Equation 3 · From Origins to Frontier: A History of Evolutionary Biology and Ecology
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for a diploid population starting at allele frequency p . This single expression carries a real consequence: small populations can fix or lose alleles by chance alone, independent of whether those alleles are beneficial, neutral, or mildly deleterious, and the smaller the effective population, the faster that random fixation proceeds. Wright’s “shifting balance” framework, in which small subpopulations explore a rugged fitness landscape partly by drift and partly by selection, remains debated in its strong form, but the underlying mathematics of drift is uncontroversial and became a permanent part of population genetics.
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