Natural Selection

The theory that organisms become adapted to their environment through the process of natural selection has become so ingrained in modern biological thought, and more generally in Western culture of the late 20th century, that it is surely one of the great scientific paradigms of the present era. Evolution and adaptation were both well-accepted concepts by the mid-19th century, at least among French and British natural philosophers. The theory of natural selection, developed by Wallace (1858) and Darwin (1859), provided a functional connection between the two processes. However, despite its logical consistency, natural selection was not accepted as a necessary or sufficient explanation for adaptation until the “evolutionary synthesis” of the mid-20th century, when knowledge from population and quantitative genetics, natural history (e.g., biogeography, ecology, behavior), systematics, and paleontology merged to form the unified theory of adaptive evolution known as neo-Darwinism (see Futuyma 1998 for a concise review of this history). Since that time, natural selection has been accepted as the universal mechanism leading to adaptation, and the two terms have become so closely associated as to be almost tautological. Adaptationist hypotheses are now fundamental to much of modern biology and are becoming increasingly apparent in more disparate fields, such as anthropology, medicine, biochemistry, and psychology (Futuyma 1999). Nevertheless, there is much that natural selection cannot explain. For example, chance events may strongly influence macroevolutionary trends (i.e., the origin and extinction of species and higher taxa), some aspects of molecular evolution, and evolution within small or subdivided populations (Mazer and Damuth, this volume, chapter 2; Nunny, this volume). For this reason, adaptationist hypotheses should be viewed with skepticism until adequately tested (Reznick and Travis, this volume). In this chapter, we carefully define natural selection and discuss methods of measuring selection in natural populations as a means of testing adaptationist hypotheses. These methods are most appropriate for testing hypotheses concerning the adaptive significance of contemporary trait distributions within and among populations (“microevolutionary” hypotheses) and thus have particular relevance for evolutionary ecologists. Readers will find many additional examples of these and other methods of testing microevolutionary adaptationist hypotheses throughout this volume, such tests being an essential component of most research programs in evolutionary ecology.