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Exaptation is a term used in evolutionary theory to supplement and contrast with the concept of adaptation. It means that a particular feature or behavior has evolved indirectly or secondarily rather than as a primary consequence of evolutionary adaptive pressures related to current functions. Exaptations are features or behaviors with biological functions that are different today than they were when they originally evolved. It is difficult to determine whether any given feature or behavior is an adaptation or an exaptation, because many conditions of evolutionary change are unknowable. However, it stands to reason that certain features and functions provide current benefits that differ substantially from the initial benefits exploited by the process of evolution by natural selection.

The term exaptation was first introduced by Stephen Gould and Elisabeth Vrba in 1982 to address this type of repurposing of adaptations in biology, to distinguish those features originally formed by selective pressures, termed “historical genesis,” from later uses of those features, termed “current utility.” Gould and others subsequently expanded this principle to sociobiology and evolutionary psychology. As pointed out by those authors, the concept of natural selection does not rule out the later repurposing of features that originally provided a benefit, and thus greater clarity accrues from distinguishing historical genesis for current utility. Neither does natural selection preclude the emergence of features that have a neutral impact on heredity. In both cases, the explanation for such emergence may lie outside the scope of immediate selective pressures. Exaptation as a concept addresses the former condition; the latter remains largely unexplored. Much debate has been spent in attempting to describe musical capacities and behaviors as adaptations, exaptations, or neutral developments.

Spandrels and Emergentism

Some forms of flight appear to have arisen through co-option or readaptation of the lift afforded by feathers, which most scientists believe originally provided a selective advantage because of their insulating qualities that aided body temperature regulation. A metaphor that is sometimes used to explain exaptation is that of a spandrel, a term borrowed from architecture, which describes the v-shaped space between two archways. The shape of a spandrel is not the direct intention of architectural plans but rather the result of placing two archways adjacent to each other. Architecture led directly to the development of arches as a feature in the construction of buildings. Spandrels were simply a by-product of this process.

Spandrels have a correlate in nature, in the principle of emergentism. A feature or characteristic may exist in the world, or emerge during evolution (phylogenesis) or cognitive development (ontogenesis), that itself is not a direct consequence of evolutionary pressures but rather the outcome of the conditions surrounding its emergence. Think of the flow of water following the path of least resistance. Water exhibits no intentionality but is simply subject to the resistances that are presented by the landscape. The shape of honeycomb cells in beehives is hexagonal, not because bees have adapted a hexagon-making gene but because the hexagonal shape completely tiles in two-dimensional space. Thus, hexagon-shaped containers in three dimensions provide greater structural integrity than round cylinders do, as the surface tensions in hexagonal cells are evenly distributed along a maximum area. Whether the bee intends to produce this shape or the roundness of cylindrical cells simply deforms under pressure into hexagons, the emergent nature of stacked hexagonal containers is the consequence of the laws of physics rather than natural selection.

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