This included variants at nine loci with known genome-wide significant T2D association (PROX1, ST6GAL1, SMARCAD1, XKR6, INS-IGF2, HMGA2, SMEK1, HMG20A, and LAMA1), as well as at two previously unreported loci with sub-genome-wide significant association, CRB2 and PGM1. To identify candidate target genes of the T2D-associated PSSE in pancreatic progenitors, we analyzed the expression of all genes within the same topologically associated domain (TAD) as the PSSE in PP2 cells and in primary human embryonic pancreas tissue (Figure 3B and Figure 3—figure supplement 1A). This evidence concerns the gene SMARCAD1 and type 2 diabetes mellitus.