We observed independent regions of association at 17 loci with P < 1 × 10−5 for FEV1 and 23 for FEV1/FVC (Figs. 1a,b and 2), including three regions (4q24 in GSTCD, 4q31 near HHIP and 15q23 in THSD4) that reached P < 5 × 10−8 in the stage 1 GWAS data alone, corresponding to a threshold of P < 0.05 after adjusting for 1 million independent tests12. SNP rs12504628, which was associated with both FEV1/FVC (P = 6.48 × 10−13; Fig. 2c and Table 2) and FEV1 (P = 1.50 × 10−10; Table 3), lies in an intergenic region upstream of HHIP and spanning ~300 kb at 4q31 that has been associated with lung function11, COPD11 and height9. Our top SNP rs12504628 was in strong linkage disequilibrium (LD; r2 = 0.97) with the previously reported SNP associated with lung function, rs13147758 (P = 5.30 × 10−10 for FEV1 and P = 1.11 × 10−12 1 for FEV1/FVC in our data), and with SNPs associated previously with height (rs6854783, r2 = 0.55; rs2055059, r2 = 0.48), suggesting a role in skeletal
