, in any form or by any means, electronic, mechanical, photocopying,
microfilming, recording or otherwise, without written permi~sion of Ple-num Publishing Company Limited.
2 ALBERT SZENT-GYORGYI
living systems are built. On its left side are the atoms which tend to give
off electrons to approach the noble gas structure, while on the right
are the atoms which te nd to take up electrons. A "trend to give off
electrons" involves that the electrons are loosely held, given off
easily, that is have a low ionization potential, are on a high energy
level. The "trend to take up electrons" means that there are low
energy levels capable of accommodating electrons. It follows that
energy can be released by electrons of an element of the left side by
interacting with an element of the right. In such interaction the
electrons of the former could go to a lower energy level of the latter
and doing so, would release energy.
What would probably strike me when taking a second look at this
table, is the extraordinary nature and position of H. It is a group by
itself. This atom is the only one which has the qualities needed to
support life. It is small, it parts easily with its electron and the proton,
left behind, can merge with the proton pool of the general solvent,
water. From this pool an electron can easily pick up a proton and form
an H with it. This would lead me to the conclusion that H had to be
the fuel of life. Looking over the elements of the right side I would
soon come to the conclusion that for many reasons oxygen only is fit
to act as general H- or electron-acceptor. However, the acceptor
qualities are not as specific as the donor ones, and in absence of O
2
various other elements mayaIso act as acceptors, and, evidently, had
to do so before O appeared in the atmosphere. About the possibilities
2
of storage of H, the Mendeleef table would tell me that only C, with
its position in the middle, is fit to build up the complex molecules
which could hold the H's.
If energy production only demands the interaction of an element
from the left ofMendeleeftable with an element ofthe right, then why
do not all atoms of the left side simply unite with the atoms on the
right side? Why do not all thermodynamically possible reactions take
place spontaneously? Should this happen then the whole biosphere
would burn up and there would be no place left for life, since no life
can exist without energy. Whatever the living machinery does, it has
to pay for it in terms of energy. Thus why do all thermodynamically
possible reactions not take place? Where is the "hang-up" which
makes life possible, and how can life make the reactions go which do
not take place spontaneously?
The hang-up is the activation energy, the fact that most chemical
reactions are, in a way, similar to a rock on the mountain side. By
rolling down, the rock would liberate great amounts of energy. If it
does not do so this is simply because in order to roll down it would
have to be loosened up, which demands a relatively small quantity of
energy which has to be invested. This small initial investment is
3
THE DEVELOPMENT OF BIOENERGETICS
which holds reactions up and makes life possible. What life does is to
"loosen up" H atoms, decrease the activation energy to a level at
which the energy of heat agitation is sufficient to make the reaction
go. This loosening up, this decrease of activation energy, is done by
enzymes, dehydrogenases, or "H-activation".
With these factors in mind I would try to follow the road on wh ich
life has built its machinery. Life builds gradually and does not reject
what it has built but builds on top. Consequently, the present cell is
comparable to an archeological excavation si te where one can find all
successive layers on top of one another, the oldest one the deepest. It
is like a pyramid. The oldest ones can be expected to be most widely
spread and anchored most firmly. By this I mean that the newer a
process, the more fancy its chemical equipment, the easier it will be
discarded, while the older and simpler on es will be retained more
tenaciously. For example: the most complete and newest energy
producing machinery is that of biologie al oxidation. It demands an
involved chemical structure. Ifthe cell divides it has to liquify and has
to give up its complex solid structures. So it gives up biologie al
oxidation and reverts for energy production to older and simpler
processes.
If we try to go back to the very beginning, the prebiotic period,
there we find a "primordial soup" with organic moleeules in it. When
life appeared it had to organize part of these moleeules into living
systems, while the others were used as "foodstuffs", a source of energy.
Ifwe imagined two molecules, side by side, a "living macromolecule"
and a foodstuffmolecule, the question would come up how could the
former take energy out of the latter?
According to what has been said before I can see one way only:
taking H atoms (or electrons) off the foodstuff moleeule. In order to
do this the binding of these H's would have to be loosened up, their
activation energy would have to be decreased. The H transfer
involves the "dehydrogenation" of the foodstuff molecule. Dehydro
genation being the oldest process we find dehydrogenases widely
spread in present living systems. They are actually the foundation of
our metabolism.
This simple scheme had to be complicated by the problem of
storage. Life needs energy all the time, but foodstuffs are accessible
only occasionally. The H's or electrons taken off the foodstuff had
thus to be stored to be used later according to need. There had to be
an H-, or electron pool. As I have pointed out in my book on
"The Living State" (now in press)* one of the most important
building blocks ofthe energy household is an H-or electron pool which
plays an important role even today and is, probably, the most
* Academic Press, New York.