Biological adaptation in light of the Lewontin-Williams (a)symmetry

Neo-Darwinism characterizes biological adaptation as a one-sided process, in which organisms adapt to their environment but not vice versa. This asymmetric relationship-here called Williams' asymmetry-is called into question by Niche Construction Theory, which emphasizes that organisms and their environments often mutually affect each other. Here, we clarify that Williams' asymmetry is specifically concerned with (quasi)-directed modifications toward phenotypes that increase individual fitness. This directedness-which drives the adaptive fit between organism and environment-entails far more than the mere presence of cause-effect relationships. We argue that difficulties with invoking fitness as the guiding principle of adaptive evolution are resolved with an appropriate definition of fitness and that objections against Williams' asymmetry reflect confusions about the nature of biological adaptation.