Details of the design, methods, and participants of UKB have been described previously [18,19]. Briefly, participants aged 40 to 75 years identified through primary care lists were recruited across 22 assessment centres throughout the UK between 2006 and 2010. At recruitment, information was collected via a standardised questionnaire and selected physical measurements. Details of the data used from UKB are provided in S1 Text. Data were subsequently linked to Hospital Episode Statistics (HES), as well as national death and cancer registries. HES uses the International Classification of Diseases (ICD) 9th and 10th revisions to record diagnosis information, and the Office of Population, Censuses and Surveys: Classification of Interventions and Procedures, version 4 (OPCS-4), to code operative procedures. Death registries include deaths in the UK, with both primary and contributory causes of death coded according to ICD-10.

Genotyping was undertaken using a custom-built genome-wide array of approximately 826,000 markers [18,20]. Imputation to approximately 96 million markers was subsequently carried out using the Haplotype Reference Consortium and UK10K/1000 Genomes reference panels [20]. Clinical biochemistry markers, including total cholesterol, HDL cholesterol, and CRP, were measured at baseline in serum samples. Full details of the biochemistry sampling, handling and quality control protocol, and assay method have been provided previously [21].

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