The Theory of Evolution by Natural Selection: A Hierarchical Expansion

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Abstract
Alleles, individuals, and species are all examples of entities possessing variation in the properties that underlie natural selection: branching (reproduction), persistence (survivorship), and heritability of characters. This suggests that the logic embodied in the theory of natural selection can be abstracted from its usual application to the level of individuals to encompass selection operating among any biological entities for which these essential properties can be meaningfully defined. This approach leads to a unified perspective of adaptation, selection, and fitness at all levels. Expanded versions of the Price covariance selection equations provide a convenient and useful conceptual vehicle for this discussion. The advantages of a hierarchical approach are twofold: it permits exploration of concepts and ideas across levels by analogy, and it focuses attention upon the mechanisms that account for different evolutionary dynamics at each level rather than obscuring these biologically unique properties with argument by extension from a single “special” level. We point out that the choice of a single measure of evolutionary change restricts the context in which “other level” processes will be perceived. We illustrate the limited forms in which higher and lower level selection can be recognized from the unique perspective provided by any given level through extensions of Price's formula. An exploration of the implications of such an approach leads us to the assertion that the development of a unified theory of evolution demands the recognition and incorporation of hierarchical structure as a conceptual foundation.

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