Ronald A. Fisher, John B. S. Haldane and Sewall Wright - which showed that mutation and natural selection together cause adaptive evolution.

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Fitness- Average number of offspring produced by a genotype and divide it by the mean number of offspring produced by the most prolific genotype

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Average fitness

In a species, only 1 of every 1000 seeds survives to reproductive age, and that those that survive produce an average of 3000 seeds. 

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Selection coefficient

A related variable is the selection coefficient (s), which is the relative intensity of selection against a  genotype

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The selection gradient measures the strength of directional selection acting on a quantitative trait.

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