DFO Library MPO - Bibliothèque III 11 1 1111 11 1111 11 1 2039492 ECOSYSTEM THEORY FOR BIOLOGICAL OCEANOGRAPHY • le.. .M1114,1»1 :7,10 • Oillseate. a .1•• Ma/ •illaW IMO • «Ma Conférence SCOR Conference 1 — Richard Haedrich, 2 — Peter Allen, 3 — Victor Smetacek, 4 — Jordi Flos, 5 — Lynn Trainor, 6 — Lionel Johnson, 7 — Marcel Fréchette, 8 — Michael Conrad. 9 — Rene Ulloa, 10 — Gilberto Rodriguez, I I — Robert Ulanowicz, 12 — Louis Legendre. 13 — Clarice Yentsch, 14 — Marta Estrada. 15 — Serge Frontier, 16 — Shubha Sathyendranath, 17 — Serge Demers, 18 — Hironori Hirata, 19 — Patricio Bernai. 20 — Curt is Davis, 21 — Juan Tundisi, 22 — Michael Kemp, 23 — Yves De La Fontaine, 24 — Henri Atlan. 25 — James Kay, 26 — Pierre Lasserre, 27 — Ramon Margalef, 28 — Trevor Platt, 29 — Jacqueline McGlade, 30 — Donald Mikulecky, 31 — Robert Rosen, 32 — Peter Wangersky. 33 — Charles Yentsch. 34 — Kenneth Denman. 35 — David F. Smith, 36 — Isabelle Lamontagne, 37 — William Calder, 38 — Walter Boynton, 39 — John Field, 40 — Steven Cousins, 41 — William Silvert, 42— Claude Joins. 43 — William Leggett. 44 — Michael Fasham. 45 — Bernard Patten. 46 — Marion Lewis, 47 — Vincent Gallucci, 48 — Fredrik Wulff, 49 — Alexander Zotin. Not pictured: Jean - Yves Bellavance, Bruce Hannon. Patricia Lane, Jeffery Watson. /-/ ?2j 3 c.)c CANADIAN BULLETIN OF FISHERIES AND AQUATIC SCIENCES 213 C Ecosystem Theory for Biological Oceanography (Proceedings of a Symposium sponsored by SCOR, NSERC, NSF, UNESCO, and the U.S. Office of Naval Research held at Laval University, Ste. Foy, Quebec, March 16-23, 1984.) anS EDITED BY ,Fp so 1* ROBERT E. ULANOWICZ et des University oMaryl nd ç , A Chesapeake Biglogical Solomons, Maryland 20688. USA AND TREVOR PLAIT Department of Fisheries and Oceans Marine Ecology Laboratory Bedford Institute of Oceanography Dartmouth, Nova Scotia B2Y 4A2 DEPARTMENT OF FISHERIES AND OCEANS Ottawa 1985 The Canadian Bulletins of Fisheries and Aquatic Sciences are designed to interpret current knowledge in scientific fields pertinent to Canadian fisheries and aquatic environments. The Canadian Journal of Fisheries and Aquatic Sciences is published in annual volumes of monthly issues. Canadian Special Publications o`Fisheries• and Aquatic Sciences are issued period- ically. These series are available from authorized bookstore agents and other bookstores, or you may send your prepaid order to the Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Ont. K I A OS9. Make cheques or money orders payable in Canadian funds to the Receiver General for Canada. Scientific Information and Publications Branch J. Watson, Ph.D. Director and Editor-in-Chief Johanna M. Reinhart, M.Sc. Scientific Evaluation and Information Systems J. Camp /G. J. Neville / Susan D. Scale Editorial and Publishing Services Editorial Office: Department of Fisheries and Oceans Scientific Information and Publications Branch 200 Kent Street, 12th Floor Ottawa, Ontario, Canada K I A 0E6 Typesetter: K. G. Campbell Corporation, Ottawa, Ontario Printer: Kromar Printing Ltd., Winnipeg, Manitoba Cover Design: Ludvic Saleh & Associates, Ottawa, Ontario ©Minister of Supply and Services Canada 1985 Available from authorized bookstore agents, other bookstores or you may send your prepaid order to the Canadian Government Publishing Centre Supply and Services Canada, Ottawa, Ont. K I A OS9. Make cheques or money orders payable in Canadian funds to the Receiver General for Canada. A deposit copy of this publication is also available for reference in public libraries across Canada. Canada: $14.95 Cat. No. Fs 94-213E Other Countries: $17.95 ISBN 0-660-11900-5 ISSN 0706-6503 Price subject to change ivithout notice Correct citation for this publication: ULANOWICZ, R. E., AND T. PLATr [ED.]. 1985. Ecosystem theory for biological oceanography. Can. Bull. Fish. Aquat. Sci. 213: 260 p. Contents Abstract / Résumé vii—viii Background and Acknowledgements ix — x Guide to the Contents. Robert E. Ulanowicz and Trevor Platt xi—xiii I. Thermodynamics Ecology, Thermodynamics, and Self-Organization: Towards a New Understanding of Complexity. P. M. Allen 3-26 Thermodynamics and Growth of Organisms in Ecosystems. A. I. Zotin 27-37 IL Statistical Mechanics Statistical Mechanics in Biology — Applications to Ecology. L. E. H. Trainor 41-51 III. Ataxonomic Aggregations Structure of the Marine Ecosystem: Its Allometric Basis. Trevor Platt 55-64 Size and Metabolism in Natural Systems. William A. Calder III 65-75 The Trophic Continuum in Marine Ecosystems: Structure and Equations for a Predictive Model. S. H. Cousins 76-93 IV. Flow Analysis Ecosystem Flow Analysis. Bruce Hannon 97 — 118 Energy Cycling, Length of Food Chains, and Direct Versus Indirect Effects in Ecosystems. Bernard C. Patten 119-138 Flow Analysis of Materials in the Marine Euphotic Zone. M. J. R. Fasham 139-162 Network Thermodynamics in Biology and Ecology: An Introduction. Donald C. Mikulecky 163-175 V. Information Theory The Statistical Basis of Ecological Potentiality. Michael Conrad 179-186 Information Theory and Self-Organization in Ecosystems. H. Atlan 187 — 199 From Hydrodynamic Processes to Structure (Information) and from Information to Process. Ramon Margalef 200-220 Information and Complexity. Robert Rosen 221 — 233 VI. Working Group Reports 1. Hypothesis Testing and Sampling Design in Exploited Ecosystems. W. C. Leggett (Chairman) . 237-240 II. Ecosystem Theory in Relation to Unexploited Marine Ecosystems. J. G. Field (Chairman) . 241-247 III. Possible Holistic Approaches to the Study of Biological-Physical Interactions in the Oceans. L. Legendre (Chairman) . 248-253 IV. Technological Developments to Implement Theory into Biological Oceanography. K. Denman (Chairman) . 254-258 V. The Design of Large-Scale Cooperative Experiments. P. A. Bernal (Chairman) . .. 259-260 vi Abstract ULANOWICZ, R. E., AND T. PLATT [ED.]. 1985. Ecosystem theory for biological ocean- ography. Can. Bull. Fish Aquat. Sci. 213: 260 p. Holistic descriptions of marine ecosystems offer an alternative to characterizing biotic communities in ternis of coupled process models. Most quantitative narratives of whole ecosystem organization and development draw from five subject areas: Thermodynamics provides the phenomenological and macroscopic perspective necessary to perceive whole system behavior. Statistical mechanics is an historical example of how microscopic observations and hypotheses may be reconciled with their macroscopic counterparts. Ataxonomic aggregations of organisms, such as classifications according to particle size or metabolic rate, rely less upon "microscopic" (i.e., taxonomie) features of the ecosystem and could be more appropriate elements with which to build holistic theories in marine ecology. Flow analysis is quantitative theory germane to the study of how the parts of an ecosystem directly and indirectly affect each other within the context of the entire system. Lastly, information theory is a formalism capable of bridging and ultimately unifying the preceding four disciplines. Holistic considerations of ecosystem behavior tend, at this early stage, to be highly abstract. The existing theories, nonetheless, have practical implications for existing bio- logical programs. Various holistic hypotheses may be tested in both exploited and non- exploited marine ecosystems. Macroscopic concepts can promote a better understanding of biological—physical interactions and suggest the development of new technological instrumentation and methods. Finally, whole community descriptions beg for the design of new, large-scale, cooperative experiments in biological oceanography. vii Résumé ULANOWICZ, R. E., AND T. PLATT [ED.]. 1985. Ecosystem theory for biological ocean- ography. Can. Bull. Fish Aquat. Sci. 213: 260 p. Les descriptions holistiques des écosystèmes marins offrent une autre possibilité pour la caractérisation des communautés biotiques sous forme de modèles de pi•ocessus jumelés. La plupart des descriptions quantitatives de l'organisation et du développement d'un écosystème complet relèvent de cinq domaines: la thermodynamique fournit la perspective macroscopique et phénoménologique nécessaire à la perception du com- portement du système entier; la mécanique statistique constitue un exemple historique de la manière dont les observations microscopiques et les hypothèses peuvent être conciliées avec leurs analogues macroscopiques; les groupements non taxonomiques des organismes, comme la classification selon la taille des particules ou le taux métabolique, s'appuient moins sur les caractéristiques «microscopiques» (c.-à-d. taxonomiques) de l'écosystème et pourraient constituer des éléments plus appropriés pour l'élaboration de théories holis- tiques en écologie marine; l'analyse du déroulement est une théorie quantitative convenant à l'étude de l'interaction directe et indirecte ente les parties d'un écosystème dans le contexte de l'écosystème entier; finalement, la théorie de l'information est un formalisme capable de relier et, en fin de compte, d'unifier les quatre disciplines précédentes. Les considérations holistiques relatives au comportement de l'écosystème ont tendance, à ce stade, à être très abstraites. Néanmoins, les théories existantes ont des portées pratiques pour les programmes biologiques actuels. On peut vérifier diverses hypothèses holistiques dans des écosystèmes marins exploités et non exploités. Par ailleurs, les concepts macroscopiques peuvent encourager une meilleure compréhension des inter- actions biologiques-physiques