The gene-based analyses found 41 genes to be significantly associated with alcohol consumption (Supplementary Table 4). The top hits were genes identified in the single SNP analyses (KLB, CADM2, GCKR) and genes in the 4q region were also found to be significantly associated with alcohol consumption at the gene level (ADH1C, C4orf17). Among the 41 loci associated at the gene level, genes of interest include the dopamine receptor D2 (DRD2) gene, previously associated with addiction phenotypes,40 chromodomain Y-like (CDYL) protein previously associated with alcohol consumption, cAMP-specific 3′,5′-cyclic phosphodiesterase 4B (PDE4B) associated with alcohol preference in rodents,41, 42 zinc-finger protein 512 (ZNF512) associated with oral cavity cancer43 and protein phosphatase 1G (PPM1G) found to be hypermethylated in individuals with AUD.44
