Notably, the neural development-related TFs, which showed consistent promoter hypermethylation in cancer cells of all patients (Fig. 2d and Supplementary Table S2), also enriched binding motifs in the normal cells, such as ASCL1, CUX1/2, DLX1/2/3/5, NEUROD1, NKX6-1, NEUROG2 (Fig. 5c and Supplementary Fig. S15c, Table S3), suggesting that dysregulation of these TFs may be involved in tumorigenesis of PDAC. Here, NKX6-1 is linked to cancer.