Heritability is the proportion of phenotypic variation due to additive genetic factors24; we therefore fitted a model in which SNPs have additive effects. Non-additive genetic variation and variation due to gene-environment interactions may exist, but they are not part of the missing heritability because they do not contribute to the heritability. Epigenetic mutations may cause resemblance between relatives and contribute to heritability if stably inherited, but in that case they would be equivalent to DNA sequence variants, would show LD with the assayed SNPs and would not contribute to missing heritability25.
