In GWAS meta-analysis of OEcontrols in the European-ancestry cohort, we observed a gene-based association for the BEND4 locus that was GWS in the gene-based test (p=9.9×10−6; Table 1; Supplementary Figure 1B). In the BEND4 gene region, we identified a genetic association that nearly reached GWS: rs9291211 on chromosome 4 (z=−5.38, p=7.2×10–8; Figure 1B; Table 1). With respect to this locus, no heterogeneity was observed among the cohorts (heterogeneity: I2=0, p=0.879; Supplementary Table 6). This variant (or LD proxies in the same ancestry group) was identified in previous GWAS of behavioral traits: alcohol consumption (rs4501255, LD proxy r2=0.94, p=5×10–10)36; neuroticism (rs9291211, p=2×10–8)35; and helping behavior (rs2880666, LD-proxy r2=0.77, p=5×10–7)37. Additionally, rs9291211 is an eQTL for SLC30A9 and BEND4 in multiple tissues (GTEx multi-tissue eQTL p = 1.2×10−26 and 2.88×10−9, respectively). The rs9291211×SLC30A9 eQTL (i.e., rs9291211 regulating the SLC30A9 expression) showed a posterior probability>90% in seven brain tissues (amygdala m=0.99; anterior cingulate cortex m=1; caudate m=1; cortex m=0.99; hypothalamus m=1; nucleus accumbens m=1, putamen m=0.99; Supplementary Figure 2A). The rs9291211×BEND4 eQTL showed posterior probabilities>90% in two brain tissues (caudate m=0.9; cortex m=1;
