Furthermore, among the 23 TFs showing TFBS enrichment, we found reduced gene expression of four TFs, namely FOXA1, NR3C1, HNF4A, and FOSL2, whose downregulation was also significantly associated with increased methylation levels at enhancer DMCpGs in PD (Fig 5B–D). The gene discussed is HNF4A; the disease is Parkinson disease.