By contrast, genes that were upregulated and hypomethylated included mucins like MUC1; SERPINA1, a serine protease inhibitor that acts in the liver and lung; HNF1B, a transcription factor that regulates in renal and pancreatic development; and ME3, a mitochondrial malic enzyme whose deletion was shown recently to confer lethality in SMAD4-deleted pancreatic adenocarcinoma [24]. Here, SMAD4 is linked to pancreatic adenocarcinoma.