We then derived the τc coefficient26 (and associated P value) of the significantly enriched natural selection annotations (i.e., the background selection metric), This represents the enrichment of an annotation over and above the enrichment of all other annotations, which is a conservative approach, as most of the categories in our model are partially overlapping. To increase our power and for additional validation, we noted that LD Score allows testing of the full range of quantitative metrics, in an extension of the partitioned heritability framework. Results of this analysis are reported in Supplementary Table 8.