AD risk variants were significantly enriched after correction for multiple testing using the Bonferroni method at DHSs containing the motifs SPI1 (PU.1) (z score = 5.53, corrected P < 1.30 × 10− 3), EGR1 (z score = 4.40, corrected P < 1.30 × 10− 3), MEF2A (z score = 4.08, corrected P = 0.023) or CEBPA (z score = 3.68, corrected P = 0.013) (Fig. 4a). Here, EGR1 is linked to Alzheimer disease.