Natural Selection, Variation, Adaptation, and Evolution: a Primer of Interrelated Concepts

Abstract

Natural selection is an elegantly simple concept but one that can manifest in complex ways. I review how the basic model of single-trait viability selection has been extended to more complex forms of selection on multiple traits and on reaction norms. Fitness is defined as the expected lifetime reproductive success for individuals with a given genotype or phenotype over a given range of environments. Since the reproductive success realized by any individual will include a stochastic departure from this expectation, selection is therefore a consistent difference in fitness between organisms with different characteristics. A clear distinction is drawn between selection, which can act on any phenotypic difference, and the response to selection, which can occur only if phenotypic differences are heritable. This distinction separates the action of natural selection in filtering variation from the origin of the novel variants on which selection acts. Since selection frequently acts on standing genetic variation or on conditionally neutral variation, both of which accumulate in populations before the imposition of selection, such variation accumulates independently of its fitness effects under the subsequent selection regime. Recent discussions of "Lamarckian" inheritance must be carefully circumscribed to avoid the implication of directed mutation, for which there is no evidence.