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Background  The control of gene expression is essential for growth and responses to environmental changes in various organisms. It is known that some meiosis-specific genes are silenced during mitosis and expressed upon nitrogen starvation in  Schizosaccharomyces pombe.  When the factors responsible for this regulation were studied, a  hip3  mutant was isolated via discovery of a defect in the transcriptional repression of meiosis-specific genes. Hip3 is a subunit of the HIRA ( hi stone  r egulatory complex  A ) complex, which consists of four subunits (Hip1, Hip3, Hip4 and Slm9) and acts as a histone chaperone that is independent of DNA replication.    Methodology/Principal Findings  In a search for mutants, the meiosis-specific gene  SPCC663.14c   +  was identified by screening for genes that are silenced during mitosis and induced upon nitrogen starvation. A reporter plasmid that expresses the  ura4   +  gene driven by the  SPCC663.14c   +  promoter was constructed. Screening for suppressor mutants was then carried out in nitrogen-rich medium without uracil. A mutant with a mutation in the  hip3 +   gene was isolated and named  hip3-1 . This mutation alleviated the transcriptional repression of the  ura4 +   gene on the reporter plasmid and of the endogenous  SPCC663.14c   +  gene in the presence of nitrogen. A ChIP assay revealed that RNA polymerase II (Pol II) and TFIIE were enriched at the  SPCC663.14c   +  locus, whereas the levels of histone H3 were decreased in  hip3-1  cells. Intriguingly, histone H3 was heavily modified at the  SPCC663.14c   +  locus in  hip3-1  cells; these modifications included tri-methylation and acetylation of H3 lysine 9 (H3K9), mono-methylation of H3 arginine 2 (H3R2), and tri-methylation of H3 lysine 4 (H3K4). In addition, the tri-methylation of H3K9 and H3K4 were strongly elevated in  hip3-1  mutants.    Conclusions  Taken together, these results indicate that Hip3 plays important roles in the control of histone modifications at meiosis-specific gene loci and induces their transcriptional repression.

The HIRA Complex Subunit Hip3 Plays Important Roles in the Silencing of Meiosis-Specific Genes in Schizosaccharomyces pombe