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Quantitative Biology of Metabolism: Models of Metabolism, Metabolic Parameters, Damage to Metabolism, Metabolic Control PDF

310 Pages·1968·7.72 MB·English
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Quantitative Biology of Metabolism Quantitative Biology of Metabolism Models of Metabolism, Metabolic Parameters, Damage to Metabolism, Metabolic Control 3rd International Symposium, Biologische Anstalt Helgoland September, 26-29, 1967 Organizers A. Locker and F. Kruger Editor A. Locker With 66 Figures Springer-Verlag Berlin· Heidelberg. New York 1968 ISBN 978-3-540-04301-0 ISBN 978-3-642-51065-6 (Ebook) DOl 10.1007/978-3-642-51065-6 All rights reserved. No part of this book may be translated or reproduced in any form without written permissen from Springer-Verlag. © by Springer-Verlag Berlin·Heidelberg 1968 Library of Congress Catalog Card Number 68-55620 The use of gen~al descriptive names, trade names, trade marks, etc. in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Title No. 1527 Opening Address Ladies and Gentlemen, This is the third time that I have the honour to welcome the participants of the International Symposium on Ouantitative Biology of Metabolism here on Helgoland. I cordially welcome you to this big rock of red sandstone in the North Sea, and to the Marine Station of the Biologische Anstalt Helgoland! We are very glad that this symposium series has proved a valuable platform for the exchange of information and ideas concerning quantitative aspects of metabolism, and that many leading scholars representing a wide variety of biological and medical disciplines have come again in order to present and to discuss new scientific results. This third symposium has attracted the largest number of participants from English-speaking countries yet - an indication, it seems, of the growing international reputation of this con- ference. Please let me say a few words about the history of this symposium series. The idea to initiate biennial international and interdisciplinary symposia devoted to the fast developing field of quantitative biology was put forward in 1962 by Drs A. LOCKER (bsterreichische Studiengesellschaft fur Atomenergie GmbH, Wicn) and F. KRUGER (Biologische Anstalt Helgoland, Zentrale, Hamburg). I did not hesitate to support their idea and to offer the assistance of the Biologische Anstalt Helgoland as spon- soring institution. Thus the first two symposia of this series were organized by LOCKER, KRUGER und myself, and held in September 1963 and 1965, respectively, on Helgoland. All papers and discussions have been published in volumes 9 and 14 of the "Helgolander wissenschaftliche Meeresunter- suchungen" . From the beginning, I intended to devote myself to getting the sym- posium off the ground. Now that it is flying, I shall restrict myself to making available, as before, the assistance of our institution. As some of you may know, I am a marine biologist and experimental ecologist; as such I should no longer exert influence on an important field which progresses rapidly beyond my professional horizon. It is also felt that the proceedings should be published, from now on, in a special publication in order to do justice to their importance, not only to marine biologists but also to physiologists, biochemists, radiation biologists, geneticists, biomathe- VI Opening Address maticians, theoretical biologists and physicists. Certainly this will make it easier to reach the great diversity of institutions and individuals interested in the papers and discussions of this conference. Weare grateful that Springer-Verlag has agreed to act as publisher, and are convinced that the proceedings of these international symposia will continue to assist all colleagues interested in quantitative aspects of metabolism, as well as to encourage further studies in this exciting new scientific frontier. I am glad to notice that some of the "old-timers", who attended the first and second symposia, have returned and equally happy to recognize several "newcomers"- well-known scholars in their scientific fields- who have come for the first time thus providing a highly welcome source of "fresh blood". May this third symposium enlarge and advance our knowledge on quantitative aspects of metabolic processes as did the two preceding ones! I wish all of you rich hours of receiving and giving, and herewith open the Third International Symposium on Quantitative Biology of Metabolism. O. KINNE Biologische Anstalt Helgoland, Zentrale, Hamburg Preface The development of the life sciences may be said to have effected a gradual transition from a more or less intuitive prescientific approach based on crude observation, via a more refined type of observation to experimentation and hence to the level of formal theories. Quantitative methods are introduced at the second level; they comprise: (a) quantitative design of experiments; (b) regrouping of experimental results; (c) evalua- tion of results by means of mathematical or special statistical techniques. The last step implies the introduction of theoretical concepts, but we are not justified in speaking of theoretical science unless true theoretical con- siderations-models or hypotheses-precede experiment, and this is then followed by an attempt to link results with theory so as to verify the theory. Biology at present seems to lie somewhere between the second and third level mentioned above, not yet having achieved the status of a theoretical science in all its branches. Thus, though the need for quantification and mathematical formulation is widely recognized, many biologists still believe that e.g. general systems theory is too abstract to be of use in handling concrete problems. Those, however, who look critically at the present state of affairs cannot adopt this attitude. Here I would like to pay may respects to the founders of theoretical bi- ology: A. J. LOTKA in his Elements of Physical Biology laid the groundwork for the theoretical approach to almost all the problems of the life sciences, espe- cially those of metabolism and related phenomena, such as occur in ecology; N. RASHEVSKY, in several now classical monographs, in Mathematical Biophysics, and through the Bulletin of Mathematical Biop~ysics, which he founded, made a decisive contribution to the subject, and his efforts are now culminating in a universal theory of "Relational Biology"; L. VON BERTALANFFY used the term "Theoretical biology" in several monographs, and we owe to him the concept of steady-state (FlieGgleichgewicht) which, generalized as an open system, led him to the principal postulate of a General Systems Theory. The late H. QUASTLER deserves a special mention for having convinced biologists that information theory may usefully be applied to biology-and psychology. As a result of this pioneering work, a world-wide research movement was set off-though the centre of gravity lies in the United States. The time is now ripe for attempts to bridge the gap which remains in biology between steps two and three as defined above. VIII Preface The chief aim of the S.ymposia on Quantitative Biology of Metabolism, held every other year at the Biologische Anstalt, Helgoland, is to promote recognition by producing acceptable results. The first (publ: Helgolander wiss. Meeresunters., Vol. 9, 1964) comprised 36 papers, most of which dealt with experimental problems, and included a long discussion on the prob- lems of metabolism under adaptive conditions. But even in 1963, the need to define and delimit the subject produced some theoretical contri- butions. The second (publ: Helgolander wiss. Meeresunters., Vol. 14, 1966) included 41 papers, many considering the general implications of models and systems, while several outlined models for metabolic regulation and control based on differential equations or binary logic. The lines of investigation pursued at the first two Symposia are found again at the third, some of the contributions achieving an even higher level of formalization. Four interrelated themes, chief among them the application of models to metabolism, can now be distinguished. The section on models is prefaced by a general review of their role in human cognition. The two main types of model capable of representing meta- bolism and the reactions of biosystems in general are, depending on the mathematics involved, either discrete and continuous or deterministic and stochastic; examples of both are found here. They show how fruitful purely theoretical considerations can be when applied to otherwise intractable problems. Certain limitations inherent in the design of computers exclude mere representation of all possible reactions in a given system; however, computer efficiency can be improved by the omission of certain terms. It is interesting that, for certain systems of differential equations, the solutions are shown not be to elementary. Several more or less experimental trends in the quantitative biology of metabolism are illustrated by papers on the effects of temperature, activity stage, adaptation to temperature and season, growth and aging. Some of these problems run parallel in animals and plants. Among noxious influen- ces, ionizing radiations playa leading role, and the quantitative aspects of the mechanisms responsible for protection and damage repair are of great interest. For models of metabolic control, the optimization of reactions is of fundamental importance and may be based on irreversible thermody- namics, as well as on the theory of automata. The link with time is of great importance in the theory of metabolic control and the topic of biological rhythmicity is also involved. Control mechanisms based on coordination of elementary reactions display oscillatory features at the cellular, inter- cellular and systemic levels; the steady state in the central nervous system is a good example of the power of such coordination. The field of thermo- regulation requires a special model. Thus we may say that these papers represent a cross-section through several of the main lines of advance of quantitative (theoretical) biology; Preface IX it is for the reader to decide to what extent they constitute a positive con- tribution to an ever-growing field of great scientific topicality. This Sym- posium, like its predecessors, was a truly interdisciplinary occasion, bring- ing together experts from such diverse fields as mathematics, computer science, theoretical physics, hiophysics, biochemistry, human, animal and plant physiology, pharmacology and psychology. The fundamental nature of the subjects of our common study enables certain general conclusions to be reached which are of philosophical significance. The Symposium was held at the Biologische Anstalt Helgoland, Marine Station, through the kindness of the Leading Director, Prof. O. KINNE, who made ayailable to us in a most hospitable manner the excellent facilities of his Institute. The local organization was in the capable hands of Prof. and Mrs. F. KRUGER, who looked after the accommodation and welfare of the participants supremely well. I wish, too, to thank the Mayor of Helgo- land and the municipalit\ for the welcome they extended to us. Grants from the Federal Ministry of Scientific Research, Bonn-Bad Godesberg, for which I herewith expre5s my profound gratitude, enabled us to reimburse some of the participants' tranl expenses. The financial contribution made by the ()sterreichische Studiengesellschaft fur Atomenergie GmbH, Vienna, greatly helped me in publishing the program of the Symposium; this Society also generouslv assumed all the expenses associated with the editorial work. ?\1 \ special thanks go to Dr. K. F. SPRINGER of Springer- Verlag, Heidelberg, for agreeing to publish the proceedings of the Sym- posium. Last but not least, ~fiss H. KRUMPHOLZ assisted me in editing and retyping some of the manuscripts. In conclusion, may I express the hope that the material presented here will come into the hands of all who are interested in quantitative and theore- tical biolog\'. Vienna, Noyember, 1968. A. LOCKER Contents 1. Models of Metabolism 1.1. Models in General 1 A. LOCKER: The Epistemological Significance of Models in Science 1 2 H. KACSER, and J. A. BURNS: Causality, Complexity and Computers 11 1.2. Discrete and Continuous Models 3 R. ROSEN: Discrete and Continuous Representations of Metabolic Models . . . . . . . . . . . . . . . . . . . . . . . . .. 24 4 M. SUGITA: Interaction of a Digital (Genetic) System with a Con- tinuous (Enzymatic) One in the Cell ............. 33 5 CH. WALTER: A Model for Sustained Rhythmic Binary Logic in Biochemical Systems . . . . . . . . . . . . . . .. 38 1.3 Stochastic and Deterministic Models 6 A. F. BARTHOLmIAY: Some General Ideas on Deterministic and Stochastic Models in Biological Systems . . . . . . . . . . .. 45 7 P.-E. E. BERGNER: Turnover Compartmentalization. An Approach to Analysis of Whole-body Retention Data .......... 66 1.4 Computer Models of Cell Metabolism 8 Z. SIMON, D. FARCAS, and A. CRISTEA: Functioning of a Cell Model 71 9 J. A. BURNS: A Digital Computer System for the Construction and Analysis of Steady State Models of Enzyme Catalysed Networks. 75 10 H. J. BRBIERMANN: Computation of Equilibria and Kinetics of Chemical SYstems with Many Species. . . . . . . . . . . .. 81 2. Metabolic Parameters 2.1 Temperature - Season - Adaptation 11 J. N. R. GRAINGER: The Relation between Heat Production, Oxy- gen Consumption and Temperature in Some Poikilotherms. . .. 86 12 H.-D. JANKOWSKY: On Structure and Metabolic Rate of the Skeletal Muscle of the Eel (Anguilla vulgaris L.) . . . . . . . . . . .. 91 13 E. ZERBST: Analvsis of Heart Rate Adaptation to Temperature.. 95 14 S. GELINEO: The Heat Production of Goldfinches and Canaries in Summer and Winter . . . . . . . . . . . . . . . . . . .. 102 15 R. E. S~IITH, BARBARA A. HORWITZ, and Y. IMAI: Quantitative Ther- mogenesis of Brown Fat in Hibernation and Cold Adaptation 106 2.2 Growtb - Age 16 F. KRUGER: Contributions to the Energetics of Animal Growth. 113 17 P. R. PAYNE, and ERICA F. WHEELER: Models of the Growth of Organisms under Nutrient Limiting Conditions . . . . . . .. 127 XII Contents 18 F. COLLOT: Biomathematical Interpretation of Organismic Growth 133 19 L. W. WOODSTOCK: Relationships between Respiration during Imbi- bition and Subsequent Growth Rates in Germinating Seeds . .. 136 20 D. ADAMIKER: Quantitative Parameters of Rat Liver Mitochondria in Relation to Age . . . . . . . . . . . . . . . . . . . .. 147 2.3 Cellular Milieu - Experimental Influences 21 E. MeLLER: Enzymes as Indicators of Intracellular Reaction Condi- tions . . . . . . . . . . . . . . . . . . . . . . . . . 151 22 H. P. T. AMMON: The Influence of Ethanol on Carbohydrate- and Fat-Metabolism of the Liver . . . . . . . . . . . . . . . 154 3. Damage to Metabolism 3.1 Damage Due to Irradiation 23 N. ARLEY: Application of Quantitative Models for the Analysis of the Mechanisms of Protection against the Damaging Effects of Ionizing Radiation . . . . . . . . . . . . . . 157 24 J. KIEFER: Intracellular Recovery after Irradiation . . . . . .. 164 3.2 Damage Due to Radiolytic or Toxic Compounds 25 O. L. KLAMERTH: Inhibitory Action of Radiolytic Compounds on Cell Function . . . . . . . . . . . . . . . . . . . . . .. 177 26 A. FISCHER, and L. TAKACS: On the Release and Elimination of Liver Glutamic-Oxalacetic-Transferase . . . . . . . . . . .. 182 3.3 Damage Due to Oxygen Lack 27 A. T. MILLER: Metabolic Effects of Acute Hypoxia on Rat and Turtle Brain. . . . . . . . . . . . . . . . . . . . . . .. 185 28 H. R. SCHOEN, and R. E. Voss: Changes in Metabolite Concentrations in Ischemic Kidneys of Rabbits and in Transplanted Kidneys of Dogs 190 4. Metabolic Control 4.1 Metabolic Control and Optimization 29 G. DETCHEV, and A. MOSKONA t: The Mutual Coaction of two Regu- latory Mechanisms Realizing Optimal Autoregulation of Metabolism 195 30 A. CRISTEA: Optimization of Structure and Metabolism in Cellular Automata . . . . . . . . . . . . . . . . . . . . . . . .. 202 4.2 Metabolic Control at the Cellular Level 31 A. BETZ: Oscillatory Control of Glycolysis as a Model for Biological Timing Processes ..................... 205 32 E. L. COE, MARY H. COE, and IN-YOUNG LEE: Evaluation of the Coor- dinated Control of Activated Glycolysis and Respiration by Ortho- phosphate and ADP in Ascites Tumor Cells . . . . . . . . .. 217 33 P. ARESE, A. BOSIA, and L. ROSSINI: Profiles of Substrate Levels of the Central Metabolic Pathways in Frog Heart ....... 221 4.3 Control at the Systemic Level 34 J. KIRK, and J. S. ORR: Analysis by Computer of Some Oscillatory Features of the Red Cell System . . . . . . . . . . . . . .. 227

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