This connection was brought into even sharper relief by two recent GWAS studies of primary sclerosing cholangitis (PSC) and inflammatory bowel disease (IBD), which both identified HDAC7 among the disease-associated loci, and also individually its immediate upstream kinases PKD and SIK2, as well as two isoforms of PKC that are upstream of PKD (Figure 6A) (Liu et al., 2013; Jostins et al., 2012). The gene discussed is PRKD1; the disease is pancreatic serous cystadenoma.