Membrane Structure and Mechanisms of Biological Energy Transduction PDF
Preview Membrane Structure and Mechanisms of Biological Energy Transduction
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. 01-9694727 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.
The list of books you might like
