Meta-analysis was carried out across all 53 cohorts using METAL83. Cochran’s Q test for heterogeneity was implemented in METAL83,84, and a heterogeneity p-value threshold of p > 1 × 10−3 applied to results. A conservative significance threshold was applied to these data, correcting for the total number of genes tested across all tissues (121,611 gene-region tests in total). This resulted in a genome-wide significance threshold of 4.1 × 10-7.