GCTA, with P-values for difference from zero computed using a likelihood ratio test. 20 principal components (PCs) were always included as fixed-effects to account for ancestry. Genetic correlation between traits in the GWAS cohort was estimated from all post-QC SNPs in the full set of 10,000 individuals with GEMMA31 (Supplementary Table 15). For the YFS, we quantified the mediating effects of trait on cis-expression by separately re-estimating cis-h2g with all analyzed traits (height, BMI, TC, TG, HDL, LDL) included as fixed-effects in addition to PCs. We did not observe significant differences in any individual gene (after accounting for 3,836 genes tested) nor in the mean estimate of cis-h2g.