was run with PLINK (ref. 55) using logistic regression adjusted for age, sex and the first three principal components as calculated by EIGENSTRAT using ancestry informative markers. Analysis was performed assuming an additive genetic model. These data were aggregated and analysed using Perl scripts and the R statistical package. A total of 1501 phenotypes were considered, for a per-SNP Bonferroni correction of 0.05/1501=3.3 × 10−5.