GPRIN3 and chronic obstructive pulmonary disease: In lung tissue, we identified two genome-wide significant genes, GPRIN3 in the 4q22 locus (P = 7.43 × 10− 7) and WNT3 in the 17q21 locus (P = 1.24 × 10− 6); the latter locus was not identified in the single variant GWAS of severe COPD (Fig. 3).