Finally, for DMRs whose associated genes had increased expression upon treatment with FQI1, we observed enrichment of GO terms for pathways in cancer (e.g. PIAS4, SMAD3, TRAF4, MAP2K1) and RNA transport (e.g. NUP188, EIF3A, NUP35, RAN) in both hyper and hypomethylated DMRs (Supplementary Table S2). Here, SMAD3 is linked to cancer.