Adaptive Evolution
Adaptive evolution—phenotypic improvement due to selection—is the central theme of Darwinian evolutionary biology. The concept of adaptive evolution by selection on heritable variation underlies every evolutionary analysis of form, function, and fitness. How biologists view adaptive evolution— how they are taught to view it—has a profound influence on thinking and research in virtually every area of biology. Despite the importance of adaptive evolution, there is still controversy over how to relate development and selection in discussions of adaptive design (e.g. see Charlesworth, 1990; Amundson, 1996). It is a controversy that began in the late nineteenth century, when biologists unfortunately began to dichotomize development and selection, as if they were opposing factors in evolution (see the discussion of gradualism in chapter 24). The modern version of this debate dichotomizes selection and developmental constraints (e.g., see Maynard Smith et al., 1985; Bell, 1989; Ridley, 1993; Amundson, 1996; see also chapter 1). Phylogenetic constraint is another version of developmental constraint, since it refers to limits imposed by ancestry (inherited developmental patterns) on current form as does the idea of developmental constraint. The dichotomy between selection and development, as if they were opposed factors in adaptive evolution, is misconceived. Adaptive evolution is a two-step process: first the generation of variation by development, then the screening of that variation by selection (Mayr, 1962; Endler and McLellan, 1988, p. 395). If an established trait (one widespread in a population) persists through phylogenetic branching events, this is not evidence against selection as an explanation for the trait (cf. Coddington, 1988), as if speciation reduces the importance of selection. Rather, it could be taken as evidence that selection is important for maintenance of the trait in more than one lineage: trait persistence over long time spans certainly does not represent absence of continued selection, since it is known that traits no longer favored by selection, such as the eyes of cave animals and walking limbs in whales, are often lost (for other classic examples, see Rensch, 1960).