The SNPTEST software program [51] was used to perform genome-wide association testing in the GEMS and CoLaus cohorts, while the Merlin software package [55] was used to perform association testing in the TwinsUK cohort. The meta-analysis of the discovery phase cohorts (CoLaus, TwinsUK and GEMS) was performed using Liptak-Stouffer's method for combination of independent tests, where P-values are converted to Z-scores by a standard normal curve and weighted by each study's sample size [56].