Epistasis hypothesis of heterosis was proposed by:
Epistasis hypothesis proposes that interactions among alleles at different loci contribute substantially to heterosis beyond single-locus dominance or overdominance. Favorable combinations of interacting genes present in parental lines act synergistically in hybrid background. Gowen in early 1950s argued that recombination of coadapted gene blocks disrupted in inbreds reconstituted in F1 enhancing regulatory networks involving auxin, cytokinin and brassinosteroid cross-talk, carbohydrate metabolism and stress signaling pathways. Modern QTL mapping and genome-wide epistasis scans in Arabidopsis
Ref: Gowen JW 1952 Heterosis epistasis; Melchinger Genetics epistatic contribution; PubMed Epistatic QTL rice heterosis analysis.