N

Assuming that total phenotypic variation is caused only by VA and VE and using the definition of the narrow-sense heritability, VA can be expressed as h2 2

V. =-— V • If h2 = 0.5, then VA in the numerator

on the right side of equation 9.27 can be replaced with VE. The final step was to utilize an estimate of the input of additive variation due to mutation available at the time that suggested V Putting this all together results in

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