. 2018 Aug 17;8:12309. doi: 10.1038/s41598-018-30089-2

Table 3.

Genotypic values and genotypic frequencies¹ in a two-locus, two-allele model with additive, dominance, and epistatic gene action.

A–locus genotype	f(A_iA_j)	B–locus genotype
		B ₁ B ₁	B ₁ B ₂	B ₂ B ₂
		f(B_kB_l)
		$q_{1}^{2}$	2q₁q₂	$q_{2}^{2}$
$A_{1} A_{1}$	$p_{1}^{2}$	$μ + a^{A} + a^{B} + a a$	$μ + a^{A} + d^{B} + a d$	$μ + a^{A} - a^{B} - a a$
$A_{1} A_{1}$	$p_{1}^{2}$	$p_{1}^{2} q_{1}^{2}$	$2 p_{1}^{2} q_{1} q_{2}$	$p_{1}^{2} q_{2}^{2}$
$A_{1} A_{2}$	$2 p_{1} p_{2}$	$μ + d^{A} + a^{B} + d a$	$μ + d^{A} + d^{B} + d d$	$μ + d^{A} - a^{B} - d a$
$A_{1} A_{2}$	$2 p_{1} p_{2}$	$2 p_{1} p_{2} q_{1}^{2}$	$4 p_{1} p_{2} q_{1} q_{2}$	$2 p_{1} p_{2} q_{2}^{2}$
$A_{2} A_{2}$	$p_{2}^{2}$	$μ - a^{A} + a^{B} - a a$	$μ - a^{A} + d^{B} - a d$	$μ - a^{A} - a^{B} + a a$
$A_{2} A_{2}$	$p_{2}^{2}$	$p_{2}^{2} q_{1}^{2}$	$2 p_{2}^{2} q_{1} q_{2}$	$p_{2}^{2} q_{2}^{2}$

Two locus genotypic frequencies were obtained by multiplication of marginal frequencies under linkage equilibrium⁶³.

μ: population mean; a: additive substitution effect; d: dominance deviation;aa, da, da and dd: additive × additive, additive × dominance, dominance × additive and dominance × dominance, gene actions respectively; p and q are major and minor allele frequencies.