We then compared the list of 43 motifs to α- and δ-cell motif enrichment and found only 5 were unique to β-cells (Fig. 6m, Supplementary Data 18): BACH2, a risk gene for T1D69; SMAD5, a protein downstream of TGF-β signaling and can autonomously promote glycolysis70; HMBOX1, a regulator of telomerase activity71; TP53, a gene connected T1D and T2D by TP53-mediated apoptosis72; and JUND, a gene associated with β-cell dysfunction73. Here, SMAD5 is linked to type 2 diabetes mellitus.