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Brown 2004 Ecology Towards a Metabolic Theory of Ecology

Brown 2004 Ecology Towards a Metabolic Theory of Ecology -...

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1771 Ecology, 85(7), 2004, pp. 1771–1789 q 2004 by the Ecological Society of America TOWARD A METABOLIC THEORY OF ECOLOGY J AMES H. B ROWN , 1,2,4 with J AMES F. G ILLOOLY , 1 A NDREW P. A LLEN , 1 V AN M. S AVAGE , 2,3 AND G EOFFREY B. W EST 2,3 1 Department of Biology, University of New Mexico, Albuquerque, New Mexico 87131 USA 2 Santa Fe Institute, 1399 Hyde Park Road, Santa Fe, New Mexico 87501 USA 3 Theoretical Division, MS B285, Los Alamos National Laboratory, Los Alamos, New Mexico 87545 USA J AMES H. B ROWN , MacArthur Award Recipient, 2002 Abstract. Metabolism provides a basis for using first principles of physics, chemistry, and biology to link the biology of individual organisms to the ecology of populations, communities, and ecosystems. Metabolic rate, the rate at which organisms take up, transform, and expend energy and materials, is the most fundamental biological rate. We have developed a quantitative theory for how metabolic rate varies with body size and temperature. Metabolic theory predicts how metabolic rate, by setting the rates of resource uptake from the environment and resource allocation to survival, growth, and reproduction, controls ecological processes at all levels of organization from individuals to the biosphere. Examples include: (1) life history attributes, including devel- opment rate, mortality rate, age at maturity, life span, and population growth rate; (2) population interactions, including carrying capacity, rates of competition and predation, and patterns of species diversity; and (3) ecosystem processes, including rates of biomass production and respiration and patterns of trophic dynamics. Data compiled from the ecological literature strongly support the theoretical predictions. Even- tually, metabolic theory may provide a conceptual foundation for much of ecology, just as genetic theory provides a foundation for much of evolutionary biology. Key words: allometry; biogeochemical cycles; body size; development; ecological interactions; ecological theory; metabolism; population growth; production; stoichiometry; temperature; trophic dynamics. 4 E-mail: [email protected]
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1772 JAMES H. BROWN ET AL. Ecology, Vol. 85, No. 7 Perspectives I NTRODUCTION The complex, spatially and temporally varying struc- tures and dynamics of ecological systems are largely consequences of biological metabolism. Wherever they occur, organisms transform energy to power their own activities, convert materials into uniquely organic forms, and thereby create a distinctive biological, chemical, and physical environment. Metabolism is the biological processing of energy and materials. Organisms take up energetic and ma- terial resources from the environment, convert them into other forms within their bodies, allocate them to the fitness-enhancing processes of survival, growth, and reproduction, and excrete altered forms back into the environment. Metabolism therefore determines the demands that organisms place on their environment for all resources, and simultaneously sets powerful con- straints on allocation of resources to all components of fitness. The overall rate of these processes, the meta-
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