For two-locus models in which the genotypic values were not functions of simple parameters, the genotypic values were entered as data, and V A and V G calculated as functions of the gene frequencies p and q. Bivariate numerical integration was undertaken using Simpson's rule by computing e.g. V A(p,q)f(p,q) over an (m+1) × (m+1) grid of equally spaced p and q values, taking m = 210 or higher power of 2 as necessary for adequate convergence. Results were computed for some models of metabolic pathways [38],[39] and for some published models obtained from QTL analysis [8].