Large-scale GWAS efforts are rapidly elucidating the genetic architecture of polygenic traits, including anthropometrics [1, 2] and diseases [3–5], as well as behavioral and psychological outcomes [6–8]. These efforts have led to new biological insights, therapeutic targets, and polygenic scores (PGS), and help to understand the complex interplay between genes and environments in shaping individual outcomes [7, 9, 10]. However, GWAS results do not yet account for a large part of the estimated heritability [1, 2, 7, 8]. This dissonance, which is referred to as the ‘missing heritability’, has received broad attention [11–17].
