Further SMR analysis treating methylation as the exposure and expression as the outcome (P-SMR < 0.05, P-SMR_multi < 0.05, P-HEIDI > 0.01) identified 3 genes (ATG16L1, MAP3K14, PMVK) where specific CpG methylation levels appeared to causally influence gene expression levels (Table 1; Supplementary Table S5), and importantly, both the CpG site and the gene’s expression were associated with odds of preeclampsia in the primary analyses. The gene discussed is ATG16L1; the disease is preeclampsia.