Thermodynamic Network Analysis Of Biological Systems

Universitext J. Schnakenberg Thermodynamic Network Analysis of Biological Systems With 13 Figures Spri nger-Verlag Berlin Heidelberg New York 1977 Professor Dr. J. Schnakenberg Institut fOr Theoretische Physik, Rhein.-Westf.-Techn. Hochschule 0-5100 Aachen ISBN-13: 978-3-540-08122-7 e-ISBN-13: 978-3-642-96394-0 DOl: 10.1007/978-3-642-96394-0 Library of Congress Cataloging in Publication Data. Schnakenberg, J. Thermodynamic network analysis of biological systems. (Universitext) Based on two lectures presented by the author at the Rheinisch-Westfiilische Technische Hochschule in Aachen. Includes bibliographical references and index. 1. Biological physics. 2. Thermodynamics. I. Title. OH505.S313 574.1'912 77-1112 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. © by Springer-Verlag Berlin Heidelberg 1977. 2153/3130-543210 Preface This book is devoted to the question: What fundamental ideas and concepts can physics contribute to the analysis of complex systems like those in biology and ecology? The book originated from two lectures which I gave during the winter term 1974/75 and the summer term 1976 at the Rheinisch-Westfalische Technische Hochschule in Aachen. The wish for a lecture with this kind of subject was brought forward by students of physics as well as by those from other disciplines like biology, physiology, and engineering sciences. The students of physics were looking for ways which might lead them from their monodisciplinary studies into the interdisciplinary field between physics and life sciences. The students from the other disciplines suspected that there might be helpful physical concepts and ideas for the analysis of complex systems they ought to become acquainted with. It is clear that a lecture or a book which tries to realize the expectations of both these groups will meet with difficulties arising from the different trainings and background knowledge of physicists and nonphysicists. For the physicists, I have tried to give a brief description of the biological aspect and significance of a problem wherever it seems necessary and appropriate and as far as a physicist like me feels authorized to do so. For the nonphysicists, the physical background information is represented in the book as far as possible, for example a brief introduction into the fundamentals of thermodynamics in Chapter 3. I would be glad if this book stimulates the physicists to further reading of physiological texts, and the nonphysicists to study a more rigorous text of irreversible thermodynamics. The latter remark leads me to pOint out the significance of thermodynamics for the intention of this book. Very rigorously, one could suspect that its title "is only a fashionable paraphrase of simply "the role of thermodynamics in 1ife sciences". For two reasons, however, this would not hit the pOint. First, it is not just thermodynamics but thermodynamics of situations very far from equilibrium which plays the role of a ground theme for all our considerations in this book. Secondly, the conditions imposed by the fundamental laws of nonequilibrium thermodynamics will automatically be satisfied by utilizing a network language for the analysis of the systems that we have in view. At this pOint, I would like to express mY deep gratitude to the late Aaron Ka tcha 1sky. The 1ecture, "Network Thermodynami cs of Membranes", whi ch he gave on the occasion of the Israel ian-Ger