Epigenetic silencing of a multifunctional plant stress regulator.

The central regulator of the ethylene (ET) signaling pathway, which controls a plethora of developmental programs and responses to environmental cues in plants, is ETHYLENE-INSENSITIVE2 (EIN2). Here we identify a chromatin-dependent regulatory mechanism at requiring two genes: ETHYLENE-INSENSITIVE6 (EIN6), which is a H3K27me3 demethylase also known as (), and EIN6 ENHANCER (EEN), the homolog of the yeast INO80 chromatin remodeling complex subunit (). Strikingly, EIN6 (REF6) and the INO80 complex redundantly control the level and the localization of the repressive histone modification H3K27me3 and the histone variant H2A.Z at the 5' untranslated region (5'UTR) intron of . Concomitant loss of EIN6 (REF6) and the INO80 complex shifts the chromatin landscape at to a repressive state causing a dramatic reduction of expression. These results uncover a unique type of chromatin regulation which safeguards the expression of an essential multifunctional plant stress regulator.