Directional selection produces a change in the population mean of a phenotype, but this shift will only be translated into a phenotypic difference between generations if the trait is heritable, in the sense that there is additive genetic variance in the trait. Thus any study of evolutionary change must concern itself both with the factors causing phenotypic change and the genetic architecture of the traits under investigation. In the case of a single trait the relevant genetic parameter is the narrow sense heritability, de®ned as the ratio of the additive genetic variance to the phenotypic variance (Falconer, 1989). Heritability can be estimated by pedigree analysis (e.g. full-sibs, half-sibs) or by response to selection, the latter estimate frequently being called the realized heritability. Estimation of the heritability from pedigree analysis is the more generally adopted method, because it is typically both less labour-intensive and quicker, requiring only one or two generations of breeding. A selection experiment is not simply an alternative method of estimating heritability but can provide insights into the genetic architecture of a trait that are not evident from the sib analysis. For example, according to the standard response equation (Response Heritability