Natural selection and adaptation

Students of the natural world have long remarked on the fact that animals and plants are well suited to the demands of their environments. ‘Adaptation’, as it is used in modern biology, can name both the process by which organisms acquire this functional match, and the products of that process. Eyes, wings, beaks, camouflaging skin pigmentation and so forth, are all ‘adaptations’ in this second sense. Modern biological orthodoxy follows Darwin in giving a central role to natural selection in explaining the production of adaptations such as these. This much is uncontroversial. But a number of more contentious conceptual questions are raised when we look in more detail at the relationship between natural selection and adaptation. One of these questions concerns how we should define adaptation. It is tempting to characterize adaptations as functional traits – eyes are for seeing, large beaks are for cracking tough seed-casings. This in turn has led many commentators in biology and philosophy to define adaptations as those traits which have been shaped by natural selection for their respective tasks. Others – especially biologists – have complained that such a definition trivializes Darwin’s claim that natural selection explains adaptation. This claim was meant to be an important discovery, not a definitional consequence of the meaning of ‘adaptation’. These worries naturally lead on to the issues of how natural selection itself is to be understood, how it is meant to explain adaptation, and how it should be distinguished from other important evolutionary processes. These topics have a historical dimension: is Darwin’s understanding of natural selection, and its relationship to adaptation, the same as that of today’s evolutionary biology? Textbook presentations often say yes, and this is surely legitimate if we make the comparison in broad terms. But differences emerge when we look in more detail. Darwin, for example, seems to make the ‘struggle for existence’ an essential element of natural selection. It is not clear whether this is the case in modern presentations. And Darwin’s presentation is largely neutral on the inheritance mechanism that accounts for parent/offspring resemblance, while modern presentations sometimes insist that natural selection always implies a genetic underpinning to inheritance.