where
is the proportion of genes contributed by ancestor i to the current population and
is the
proportion of founder i's genes that are retained in the current population. Like
,
accounts for
unequal founder contributions. Unlike
,
also accounts for the fraction of founder genomes lost from the
pedigree through drift during bottlenecks. Although
is the more accurate description of the amount of founder variation
present in a population, it can only be calculated directly for simple pedigrees. For large or complex pedigrees, the number of
founder genome equivalents must be approximated based on computer simulation of a large number of segregations through the
pedigree. This is done by assigning each founder a unique pair of alleles and randomly transmitting those alleles through the
pedigree [MacCluer, VandeBerg, Read, and RyderMacCluer
et al.1986]. The number of founder genome equivalents is similar to the effective founder number, but the former
has been devalued based on the proportion of its genome that has probably been lost to drift [LacyLacy1989].