The Organization of Cell Metabolism PDF
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The Organization of Cell Metabol ism NA TO ASI Series Advanced Science Institutes Series A series presenting the results of activities sponsored by the NA TO Science Committee, which aims at the dissemination of advanced scientific and technological knowledge, with a view to strengthening links between scientific communities. The series is published by an international board of publishers in conjunction with the NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics New York and London C Mathematical D. Reidel Publishing Company and Physical Sciences Dordrecht, Boston, and Lancaster o Behavioral and Social Sciences Martinus Nijhoff Publishers E Engineering and The Hague, Boston, Dordrecht, and Lancaster Materials Sciences F Computer and Systems Sciences Springer-Verlag G Ecological Sciences Berlin, Heidelberg, New York, London, H Cell Biology Paris, and Tokyo Recent Volumes in this Series Volume 123-The Molecular Basis of B-Cell Differentiation and Function edited by M. Ferrarini and B. Pernis Volume 124-Radiation Carcinogenesis and DNA Alterations edited by F. J. Burns, A. C. Upton, and G. Silini Volume 125-Delivery Systems for Peptide Drugs edited by S. S. Davis, L. ilium, and E. Tomlinson Volume 126-Crystallography in Molecular Biology edited by Dino Moras, Jan Drenth, Bror Strandberg, Dietrich Suck, and Keith Wilson Volume 127-The Organization of Cell Metabolism edited by G. Rickey Welch and James S. Clegg Volume 128-Perspectives in Biotechnology edited by J. M. Cardoso Duarte, L. J. Archer, A. T. Bull, and G. Holt Volume 129-Cellular and Humoral Components of Cerebrospinal Fluid in Multiple Sclerosis edited by A. Lowenthal and J. Raus Series A: Life Sciences The Organization of Cell Metabolism Edited by G. Rickey Welch University of New Orleans New Orleans, Louisiana and James S. Clegg University of California, Davis Davis, California Plenum Press New York and London Published in cooperation with NATO Scientific Affairs Division Proceedings of a NATO Advanced Research Workshop on the Organization of Cell Metabolism, held August 31-September 4, 1985, in Hanstholm, Denmark Library of Congress Cataloging in Publication Data NATO Advanced Research Workshop on the Organization of Cell Metabolism (1985: Hanstholm, Denmark) The organization of cell metabolism. (NATO ASI series. Series A, Life sciences; v. 127) "Published in cooperation with NATO Scientific Affairs Division." Proceedings of a NATO Advanced Research Workshop on the Organization of Cell Metabolism, held August 31-September 4, 1985, in Hanstholm, Denmark" - T.p. verso. Bibliography: p. Includes index. 1. Cell metabolism-Congresses. I. Welch, G. Rickey. II. Clegg, James S., 1933- . III. North Atlantic Treaty Organization. Scientific Affairs Division. IV. Title. V. Series. QH634.5.N38 1985 574.87'61 87-2468 ISBN-13: 978-1-4684-5313-3 e-ISBN-13: 978-1-4684-5311-9 DOl: 10.1007/978-1-4684-5311-9 © 1986 Plenum Press, New York Softcover reprint of the hardcover 1s t edition 1986 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 This volume is dedicated to the memory of the late Dr. Mario di Lullo, Programme Director in the NATO Scientific Affairs Division. Our workshop (like many others in the past) benefited greatly from his advice, generous assistance, and personal care. FOREWORD The NATO Advanced Research Workshop on nThe Organization of Cell Metabolismn was held 31 August - 4 September, 1985, at the Hotel Hansthol m in Hansthollllt Denmark. From areas of cell biology, bio chemistry, enzymology, and biophysical chemistry, the Workshop brought together workers whose research focuses on the character of the metabolic infrastructure of the living cell. The organizing committee was composed of the following members: James S. Clegg Douglas B. Kell Paul A. Srere G. Rickey Welch (Chairman) There has now arisen an edifice of proof that cell metabolism is highly ordered in space and time. The refinement of cytological extraction methodology and the development of more gentle isolation techniques have led to the demonstration of structural organization for most of the primary metabolic pathways. Moreover, the study of isolated enzyme complexes and membranous arrays has revealed unique functional properties of the organized state. Interest in metabolic organization has heightened very recently, due to the emergence of exciting new cytochemical and electron-microscopic advances in the elucidation of the cytOOIatrix and associated cell water. The hyaloplasmic space of eukaryotic cells has now been shown to be laced with a dense network of various filamentous structures - one role of which appears to be that of a structural support for the microcompartmentation of metabolic The supramolecular organization of certain processes may processe~ exceed the physical confines of intracellular particulates, involving large-scale entrainment of the cytoplasm in space and time. The Workshop dealt with the following major areas: 1) Organization Qf the cytomatrix ~ aqueous compartments. The aqueous milieu in the metabolic microenvironments of the cell may be different from the bulk aqueous condition in~. Cell-associated water must play an important role in the integration of metabolic In processe~ addition to their roles in scaffolding, organelle motility, and cellular motion, the cytomatrix may provide a structural setting for the organization of metabolic processes. 2) Organization Qf macromolecular synthesis. Macromolecular (e.~, protein) assemblage is an extremely complex process, invol ving the coordination of a multitude of enzymatic events. In eukaryotic cells many of these processes are associated with the cytomatrix. vii 3) Organization .2I. metabolic pathways. Organization of the mul tienzyme systems in intermediary metabolism may lead to such physiological advantages as "channeling" of intermediates (with the attendant elimination of diffusional transit, segregation of metabolic flow, etc.), unique modes of free-energy transduction, and coordination of metabolic regulatiOn. The organizational state may invol ve static spatial designs with protein-protein, protein-membrane, and/or protein-cytomatrix interactions. In some cases the organization may not be localized statically, but entail periodic spatio-temporal order extending throughout the bulk cytoplasm. 4) Experimental ~ theoretical modeling ~ metabolic organization. Realization of the organized nature of cellular metabolism necessitates the construction of theoretical paradigms which will prov ide us with a better qualitative and quantitative understanding of the workings of the cell. In addi tion, theoretical model ing opens up new vistas for experimental analysis of metabolic processes. Moreover, the emergence of immobilized-em:yme technology has fostered the usage of macroscopic model systems for exploring experimentally some of the unique advantages of the organized state. There were main lectures in each of these four areas. In. addi tion, there were round-table discussions focusing on topics of interest wi thin these headings. The round-tabl es were steered by inv ited experts. And, there was a poster session. * These publ ished proceedings incl ude the main 1 ectures, as well as presentations by the invited round-table panelists. As is usual in successful scientific gatherings, the discussions that took place outside the formal structure of the program were at least as important. As ev inced from the tabl e of contents herein, this Workshop was rather unique in the scope and breadth of coverage of the subject of "organization of cell metabolism". Our sincere thanks go to the contributing authors, for their time and energy invested in bringing this proj ect to frui tion. G. Rickey Welch, New Orleans James S. Clegg, Bodega Bay * Manuscripts were not sutmitted for the follOWing two lectures: "Structure and Evol ution of Ribosomes and Their Components" (by Dr. H. Wittmann) and "Time Patterns in Cellular Metabolism" (by Dr. B. Hess). viii ACKNGlLEDGEMENTS We would like to express our sincere thanks to NATO for major financial support. In particular, we are indebted to Dr. Mario Di Lullo, Programme Director in the Scientific Affairs Division, for his continued interest and advice. Additional financial support came from the following sources: Commission ..2.t: European Communities. A special note of thanks is due ~ to Paolo Fasella, Director-General for Science, Research, and D~ Development, for his interest and assistance. He himself has contributed significantly to the scientific subject of this Workshop. Regrettably, he was unable to attend due to pressing obligations of his office. International J1n1Qn..Q.( Biochemistry. We are especially grateful to Dr. W. J. Whelan of the Executive Committee, for his encouragement and interest and for his effort in the expeditious handling of our application to the IUB • ..NQI.Q Industri ~ (Denmark). The kind assistance of Dr. Villy Jensen (Research and Development Division) is much appreciated. We are very grateful to Mr. Kjeld Olsen and his staff at the Hotel Hans thol m, for their important role in the execution of our Workshop. We cannot say enough about their hospital ity and cooperation. The peaceful seclusion of the North Sea v illage of Hansthol m was perfect for a small, intensive scientific meeting. We would like to thank Mrs. Barbara Kester of the International Transfer of Science and Technology Office (Belgium), for much logistical help in the early planning of the Workshop. Last, but not least, we thank the Department of Biological Sciences (Chairman: Dr. Michael A. Poirrier), University of New Orleans, for help with the extensive costs of telephoning, mailing, photocopying, and printing, as well as for expert typing/clerical assistance. In particular, the warm smile, care, and skilled typing work of Mrs. Marie LeBlanc are beyond words. ix CONTENTS The Organization of Cell Metabolism: A Historical Vignette • • • • • 1 G.R. Welch and J.S. Clegg ORGANIZATION OF THE CYTOMATRIX AND AQUEOUS COMPARTMENTS Structural Organization of the Cytomatrix • • • • 9 K.R. Porter The Cytoskeleton • • • • 27 M. Osborn and K. Weber Interactions within the Cytoskeleton 37 A.B. Fulton On the Physical Properties and Potential Roles of Intracellular Water • • • • 4l J.S. Clegg Diffusion of a Small Molecule in the Aqueous Compartment of Mammalian Cells • • • • • 57 A.M. Mastro and D.J. Hurley Restricted Motion of Cellular ~ in Permeabilized Cells • • • • • 75 M. Kellermayer and C.F. Hazlewood The Intracellular Distribution of Adenosine Triphosphate • • • • • • • • • • • • • 79 S.B. Horowitz and D.S. Miller ORGANIZATION OF MACROMOLECULAR SYNTHESIS New Views of Cell and Tissue Cytoarchitecture: Embedment-Free Electron Microscopy and Biochemical Analysis • • • • • • • • • 89 E.G. Fey and S. Penman xi Understanding the Organization of Cell Metabolism in Early Embryonic S,ystems: Developmental Implications • • • • • • • • • 101 R. Marco, B. Batuecas, M. Calleja, M. Carra tal a, M. Cervera, A. Domingo, C. Ferreiro, R. Garesse, C. Urq ui a, and I. Vernos OBlANIZATION OF BIOSYNTHETIC AND BIODEGRADATIVE PROCESSES Organization of Enzymes in the Tryptophan Pathway 109 J.A. DeM>ss Implications of Metabolic Compartmentation in Prokaryotic Cells • • • • • • • • • 121 V. Moses Organization of Glucose Metabolism: A Model of Compartments by Poly-Isozymic Complexes 131 T. Ureta and J. Radojkovic Direct Transfer of Metabolites via Enzyme-Enzyme Complexes: Evidence and Physiological Significance 143 S.A. Bernhard and D.K. Srivastava Energetic Consequence of Dynamic Enzyme-Enzyme Interaction • T. Keleti Mitochondrial Hexokinase: Interactions between Glycolysis and Oxidative Phosphorylation • 169 J.E. Wilson Diffusion and Perfusion in the Living Cell: Implications for Metabolic Regulation and Organization • • • • • • • • • • • 171 D.N. Wheatley and P.C. Malone OBlANIZATION OF ENERGY METABOLISM: ENZYMOLOGICAL APPROACHES Interactions of Mitochondrial Matrix Enzymes with Mitochondrial Inner Membranes • • • • 177 R.L. TYiska, J.S. Williams, L.G. Brent, A.P. Hudson, B.J. Clark, J.B. Robinson, Jr., and P.A. Srere Some Aspects of Enzyme Organization in the Citric Acid Cycle • • • • 191 P.D.J. Weitzman and S.J. Barnes Enzyme Interactions in the Citric Acid Cycle and the Aspartate-Malate Shuttle • • • • 199 S. Beeckmans and L. Kanarek xii
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