Colocalization analysis was performed using the coloc R package42 for genes that were significant from the TWAS results of brain tissues with gene expression data from GTEx Release V8. The coloc.abf function was used to test for shared causal loci under four alternative hypotheses. Loci with posterior probability >90% were considered as strong evidence for the H4 hypothesis, i.e. both traits are associated and share a single causal variant.