Putative novel signals of association from stage 1 were followed up in three independent sets of samples (stage 2): (i) an independent subset of UK Biobank participants (UK Biobank, n=49,727), (ii) a population-based consortium (SpiroMeta, n=38,199)15 and (iii) UK Households Longitudinal Study (UKHLS, n=7,449). We did not include these studies in Stage 1 as: (ii) was to be utilised for independent replication and; (i) and (iii) were not yet available when Stage 1 was undertaken. Each signal was followed-up only for the trait with which it was most strongly associated in Stage 1. The first tranche of genotype data and imputation output (merged 1000 Genomes Project Phase 3 and UK10K imputation panel) from UK Biobank was released May 2015 (see URLs) and comprised the 49,979 individuals originally genotyped for UK BiLEVE (an unrelated subset of 48,943 of which were used as discovery in this study) and an additional 102,757 individuals selected at random from the entire UK Biobank. From these 102,757 individuals, we initially selected 51,117 samples that had lung function measurements (FEV1 and FVC) meeting ATS/ERS criteria and had