Hemimethylation of CpG dyads is characteristic of secondary DMRs associated with imprinted loci and correlates with 5-hydroxymethylcytosine at paternally methylated sequences.

In contrast, the difference in hemimethylation levels at either paternally or maternally methylated primary DMRs when compared to any of the secondary DMRs we analyzed was highly significant (Fig. 5 d) and the differences were notably more significant between primary vs. secondary DMRs than between secondary DMRs (Fig. 5 c).

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