Plant-specific RNA Polymerase V (Pol V) transcribes noncoding RNAs in the RNA-directed DNA methylation pathway, thereby influencing gene expression and genome stability by controlling de novo DNA methylation. However, the mechanisms governing precise chromatin localization and transcriptional activities of Pol V remain elusive. Here we show that Pol V localization is spatially constrained by the chromatin regulators microrchidia (MORC) and MORPHEUS' MOLECULE 1 (MOM1). MORC and MOM1 promote Pol V occupancy at sites near active chromatin, whereas their loss leads to redistribution of Pol V into CMT3-enriched heterochromatin, accompanied by noncoding RNA transcription, small RNA production and DNA methylation. Our findings reveal a combinatorial model in which recruitment, spatial constraint and DNA methylation feedback collectively define Pol V chromatin distribution and epigenetic function.
He, X., Li, Z., Xue, Y., Guo, J., Liu, X., Feng, S., Zhong, Z., Jacobsen, S. E.
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