Of these gene-linked cCREs, 72.2% were located at least 2 Kb away from annotated promoter regions of protein-coding genes, a proportion supported by CRISPR perturbations of enhancer-promoter connections.27 Notably, the linked genes were enriched for previously identified AD seed genes28 (Fisher’s exact test P = 0.0033), including APOE, TGFB1, PICALM and APP (Figure S6 and Table S3). Here, PICALM is linked to Alzheimer disease.