The study shows that silencing of NOR2 rRNA genes in Arabidopsis thaliana depends primarily on CHH-context cytosine methylation, particularly mediated by CMT2 and the chromatin remodeler DDM1, rather than CG or CHG methylation. Comparative promoter analysis revealed a prevalence of CHH sites in plant rDNA promoters, explaining why CHH methylation mutants disrupt NOR2 silencing more strongly, while NOR2 loci are hyper‑methylated and more condensed than NOR4.

The study examined transposable element (TE) silencing in the duckweed Spirodela polyrhiza, which exhibits unusually low DNA methylation, scarce 24‑nt siRNAs, and missing RdDM components. While degenerated TEs lack DNA methylation and H3K9me2, they retain heterochromatin marks H3K9me1 and H3K27me1, whereas the few intact TEs show high DNA methylation and H3K9me2, indicating a shift in RdDM focus toward potentially active TEs and suggesting heterochromatin can be maintained independently of DNA methylation in flowering plants.