Gametic Disequilibrium Measures: Proceed With Caution

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Abstract
Five different measures of gametic disequilibrium in current use and a new one based on R. C. Lewontin's D′, are examined and compared. All of them, except the measure based on Lewontin's D′, are highly dependent upon allelic frequencies, including four measures that are normalized in some manner. In addition, the measures suggested by A. H. D. Brown, M. F. Feldman and E. Nevo, and T. Ohta can have negative values when there is maximum disequilibrium and have rates of decay in infinite populations that are a function of the initial gametic array. The variances were large for all the measures in samples taken from populations at equilibrium under neutrality, with the measure based on D′ having the lowest variance. In these samples, three of the measures were highly correlated, D2, D* (equal to the correlation coefficient when there are two alleles at each locus) and the measure X(2) of Brown et al. Using frequency-dependent measures may result in mistaken conclusions, a fact illustrated by discussion of studies inferring recombinational hot spots and the effects of population bottlenecks from disequilibrium values.