Furthermore, among these 19 depression genes, 7 did not have evidence for association with depression at the mRNA-level in the TWAS, including those not heritable and thus not included in the TWAS (Table 1, genes without asterisk). Interestingly, 4 of the these 7 were significant in the discovery PWAS and were replicated (CTNND1, PSMB4, P2RX7, and CACNA2D2, Table 2) suggesting that the PWAS contribute new insights into the pathogenesis of depression beyond the TWAS.