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Proponents of optimal foraging theory attempt to predict the behavior of animals while they are foraging; this theory is based on a number of assump tions ( 133 , 155 , 2 10, 23 1 ) . First, an individual's contribution to the next generation (i.e. its fitness) depends on its behavior while foraging. This contribution may be measured genetically or culturally as the proportion of an individual's genes or ideas, respectively, in the next generation. In the former case, the theory is simply an extension of Darwin's theory of evolution. Second, it is assumed that there should be a heritable component of foraging behavior, i.e. an animal that forages in a particular manner should be likely to have offspring that tend to forage in the same manner. This heritable compo nent can be either the actual foraging responses made by an animal or the rules by which an animal learns to make such responses. In other words, optimal foraging theory may apply regardless of whether the foraging behavior is learned or innate. Given these first two assumptions, it follows that the proportion of individuals in a population foraging in ways that enhance their fitness will tend to increase over time. Unless countervailed by sufficiently strong group selection (see 287, 242), foraging behavior will therefore evolve, and the average foraging behavior will increasingly come to be characterized by those characteristics that enhance individual fitness. The third assumption is that the relationship between foraging behavior and fitness is known. This relationship is usually referred to as the currency of fitness (23 1 ) . In general, any such currency will include a time scale, although in some cases it may be assumed that fitness is a function of some rate.
Graham H. Pyke (Thu,) studied this question.