SPDEF and pancreatic intraductal papillary-mucinous neoplasm: LGD IPMN was characterized by the expression of transcription factors HOXB3 and ZNF117, Borderline IPMN showed high expression of SPDEF, NR4A1, and NR4A2; in contrast HGD Gastric IPMN was characterized by the high expression of NKX6-2 (Fig. 3A).