mixed model test implemented in SAIGE114 with birth year and the top four principal components (computed from the white British subset) as covariates. For the pLOF burden tests, for each autosomal gene with at least two rare pLOF variants (n = 12,052 genes), a burden variable was created in which dosages of rare pLOF variants were summed for each individual. This sum of dosages was tested for association with the 1,419 traits using SAIGE. The same covariates used in the single-variant tests were included. For both the single-variant and the burden tests, we used 5 × 10−8 as the genome-wide significance threshold.
