We also computed module scores for IPMN signatures (as shown in Fig. 3B), which were derived from these markers associated with different types of IPMN (LGD IPMN signature: HOXB3, ZNF117, IGFBP3, GABRP, PDLIM3; Borderline IPMN signature: SPDEF, NRA4A1, NR4A2, DUSP1, PGC; HGD Gastric IPMN signature: NKX6-2, PSCA, SULT1C2, VSIG1; HGD Intestinal IPMN signature: REG4, SPINK4, CLCA1, RETNLB, ITLN1) that specifically associated with the IPMN type. Here, GABRP is linked to pancreatic intraductal papillary-mucinous neoplasm.