AD risk variants were significantly enriched at ATAC-seq peaks containing motifs for RUNX (z score = 5.22, corrected P < 1.9 × 10− 3), SPI1 (PU.1) (z score = 5.02, corrected P < 1.9 × 10− 3) and Spdef (z score = 3.80, corrected P = 0.027) after correction for multiple testing using the Bonferroni method (Fig. 4b). Here, SPI1 is linked to Alzheimer disease.