FACT complex is required for DNA demethylation at heterochromatin during reproduction in Arabidopsis

Significance The chromatin remodeling activities of the FACT (facilitates chromatin transactions) complex are required for many cellular functions, including transcription, DNA replication, and repair. Here, we demonstrate that the two FACT subunits, SSRP1 and SPT16, are also required for genome-wide DNA demethylation and regulation of gene imprinting during Arabidopsis reproduction. Without FACT, Arabidopsis seeds undergo abnormal development and exhibit aberrant DNA hypermethylation, including at imprinting control region loci. We show that FACT associates with the DEMETER (DME) DNA demethylase, facilitating DNA demethylation at over half of DME’s targets, specifically those which reside in heterochromatin. These results provide insight into upstream events in the DNA demethylation pathway and reveal the importance of chromatin remodeling for DNA demethylation during Arabidopsis reproduction.