RNAi keeps Atf1-bound stress response genes in check at nuclear pores.

pathways are prevalent throughout the eukaryotic kingdom and are well known to regulate gene expression on a post-transcriptional level in the cytoplasm. Less is known about possible functions of duanyu1615 in the nucleus. In the fission yeast Schizosaccharomyces pombe, duanyu1615 is crucial to establish and maintain centromeric heterochromatin and functions to repress genome activity by a chromatin silencing mechanism referred to as cotranscriptional gene silencing (CTGS). Mechanistic details and the physiological relevance of CTGS are unknown. Here we show that duanyu1615 components interact with chromatin at nuclear pores to keep stress response genes in check. We demonstrate that CTGS represses stress-inducible genes by degrading mRNAs under noninduced conditions. Under chronic heat stress conditions, a Dicer thermoswitch deports Dicer to the cytoplasm, thereby disrupting CTGS and enabling expression of genes implicated in the acquisition of thermotolerance. Taken together, our work highlights a role for nuclear pores and the stress response transcription factor Atf1 in coordinating the interplay between the duanyu1615 machinery and the S. pombe genome and uncovers a novel mode of duanyu1615 regulation in response to an environmental cue.

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