With respect to the single-variant associations observed, the strongest bioinformatics support from other studies was observed for rs9291211, identified in the European-ancestry GWAS meta-analysis of the OEcontrols phenotype. Although it reached only a suggestive GWS threshold (p=7.2×10–8), this variant was the leading signal in the significant BEND4 gene-based association and also showed strong regulatory effects on the brain-specific transcriptomic profile of BEND4 and SLC30A9. The function of BEND4 gene is unclear, but previous GWAS identified several variants at this locus (including rs9291211) that were associated with depression43, alcohol consumption36, autism spectrum disorder43, neuroticism35, height44, and helping behavior37. SLC30A9 encodes a zinc transporter involved in intracellular zinc homeostasis, which also plays a role in transcriptional activation of Wnt-responsive genes45. In previous GWAS, variants located in SLC30A9 gene (rs9291211 is located in BEND4 but also affects SLC30A9 gene expression) were associated with neuroticism44 and depression46. The phenome-wide scan of rs9291211 in the UK Biobank showed an effect of this SNP on a wide range of complex traits, some with an easy-to-conceptualize relationship to OD such as alcohol consumption, neuroticism, depression, and anxious
