Colocalization analysis suggested campesterol and gallstones share the same causal variant in this region (RCP = 0.65; Fig. 5a) and nominated rs6544713 (SCP = 0.45) as the most likely causal variant. rs6544713 resides in active regulatory units in intestinal tissue45 and its campesterol-decreasing allele is associated with higher ABCG8 expression in colon tissue46 (β = 0.37, P = 7.2 × 10−16), but not in the liver46. The gene discussed is ABCG8; the disease is gallstones.