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.

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Toward a metabolic theory of ecology

Body size is manifestly one of the most important attributes of an organism from an ecological and evolutionary point of view. Size has a predominant influence on an animal's energetic requirements, its potential for resource exploitation, and its susceptibility to natural enemies. A large literature now exists on how physiological, life history, and population parameters scale with body dimensions (24, 131). The ecological literature on species interactions and the structure of animal communities also stresses the importance of body size. Differences in body size are a major means by which species avoid direct overlap in resource use (153), and size-selective predation can be a primary organizing force in some communities (20, 70). Size thus imposes important constraints on the manner in which an organism interacts with its environment and influences the strength, type, and symmetry of interactions with other species (152, 207). Paradoxically, ecologists have virtually ignored the implications of these observations for interactions among species that exhibit size-distributed populations. For instance, it has been often suggested that competing species

https://www.jstor.org/stable/pdf/2096954.pdf

The ontogenetic niche and species interactions in size-structured populations

An understanding of insect ecology has been hampered by an inadequate knowledge of nutritional physiology. The effects of food quality upon physi­ ology and behavior (24, 102, 185, 285) and at the population and ecosystem levels (107, 209) are too often neglected. Nutritional ecology is central to proper interpretations of life history phenomena (e.g. manner of feeding, habitat selection, defense, and reproduction) both in ecological and evolu­ tionary time. Careful consideration of present and future information on this topic can lead to better predictive capabilities in both coevolutionary theory and pest management. In this review we assess the importance of food quality relative to other environmental factors and organism adaptations that influence post-inges­ tive food utilization and growth performance of immature arthropods. Reviewed elsewhere are pre-ingestive factors affecting food choice and con­ sumption (52, 73, 142, 154), nutritional requirements (67, 129, 226, 252), and the physiology of food consumption and utilization (19, 130, 270).

The nutritional ecology of immature insects

Many species are experiencing sustained environmental change mainly due to human activities. The unusual rate and extent of anthropogenic alterations of the environment may exceed the capacity of developmental, genetic, and demographic mechanisms that populations have evolved to deal with environmental change. To begin to understand the limits to population persistence, we present a simple evolutionary model for the critical rate of environmental change beyond which a population must decline and go extinct. We use this model to highlight the major determinants of extinction risk in a changing environment, and identify research needs for improved predictions based on projected changes in environmental variables. Two key parameters relating the environment to population biology have not yet received sufficient attention. Phenotypic plasticity, the direct influence of environment on the development of individual phenotypes, is increasingly considered an important component of phenotypic change in the wild and should be incorporated in models of population persistence. Environmental sensitivity of selection, the change in the optimum phenotype with the environment, still crucially needs empirical assessment. We use environmental tolerance curves and other examples of ecological and evolutionary responses to climate change to illustrate how these mechanistic approaches can be developed for predictive purposes.

/pdf/adaptation-plasticity-and-extinction-in-a-changing-1gcff0hfmv.pdf

Adaptation, plasticity, and extinction in a changing environment: towards a predictive theory.

▪ Abstract In large-herbivore populations, environmental variation and density dependence co-occur and have similar effects on various fitness components. Our review aims to quantify the temporal variability of fitness components and examine how that variability affects changes in population growth rates. Regardless of the source of variation, adult female survival shows little year-to-year variation [coefficient of variation (CV 30%). Old females show senescence in both survival and reproduction. These patterns of variation are independent of differences in body mass, taxonomic group, and ecological conditions. Differences in levels of maternal care may fine-tune the temporal variation of early survival. The immature stage, despite a low relative impact on population growth rate compared with the adult stage, ma...

https://www.jstor.org/stable/pdfplus/221737.pdf

Temporal Variation in Fitness Components and Population Dynamics of Large Herbivores

Patterns in life history phenomena may be demonstrated by examining wide ranges of body weight. Positive relationships exist between adult body size and the clutch size of poikilotherms, litter weight, neonate weight life span, maturation time and, for homeotherms at least, brood or gestation time. The complex of these factors reduces r
max in larger animals or, in more physiological terms, r
max is set by individual growth rate. Comparison of neonatal production with ingestion and assimilation suggests that larger mammals put proportionately less effort into reproduction. Declining parental investment and longer development times would result if neonatal weight is scaled allometrically to adult weight and neonatal growth rate to neonatal weight. Body size relations represent general ecological theries and therefore hold considerable promise in the development of predictive ecology.

S. Sams

Papers

Relationships between body size and some life history parameters.