Ecology is that branch of science that aims at explaining the relationships between organisms and their physical and biotic environments. Model building is generally used in ecology as the ftrst step towards meeting this goal. The model should be a simplified representation of reality that provides insight into the structure and function of often complex systems (49). Of the many models applied to ecological problems, those based on the principle of optimization, borrowed from economics, have frequently been used (88) . In economic models, a dependent variable (money) is maximized, using some cost-beneftt function. Ecologists' reliance on the optimality construct is related to the fact that its structure is similar to the evolutionary biologist's view of the mechanics of evolutionary processes-that natural selection, operating on phenotypic variance, drives populations towards optimal, fine-tuned responses to environmental variables. Instead of money, the dependent variable that will be maximized here is Darwinian fitness. The methodology and problems associated with the application of optimality theory to ecological problems have received extensive treatment in several reviews (16, 52, 62, 77, 76). Despite some criticisms (see 88) , many ecologists think that optimization models have been, and are likely to remain, the principal conceptual framework for thinking about evolutionary trends at the phenotypic level (78).
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Riechert et al. (1983) studied this question.
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