“…The study of epistasis can be interesting because epistatic interactions aid understanding of the functional relations of genes involved in the trait under study and sometimes can compensate for a lack of additive genome-wide studies in quantitative traits (Visscher et al 2012(Visscher et al , 2017Lee et al 2016). Distinct genes in chickens control plumage color traits (Li et al 2019a,b) and the genes associated with plumage pigmentation, such as melanocortin 1 receptor (MC1R; Kerje et al 2003;Hoque et al 2013;Zhang et al 2013;Ran et al 2016;Tu et al 2019;Yang et al 2019), tyrosinase (TYR;Yang et al 2019;Zheng et al 2020), premelanosome protein (PMEL; Kerje et al 2004;Abolins-Abols et al 2018;Zheng et al 2020), melanophilin (MLPH;Vaez et al 2008;Bed'hom et al 2012), Agouti signaling protein (ASIP; Robic et al 2019;Yang et al 2019), SRY-box (SOX families; Harris et al 2010;Gunnarsson et al 2011), solute carrier family member 2 (SLC45A2; Gunnarsson et al 2007;Zheng et al 2020) and endothelin B2 receptor (EDNRB2; Kinoshita et al 2014;Li et al 2015;Wu et al 2017;Xi et al 2020), are well known. Accordingly, multiple responsible genes have been reported for plumage color variation, but studies have focused on single loci with a major effect, rather than effective modifier loci with small or epistatic effects that influence plumage color more delicately.…”