GATA4 and cancer: Inspection of individual genes that were either significantly increased or decreased in 5hmC density revealed genes implicated in normal pancreas development, for instance the transcription factors GATA4, GATA6, PROX1, ONECUT1/2, in addition to genes whose increased expression is implicated in cancer, such as YAP1, TEAD, PROX1, ONECUT2, ONECUT1 and IGF1. The relative 5hmC increase in transcription factor genes like GATA4, GATA6, PROX1, ONECUT1/2, MEIS2, which were previously reported to be involved in early pancreatic development29–31, suggest a reversion to a stem-like state in PDAC samples.