European Society for Comparative Physiology and Biochemistry Third Congress Organizing Committee: A. D. F. ADDINK, President Animal Physiology, Gorlaeus Laboratory, Rijksuniversiteit Leiden C. M. BALLINTIJN Zoological Laboratory, Rijksuniversiteit Groningen A. P. VAN OVERBEEKE Zoological Laboratory, Katholieke Universiteit Nijmegen L. M. SCHOONHOVEN Animal Physiology, Landbouw Hogeschool Wageningen N. SPRONK, Secretary Biological Laboratory, Vrije Universiteit Amsterdam D. I. ZANDEE Chemical Animal Physiology, Rijksuniversiteit Utrecht Sponsors: The Commission of the European Community Ministerie van Landbouw en Visserij Technische Hogeschool Delft Technische Hogeschool Twente Landbouwhogeschool Wageningen Rijksuniversiteit Leiden Rijksuniversiteit Utrecht Universiteit van Amsterdam Vrije Universiteit Proceedings of the Third Congress of the European Society for Comparative Physiology and Biochemistry, August 31-September 3, 1981, Noordwijkerhout, Netherlands Exogenous and Endogenous Influences on Metabolic and Neural Control of • RESPIRATION • FEEDING • ACTIVITY AND ENERGY SUPPLY IN MUSCLES • ION- AND OSMOREGULATION • REPRODUCTION • PERCEPTION AND ORIENTATION Volume 1 Invited Lectures Editors A. D. F. ADDINK Animal Physiology Gorlaeus Laboratory Rijksuniversiteit Leiden, Netherlands N. SPRONK Biological Laboratory Vrije Universiteit, Amsterdam, Netherlands PERGAMON PRESS OXFORD • NEW YORK • TORONTO • SYDNEY • PARIS • FRANKFURT U.K. Pergamon Press Ltd., Headington Hill Hall, Oxford OX3 0BW, England U.S.A. Pergamon Press Inc., Maxwell House, Fairview Park, Elmsford, New York 10523, U.S.A. CANADA Pergamon Press Canada Ltd., Suite 104, 150 Consumers Road, Willowdale, Ontario M2J 1P9, Canada AUSTRALIA Pergamon Press (Aust.) Pty. Ltd., P.O. Box 544, Potts Point, N.S.W. 2011, Australia FRANCE Pergamon Press SARL, 24 rue des Ecoles, 75240 Paris, Cedex 05, France FEDERAL REPUBLIC Pergamon Press GmbH, 6242 Kronberg-Taunus, OF GERMANY Hammerweg 6, Federal Republic of Germany Copyright © 1982 Pergamon Press Ltd. 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, electrostatic, magnetic tape, mechanical, photocopying, recording or otherwise, without permission in writing from the publishers. First edition 1982 Library of Congress Cataloging in Publication Data European Society for Comparative Physiology and Biochemistry. Congress (3rd: 1981: Noordwijkerhout, Netherlands) Exogenous and endogenous influences on metabolic & neural control of respiration, feeding, activity and energy supply in muscles, ion- and osmoregulation, reproduction, perception and orientation. Includes indexes. Contents: v. 1. Invited lectures — v. 2. Abstracts. 1. Metabolic regulation—Congresses. 2. Neurotrophic functions—Congresses. 3. Bio logical control systems—Congresses. I. Addink, A. D. F. (Alberti Daniel Francois), 1935- II. Spronk, N. III. Title. QP171.E97 1981 591.1'88 82-3657 AACR2 British Library Cataloguing in Publication Data European Society for Comparative Physiology and Biochemistry. Congress (3rd: 1981: Noordwijkerhout) Exogenous and endogenous influences on metabolic & neural control. Vol. 1: Invited lectures 1. Physiology, Comparative—Congresses I. Title II. Addink, A. D. F. III. Spronk, N. 574.1 QP1 ISBN 0-08-027986-4 (vol. 1) ISBN 0-08-028845-6 (vol. 2) In order to make this volume available as economically and as rapidly as possible the authors' typescripts have been reproduced in their original forms. This method unfortunately has its typographical limitations but it is hoped that they in no way distract the reader. Printed in Great Britain by A. Wheaton & Co. Ltd., Exeter PREFACE The control of physiological activities in animals is essential to life. During this Congress of the exogenous and endogenous influences on these metabolic and neural processes of control are discussed fundamentally, followed by an applied aspect. To the organizers the interaction of fundamental and applied aspects is essential (not only) during such a congress; it proved to be a basis for a fruit- full exchange of ideas for future research, in many ways, as was learned from the discussions. Especially in the context of this European Society for Comparative Physiology and Biochemistry it is important to cooperate and to prevent possible duplications in research. It will certainly lead to scientific agreements within Europe, for ap­ plied or fundamental projects. ACKNOWLEDGEMENTS We want to thank the following members of the Organizing Committee: Dr. Ballintijn, Dr. Van Overbeeke, Dr. Schoonhoven and especially Dr. Zandee for their invaluable activities in organizing, editing and indexing the separate book, containing the abstracts of posters and invited lectures. It was meant and proved to be the es­ sential basis for the 4 days of the Congress, August 31 to September 3, 1981 in Noordwijkerhout, The Netherlands. It is with great pleasure, that we remember the enthusiastic cooperation and at­ tention with increasing intensity during the past two years which resulted in the 3rd Congress of the European Society for Comparative Physiology and Biochemistry. The support and experience of Dr. Munday and Dr. Gilles our President and Executive officer of the Society were essential in the process of organization. Furthermore the incentive activities are gratefully acknowledged of the chairmen in the six Sections in the Congress: Dr. Dejours, Dr. De Ruiter, Dr. Beenakkers, Dr. Gilles, Dr. Laverack and Dr. Voogt. The Congress organization has had an explicit benefit of the accurate, helpfull and devoted activities of Trix van footen, Thea Laan, Danielle Zandee and Marc Spronk: not only the secretarial and financial administration before the Congress, but also their reception in a friendly manner at the Congress desk. For the final editing and indexing of the total manuscript of the Proceedings we are much obliged to Trix van Kboten for her expert typing activities. A.D.F. ADDINK and N. SPRONK Leiden, November 1981. PREFACE The control of physiological activities in animals is essential to life. During this Congress of the exogenous and endogenous influences on these metabolic and neural processes of control are discussed fundamentally, followed by an applied aspect. To the organizers the interaction of fundamental and applied aspects is essential (not only) during such a congress; it proved to be a basis for a fruit- full exchange of ideas for future research, in many ways, as was learned from the discussions. Especially in the context of this European Society for Comparative Physiology and Biochemistry it is important to cooperate and to prevent possible duplications in research. It will certainly lead to scientific agreements within Europe, for ap­ plied or fundamental projects. ACKNOWLEDGEMENTS We want to thank the following members of the Organizing Committee: Dr. Ballintijn, Dr. Van Overbeeke, Dr. Schoonhoven and especially Dr. Zandee for their invaluable activities in organizing, editing and indexing the separate book, containing the abstracts of posters and invited lectures. It was meant and proved to be the es­ sential basis for the 4 days of the Congress, August 31 to September 3, 1981 in Noordwijkerhout, The Netherlands. It is with great pleasure, that we remember the enthusiastic cooperation and at­ tention with increasing intensity during the past two years which resulted in the 3rd Congress of the European Society for Comparative Physiology and Biochemistry. The support and experience of Dr. Munday and Dr. Gilles our President and Executive officer of the Society were essential in the process of organization. Furthermore the incentive activities are gratefully acknowledged of the chairmen in the six Sections in the Congress: Dr. Dejours, Dr. De Ruiter, Dr. Beenakkers, Dr. Gilles, Dr. Laverack and Dr. Voogt. The Congress organization has had an explicit benefit of the accurate, helpfull and devoted activities of Trix van footen, Thea Laan, Danielle Zandee and Marc Spronk: not only the secretarial and financial administration before the Congress, but also their reception in a friendly manner at the Congress desk. For the final editing and indexing of the total manuscript of the Proceedings we are much obliged to Trix van Kboten for her expert typing activities. A.D.F. ADDINK and N. SPRONK Leiden, November 1981. The Origin of Physiology* A. D. F. Addink Department of Animal Physiology, Gorlaeus Laboratoria, Rijksuniversiteit, Wassenaarseweg 76, 2300 RA Leiden, The Netherlands INTRODUCTION I. THE ORIGIN OF PHYSIOLOGY AS A SCIENCE. Scientifically speaking you are, attending this European Congress in Holland, descendants of famous scientists in physiology and comparative physiology, such as just to mention a few: Galvani working on electricity and muscle movement in Italy and writing in Latin; Claude Bernard in France, who really introduced the experimental methodology in biology and the concept of the "milieu interieur"; Pavlov in Russia training dogs; Fick in Germany doing research on heart physiology and in Holland Jordan explored the muscle contraction (Postma 1980). Concerning biochemistry Priestley in England discovered oxygen and oxygen consumption in animals about 200 years ago; around 1935 Embden and Meyerhof established the glycolysis and fermentation, while Krebs postulated the citric acid cycle. Thereafter Florkin in Belgium is of importance for the comparative bio­ chemistry. As time does not permit me, I realize that you can extend this list much, much more, but what you will find is that for a long time -some 200 years- this all happened in Europe. Perhaps it is also for that reason, that we all enjoy the existence of this European Society for Comparative Physiology and Biochemistry. We hope that the lectures on fundamental and applied subjects, summarizing results of yesterday and today as well as the postersessions and round tables -organized along your personal interests- will inspire you for the research in the future. The day of tomorrow has a grey sky, but we do hope and believe, that through cooperation between research groups inside Europe a more favourable climate can be created, as far as scientific output is concerned. It is really possible to shape your own scientific future in physiology and compa­ rative biochemistry as Science is made by humans. II. THE ORIGIN OF PHYSIOLOGY AS A SCIENTIFIC PHENOMENON. Looking ahead for the scientific problems to be resolved such as the origin of life and its subsequent evolution into e.g. highly organized vertebrate animals * Dedicated to Prof. Dr H.J. Vonk. 1 2 A. D. F. Addink together with their physiological implications I will give a short speculative view on the future of animal physiology and comparative biochemistry to describe a living animal more than just a temporary adaptive self-reproducing battery of (sun)energy. If we will understand the course of events starting with organic molecules, follo­ wed by macromolecules and more complex forms of prebiotic situations such as com­ binations of nucleotides and proteins it will be instructive to unravel physiolo­ gical regulation mechanisms. In order to arrange nucleotides and proteins stereochemically a third factor is necessary. This third factor is basically the original part of the earth itself: the inorganic compounds more or less crystallized. So in a concise way I will point out two theories: at 1. the inorganic/organic interactions 2. the organic /macromolecular organization Ad 1. Experiments with silent discharges, ultraviolet or electric discharges (lightning), passing through a gaseous mixture of methane, ammonia, hydrogen and water vapour resulted in the formation of amino acids, as racemic mixtures (Miller 1960 ; Fox 1965, 1972 and Oparin 1957). In the presence of clays a preference for the formation of 1-amino acids was established. The formation of polymers from aminoacids was established by Degens and Matheja (1971) in the presence of inorganic templates such as kaolinite (Al, OH ). (Si.O ) and montmorillonate ((Al, Mg) OH ) (Si0 ). ((Na, Ca) n^O) . |U 4 1Q These clays serve the same function as the ribosomes in a biological system. As we know quite a number of enzymes contain (divalent) metalions in their active centers such as zinc, copper or iron. Also rather primitive organic catalysts have been described for instance an imidazole-zinc complex, which binds C0 just as the ? active site in the zinc containing enzyme carbonic anhydrase of 36000 daltons (Dobry-Duclaux 1966, 1970; Addink 1968). The aminoacid configuration of the apoprotein is such that the metals fit in a stable manner through coordination. In my opinion during prebiotic conditions the primary structure of such a protein was formed on zinc containing geochemical matrix -zinc is coordinated by histidin/imidazole-. This matrix is not just a clay mineral, but a silicate of the amphibole group (Deer et al 1963) including zinc silicate such as hemimorphite or calamine (Zn.Si 0 (OH) .HO). The amphibole silicates are composed of double chains of linked tetrahedras; (Si.O ) , repeating at 5.3 8. Octohedrally coordinated are arrays of five cations eg iron, magnesium, zinc, copper or calcium, possibly cobalt (indicated by MM). It seems to me not unlikely, that such minerals were involved in the stereochemical allignment of proteins, which catalyzed the nucleic acid polymerization. It is known that DNA-polymerase of Escherichia coli contains zinc. In the presence of sodium connected to the phosphate groups of the polynucleotide strand a double stranded macromolecule will be the result. Through adsorption of polynucleotides and proteins on these specific amphibole silicates and their coordinated metals it is most likely, that catalysis could take place, as presented in the scheme. This configuration resulted in the only basic molecular machinery in every cell: the universal genetic code. AMINO ACIDS PROTEINS GEOCHEMICAL ^- ^ TEMPLATE1 SILICATES-METALS -> NMUMC-LSEIOLPIRCOATTEEISN-SM M + MM-P^R O^T E_I NTSO NUCLEOTIDES POLYNUCLEOTIDES The Origin of Physiology 3 My conclusion is that the geochemical matrix (MM) has been the basic directing force towards both nucleotides and proteins, resulting finally in selfreproducing organisms. The molecular inorganic-organic interactions form in many physiological functions the regulatory control, which we are discussing during this congress. The original geochemical template (MM-silicates) is left outside the cell. Ad 2. The organization of the organic/macromolecular phase resulting into self- reproduction can be pointed out as follows. It is based on a theoretical approach of Manfred Eigen, who got a Nobel prize for his research on fast chemical reactions. This theory is called the HYPERCYCLE, which is meant to be "an explanation for the origin of life, or of selforganization of matter and evolution of biological macromolecules" to quote Eigen (1979; 1971). If we follow his line of reasoning we can depict several types of cycles. In fig. 1 the simple cycle, one in which the catalyst E forms a complex ES with substrate S. As the activation energy is lowered in the complex the substrate is converted to the product, which dissociates and the catalyst is back. The product might be a protein. I have modified the diagrams of Eigen to show the essential influence of the geochemical matrix in prebiotic conditions of amphibole silicates containing speci­ fic metals (MM); as indicated above nucleotidephosphate binding by Ca-Mg-Al-sili- cates as well as polypeptide adsorption or coordination (proteins and enzymes) by zincsilicates, including hemimorphite or calamine. In fig. 2 the catalytic cycle is composed of enzymes; these proteins form the catalytic unit in the following steps 1, 2, 3, 4, and n. Still the question could be asked even if we take into account the many years for the development of the first unicellular living creature, on what occasion did it start. A geochemical template is an answer to that question. Fig. 1 The common catalytic cycle of an enzyme E. The substrate is indicated by S; ES is the enzyme-substrate-complex; EP is the enzyme-product-complex and P stands for product. E is the enzyme, which may bind again a substrate mo­ lecule. MM stands for the geochemical adsorptive template containing also essential metals. Fig.2 The catalytic cycle is composed of several enzymes E , which catalyze the formation of the next enzyme from a substrate S. This cycle is a self- instructive unit, indicated by the symbol I, if it is fully active as an autocatalyst. MM stands for essential metals. 4 A. D. F. Addink * S* Fig. 3 The Hypercycle, an catalytic cycle consisting of selfinstructive units I , is capable of selfreproduction of the complete cycle, using also energy- rich substrates. MM stands for the essential metals, which are incorporated in proteins or nucleoproteins. Fig. 4 The Hypercycle in its essential appearance. MM stands for the essential me­ tals incorporated in the units. In Fig. 3 and 4 a hypercycle is built of catalytic cycles of the type of Fig. 2. These cycles are also selfinstructing for their own replication. They consist of proteins and ribonucleic acids. These autocatalytic units are able to reproduce themselves and catalyze also the next step! "I" stands for such informative units. In Fig.5 rearrangement of the diagram in Fig.4 is a biologically acceptable second degree hypercycle composed of nucleoprotein-catalysts, and enzymes after the translation. Now I will extend this theory to celldifferentiation and resul­ ting multicellular animals. You will need many more of these hypercycles to form such an organism, including its sensory elements directed towards the environment. The energy comes indirectly or directly from the sun. Fig. 5 The hypercycle of a second degree. I are selfinstructive information units and E are synthesized enzymes relevant for metabolism and translation { h) ^2 processes. MM stands for incor­ porated essential metals of geochemical origin. Fig. 6 Schematic second degree hypercycle with a complete set of characteristics I and E represented by sets of (wooden) bars; probably four coils are needed for a vertebrate organism. The axis indicates the energy supply via substrates. Essential metals are not indicated, (page 5) The Origin of Physiology 5 x^% Fig. 6 VAX Fig. 7 A fish diagram, after rearranging the second degree hypercycles of Fig. 6.