SMAD3 and keratoconus: These included SNPs at ZNF469, FOXO1, MPDZ, SMAD3, and COL5A1, consistent with previous findings14 and with the expected direction of effect; but also SNPs at novel CCT-loci: LOX and FST. Further, an additional 18 were associated with keratoconus at a nominal level (P < 0.05).