The replication and combined meta-analyses are performed within METAL software66 using fixed effects inverse variance weighted meta-analysis (Supplementary Note). The combined meta-analysis of both the UKB discovery (N = 140,886) and GWAS replication meta-analysis (max N = 190,070) include a total maximum sample size of N = 330,956. For the exome combined meta-analysis, we synthesize data from the UKB discovery exome content (max N=140,866), with the replication dataset from both exome consortia (total max N=281,718), giving a maximum sample size of N=422,604.