The fundamental theorem of natural selection

R. A. Fisher's Fundamental Theorem of Natural Selection states that the rate of increase in the mean fitness of a population ascribable to gene-frequency changes is exactly equal to the additive genetic variance in fitness. It has been widely misunderstood, though clarification has gradually come about particularly through the work of G. R. Price, W. J. Ewens, and S. Lessard. Building on their interpretations we here explain the approach adopted by Fisher (1941), devising a figure as an aid to understanding this important paper.