Wright’s Adaptive Landscape, Fisher’s Fundamental Theorem
Fitness maximization, or optimization, is a controversial idea in evolutionary biology. One classical formulation of this idea is that natural selection will tend to push a population up a peak in an adaptive landscape, as Sewall Wright first proposed. However, the hill-climbing property only obtains under particular conditions, and even then the ascent is not usually by the steepest route; this shows why it is misleading to assimilate the process of natural selection to a process of goal-directed choice. A different formulation of the idea of fitness-maximization is R. A. Fisher’s ‘fundamental theorem of natural selection’. However, the theorem points only to a weak sense in which selection is an optimizing process, for it requires that ‘environmental constancy’ be understood in a highly specific way. It does not vindicate the claim that natural selection has an intrinsic tendency to produce adaptation.