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Evolution__3rd_Edition

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..<br />

Parent–offspring similarity depends<br />

on the additive genetic variance<br />

Table 9.1<br />

The covariances between several different classes of relatives.<br />

Relatives Covariance<br />

1<br />

One parent and offspring VA 2<br />

1<br />

Mid-parent and offspring VA 2<br />

1<br />

Half sibs VA 4<br />

1 1<br />

Full sibs VA + VD 2 4<br />

1<br />

covOP =<br />

2<br />

1 /2 ∑ AD = 0 because A and D have been so defined as to be uncorrelated. If an individual<br />

has a big value of A, we know nothing about whether its value of D will be big<br />

or small. That leaves:<br />

cov OP =<br />

1<br />

A2+ AD<br />

2<br />

∑ ∑<br />

1 2 1<br />

∑ A = VA 2 2<br />

CHAPTER 9 / Quantitative Genetics 235<br />

In words, the covariance of an offspring and one of its parents is equal to half the<br />

additive genetic variance of the character in the population.<br />

The expression for the covariance between one parent and its offspring is true for<br />

each parent. It is a small step (though we shall not go into it here) to show that the same<br />

expression also gives the covariance between offspring and the midparental value: it is<br />

also 1 /2V A . Other expressions can be deduced, by similar arguments, for the covariance<br />

between other classes of relatives (Table 9.1). The formulae are useful for estimating the<br />

additive variances of real characters. However, the estimates become most interesting,<br />

for the evolutionary biologist, when expressed in terms of the statistic called heritability.<br />

9.6 Heritability is the proportion of phenotypic variance<br />

that is additive<br />

The similarity between relatives in general, and between parents and offspring in<br />

particular, is governed by the additive genetic variance of the character. If a character<br />

has no additive genetic variance in a population, it will not be inherited from parent to

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