PFN3 and Wilson disease: Genes near the 10 DMRs with overlapping locations and common direction in blood and liver included CSAD, ITGB7, LSM12, PSMD9, RRN3P2, SNX29P2, and PFN3. In particular, a liver DMR in an intron of HDAC5 and upstream of LSM12 was specifically and significantly hypermethylated in WD liver and blood (Additional file 1: Fig. S8b–d).