SMC4 and cyclic hematopoiesis: Linkage disequilibrium (LD)-based clumping of 10,013,700 common autosomal and X chromosomal variants identified seven independent (r2 < 0.05) genome-wide significant loci (lead variant P < 5 × 10−8) associated with risk of developing CH, including three CH loci previously reported17 in European-ancestry populations: two at 5p15.33-TERT and one at 3q25.33-SMC4 (Fig. 4a and Supplementary Table 16).