protein coding gene - clr4 (SPBC428.08c) - histone lysine H3-K9 methyltransferase (Suv39) Clr4

Gene summary

Standard name

clr4

Systematic ID

SPBC428.08c

Product

histone lysine H3-K9 methyltransferase (Suv39) Clr4

Organism

Schizosaccharomyces pombe (fission yeast)

UniProt ID

O60016

ORFeome ID

23/23G02

Characterisation status

biological role published

Feature type

mRNA gene

Genomic location

chromosome II: 453996..458410 reverse strand

Annotation

Protein features

Orthologs

• SUV39H1 (HGNC:11479) Homo sapiens
• SUV39H2 (HGNC:17287) Homo sapiens
• clr4 (SJAG_04574) Schizosaccharomyces japonicus

References / Literature

PMID:16428807 - Hip3 interacts with the HIRA proteins Hip1 and Slm9 and is required for transcriptional silencing and accurate chromosome segregation.

Greenall A et al. J Biol Chem 2006 Mar 31;281(13):8732-9

PMID:30051891 - Automethylation-induced conformational switch in Clr4 (Suv39h) maintains epigenetic stability.

Iglesias N et al. Nature 2018 Aug;560(7719):504-508

PMID:33693625 - Dri1 mediates heterochromatin assembly via RNAi and histone deacetylation.

Ban H et al. Genetics 2021 May 17;218(1)

PMID:11069763 - Live analysis of lagging chromosomes during anaphase and their effect on spindle elongation rate in fission yeast.

Pidoux AL et al. J Cell Sci 2000 Dec;113 Pt 23:4177-91

PMID:23950735 - Global analysis of fission yeast mating genes reveals new autophagy factors.

Sun LL et al. PLoS Genet 2013;9(8):e1003715

PMID:36200823 - The chromatin remodeler RSC prevents ectopic CENP-A propagation into pericentromeric heterochromatin at the chromatin boundary.

Tsunemine S et al. Nucleic Acids Res 2022 Oct 28;50(19):10914-10928

PMID:37400983 - Cooperative DNA-binding activities of Chp2 are critical for its function in heterochromatin assembly.

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PMID:33823663 - A TOR (target of rapamycin) and nutritional phosphoproteome of fission yeast reveals novel targets in networks conserved in humans.

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PMID:15372076 - A chromodomain protein, Chp1, is required for the establishment of heterochromatin in fission yeast.

Sadaie M et al. EMBO J 2004 Oct 01;23(19):3825-35

PMID:38181050 - The Cross-Regulation Between Set1, Clr4, and Lsd1/2 in Schizosaccharomyces pombe.

Liu H et al. PLoS Genet 2024 Jan 05;20(1):e1011107

PMID:39945308 - Characterization of the Swi6/HP1 binding motif in its partner protein reveals the basis for the functional divergence of the HP1 family proteins in fission yeast.

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PMID:37787768 - Broad functional profiling of fission yeast proteins using phenomics and machine learning.

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PMID:25831549 - Epigenetics. Epigenetic inheritance uncoupled from sequence-specific recruitment.

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PMID:28765164 - Selective termination of lncRNA transcription promotes heterochromatin silencing and cell differentiation.

Touat-Todeschini L et al. EMBO J 2017 Sep 01;36(17):2626-2641

PMID:27738016 - RNA interference is essential for cellular quiescence.

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PMID:25957277 - Intrinsic Toxicity of Unchecked Heterochromatin Spread Is Suppressed by Redundant Chromatin Boundary Functions in Schizosacchromyces pombe.

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PMID:33511417 - Repression of a large number of genes requires interplay between homologous recombination and HIRA.

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PMID:16540522 - Nuclear RanGAP is required for the heterochromatin assembly and is reciprocally regulated by histone H3 and Clr4 histone methyltransferase in Schizosaccharomyces pombe.

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PMID:21760946 - Identification of genes affecting the toxicity of anti-cancer drug bortezomib by genome-wide screening in S. pombe.

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PMID:20935472 - Laser microsurgery provides evidence for merotelic kinetochore attachments in fission yeast cells lacking Pcs1 or Clr4.

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PMID:9572142 - Defective meiosis in telomere-silencing mutants of Schizosaccharomyces pombe.

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PMID:16246721 - The nucleation and maintenance of heterochromatin by a histone deacetylase in fission yeast.

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PMID:18923422 - Rad51 suppresses gross chromosomal rearrangement at centromere in Schizosaccharomyces pombe.

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PMID:29084823 - Phosphorylation of the RNA-binding protein Zfs1 modulates sexual differentiation in fission yeast.

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PMID:34984977 - Functional profiling of long intergenic non-coding RNAs in fission yeast.

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PMID:29686279 - Iron homeostasis regulates facultative heterochromatin assembly in adaptive genome control.

Gallagher PS et al. Nat Struct Mol Biol 2018 May;25(5):372-383

PMID:15607976 - Two RNAi complexes, RITS and RDRC, physically interact and localize to noncoding centromeric RNAs.

Motamedi MR et al. Cell 2004 Dec 17;119(6):789-802

PMID:17434129 - S. pombe LSD1 homologs regulate heterochromatin propagation and euchromatic gene transcription.

Lan F et al. Mol Cell 2007 Apr 13;26(1):89-101

PMID:22683269 - HP1(Swi6) mediates the recognition and destruction of heterochromatic RNA transcripts.

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PMID:24240238 - Elimination of shelterin components bypasses RNAi for pericentric heterochromatin assembly.

Tadeo X et al. Genes Dev 2013 Nov 15;27(22):2489-99

PMID:26173815 - Genome-wide screen of fission yeast mutants for sensitivity to 6-azauracil, an inhibitor of transcriptional elongation.

Zhou H et al. Yeast 2015 Oct;32(10):643-55

PMID:28143796 - Clr4 specificity and catalytic activity beyond H3K9 methylation.

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PMID:29214404 - The 19S proteasome regulates subtelomere silencing and facultative heterochromatin formation in fission yeast.

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PMID:8001791 - Three additional linkage groups that repress transcription and meiotic recombination in the mating-type region of Schizosaccharomyces pombe.

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PMID:19164572 - Msc1 links dynamic Swi6/HP1 binding to cell fate determination.

Lawrence RJ et al. Proc Natl Acad Sci U S A 2009 Jan 27;106(4):1163-8

PMID:20661445 - Rad3 decorates critical chromosomal domains with gammaH2A to protect genome integrity during S-Phase in fission yeast.

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PMID:29216371 - Fission yeast Ccq1 is a modulator of telomerase activity.

Armstrong CA et al. Nucleic Acids Res 2018 Jan 25;46(2):704-716

PMID:22144463 - RNA elimination machinery targeting meiotic mRNAs promotes facultative heterochromatin formation.

Zofall M et al. Science 2012 Jan 06;335(6064):96-100

PMID:17627824 - The kinetochore proteins Pcs1 and Mde4 and heterochromatin are required to prevent merotelic orientation.

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PMID:23697806 - A genome-wide resource of cell cycle and cell shape genes of fission yeast.

Hayles J et al. Open Biol 2013 May 22;3(5):130053

PMID:8138176 - Mutations in rik1, clr2, clr3 and clr4 genes asymmetrically derepress the silent mating-type loci in fission yeast.

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PMID:31206516 - Regulation of ectopic heterochromatin-mediated epigenetic diversification by the JmjC family protein Epe1.

Sorida M et al. PLoS Genet 2019 Jun;15(6):e1008129

PMID:20705239 - The methyltransferase activity of Clr4Suv39h triggers RNAi independently of histone H3K9 methylation.

Gerace EL et al. Mol Cell 2010 Aug 13;39(3):360-72

PMID:22990236 - Hrp3 controls nucleosome positioning to suppress non-coding transcription in eu- and heterochromatin.

Shim YS et al. EMBO J 2012 Nov 28;31(23):4375-87

PMID:22319459 - Raf1 Is a DCAF for the Rik1 DDB1-like protein and has separable roles in siRNA generation and chromatin modification.

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PMID:25972440 - Global regulation of heterochromatin spreading by Leo1.

Verrier L et al. Open Biol 2015 May;5(5)

PMID:34010645 - The histone H3K9M mutation synergizes with H3K14 ubiquitylation to selectively sequester histone H3K9 methyltransferase Clr4 at heterochromatin.

Shan CM et al. Cell Rep 2021 May 18;35(7):109137

PMID:23151475 - RNAi triggered by specialized machinery silences developmental genes and retrotransposons.

Yamanaka S et al. Nature 2013 Jan 24;493(7433):557-60

PMID:19693008 - Histone H2A.Z cooperates with RNAi and heterochromatin factors to suppress antisense RNAs.

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PMID:23260662 - Lsd1 and lsd2 control programmed replication fork pauses and imprinting in fission yeast.

Holmes A et al. Cell Rep 2012 Dec 27;2(6):1513-20

PMID:26537787 - Targeting of SUMO substrates to a Cdc48-Ufd1-Npl4 segregase and STUbL pathway in fission yeast.

Køhler JB et al. Nat Commun 2015 Nov 05;6:8827

PMID:11273706 - Solution structure, domain features, and structural implications of mutants of the chromo domain from the fission yeast histone methyltransferase Clr4.

Horita DA et al. J Mol Biol 2001 Mar 30;307(3):861-70

PMID:21844224 - Spt6 is required for heterochromatic silencing in the fission yeast Schizosaccharomyces pombe.

Kiely CM et al. Mol Cell Biol 2011 Oct;31(20):4193-204

PMID:21892171 - Defects in RNA quality control factors reveal RNAi-independent nucleation of heterochromatin.

Reyes-Turcu FE et al. Nat Struct Mol Biol 2011 Sep 04;18(10):1132-8

PMID:27451393 - Role of Inner Nuclear Membrane Protein Complex Lem2-Nur1 in Heterochromatic Gene Silencing.

Banday S et al. J Biol Chem 2016 Sep 16;291(38):20021-9

PMID:18957202 - Lid2 is required for coordinating H3K4 and H3K9 methylation of heterochromatin and euchromatin.

Li F et al. Cell 2008 Oct 17;135(2):272-83

PMID:29722648 - Distinct 'safe zones' at the nuclear envelope ensure robust replication of heterochromatic chromosome regions.

Ebrahimi H et al. Elife 2018 May 03;7

PMID:32499408 - The molecular chaperone Hsp90 regulates heterochromatin assembly through stabilizing multiple complexes in fission yeast.

Sun L et al. J Cell Sci 2020 Jul 07;133(13)

PMID:34524082 - SUV39 SET domains mediate crosstalk of heterochromatic histone marks.

Stirpe A et al. Elife 2021 Sep 15;10

PMID:37445861 - Resistance to Chemotherapeutic 5-Fluorouracil Conferred by Modulation of Heterochromatic Integrity through Ino80 Function in Fission Yeast.

Lim KK et al. Int J Mol Sci 2023 Jun 26;24(13)

PMID:18345014 - Roles of the Clr4 methyltransferase complex in nucleation, spreading and maintenance of heterochromatin.

Zhang K et al. Nat Struct Mol Biol 2008 Apr;15(4):381-8

PMID:18252195 - Methylations of histone H3 lysine 9 and lysine 36 are functionally linked to DNA replication checkpoint control in fission yeast.

Kim HS et al. Biochem Biophys Res Commun 2008 Apr 04;368(2):419-25

PMID:39367033 - Quantitative proteomics and phosphoproteomics profiling of meiotic divisions in the fission yeast Schizosaccharomyces pombe.

Sivakova B et al. Sci Rep 2024 Oct 04;14(1):23105

PMID:40446033 - Mechanistic insights into the stimulation of the histone H3K9 methyltransferase Clr4 by proximal H3K14 ubiquitination.

Du Y et al. Sci Adv 2025 May 30;11(22):eadu1864

PMID:25245948 - Tls1 regulates splicing of shelterin components to control telomeric heterochromatin assembly and telomere length.

Wang J et al. Nucleic Acids Res 2014 Oct;42(18):11419-32

PMID:15317843 - Schizosaccharomyces pombe RanGAP homolog, SpRna1, is required for centromeric silencing and chromosome segregation.

Kusano A et al. Mol Biol Cell 2004 Nov;15(11):4960-70

PMID:18030666 - Mutations in deoxyribonucleotide biosynthesis pathway cause spreading of silencing across heterochromatic barriers at the mating-type region of the fission yeast.

Singh G et al. Yeast 2008 Feb;25(2):117-28

PMID:27984744 - Survival in Quiescence Requires the Euchromatic Deployment of Clr4/SUV39H by Argonaute-Associated Small RNAs.

Joh RI et al. Mol Cell 2016 Dec 15;64(6):1088-1101

PMID:31269446 - CPF Recruitment to Non-canonical Transcription Termination Sites Triggers Heterochromatin Assembly and Gene Silencing.

Vo TV et al. Cell Rep 2019 Jul 02;28(1):267-281.e5

PMID:36617881 - Tandemly repeated genes promote RNAi-mediated heterochromatin formation via an antisilencing factor, Epe1, in fission yeast.

Asanuma T et al. Genes Dev 2022 Nov;36(21-24):1145-1159

PMID:38289024 - Heat stress-induced activation of MAPK pathway attenuates Atf1-dependent epigenetic inheritance of heterochromatin in fission yeast.

Sun L et al. Elife 2024 Jan 30;13

PMID:23613586 - Structural analysis of Stc1 provides insights into the coupling of RNAi and chromatin modification.

He C et al. Proc Natl Acad Sci U S A 2013 May 21;110(21):E1879-88

PMID:17114925 - A cullin E3 ubiquitin ligase complex associates with Rik1 and the Clr4 histone H3-K9 methyltransferase and is required for RNAi-mediated heterochromatin formation.

Hong EJ et al. RNA Biol 2005;2(3):106-11

PMID:31784357 - Native Chromatin Proteomics Reveals a Role for Specific Nucleoporins in Heterochromatin Organization and Maintenance.

Iglesias N et al. Mol Cell 2020 Jan 02;77(1):51-66.e8

PMID:18204818 - Identification of small molecules inducing apoptosis by cell-based assay using fission yeast deletion mutants.

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PMID:37913773 - TOR inactivation triggers heterochromatin formation in rDNA during glucose starvation.

Hirai H et al. Cell Rep 2023 Nov 28;42(11):113320

PMID:24210919 - Mtr4-like protein coordinates nuclear RNA processing for heterochromatin assembly and for telomere maintenance.

Lee NN et al. Cell 2013 Nov 21;155(5):1061-74

PMID:41258116 - TORC2 inactivation promotes heterochromatin formation in rDNA and prolongs viability of quiescent fission yeast cells.

Hirai H et al. Commun Biol 2025 Nov 19;8(1):1606

PMID:18761674 - Fission yeast chromatin assembly factor 1 assists in the replication-coupled maintenance of heterochromatin.

Dohke K et al. Genes Cells 2008 Oct;13(10):1027-43

PMID:21151114 - Dicer associates with chromatin to repress genome activity in Schizosaccharomyces pombe.

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PMID:23101633 - Quantitative analysis of fission yeast transcriptomes and proteomes in proliferating and quiescent cells.

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PMID:27334362 - Inner nuclear membrane protein Lem2 augments heterochromatin formation in response to nutritional conditions.

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PMID:31468675 - H3K14 ubiquitylation promotes H3K9 methylation for heterochromatin assembly.

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PMID:21385875 - Roles of fission yeast Grc3 protein in ribosomal RNA processing and heterochromatic gene silencing.

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PMID:25720772 - Quantitative phosphoproteomics reveals pathways for coordination of cell growth and division by the conserved fission yeast kinase pom1.

Kettenbach AN et al. Mol Cell Proteomics 2015 May;14(5):1275-87

PMID:27538348 - Set3 contributes to heterochromatin integrity by promoting transcription of subunits of Clr4-Rik1-Cul4 histone methyltransferase complex in fission yeast.

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PMID:22474355 - Heterochromatin protein 1 homologue Swi6 acts in concert with Ers1 to regulate RNAi-directed heterochromatin assembly.

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PMID:10835380 - Four chromo-domain proteins of Schizosaccharomyces pombe differentially repress transcription at various chromosomal locations.

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PMID:23851719 - Spt6 prevents transcription-coupled loss of posttranslationally modified histone H3.

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PMID:15218150 - RNAi-independent heterochromatin nucleation by the stress-activated ATF/CREB family proteins.

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PMID:15537537 - Heterochromatin regulates cell type-specific long-range chromatin interactions essential for directed recombination.

Jia S et al. Cell 2004 Nov 12;119(4):469-80

PMID:21211723 - Asf1/HIRA facilitate global histone deacetylation and associate with HP1 to promote nucleosome occupancy at heterochromatic loci.

Yamane K et al. Mol Cell 2011 Jan 07;41(1):56-66

PMID:26988418 - The proper connection between shelterin components is required for telomeric heterochromatin assembly.

Wang J et al. Genes Dev 2016 Apr 01;30(7):827-39

PMID:12618370 - Schizosaccharomyces pombe essential genes: a pilot study.

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PMID:15475954 - RITS acts in cis to promote RNA interference-mediated transcriptional and post-transcriptional silencing.

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PMID:15947136 - RNA polymerase II is required for RNAi-dependent heterochromatin assembly.

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PMID:23849629 - Division of labor between the chromodomains of HP1 and Suv39 methylase enables coordination of heterochromatin spread.

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PMID:25452419 - Parallel profiling of fission yeast deletion mutants for proliferation and for lifespan during long-term quiescence.

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PMID:29136238 - Regulation of transcriptional silencing and chromodomain protein localization at centromeric heterochromatin by histone H3 tyrosine 41 phosphorylation in fission yeast.

Ren B et al. Nucleic Acids Res 2018 Jan 09;46(1):189-202

PMID:22681890 - Hierarchical modularity and the evolution of genetic interactomes across species.

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PMID:29237752 - Ser7 of RNAPII-CTD facilitates heterochromatin formation by linking ncRNA to RNAi.

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PMID:16823372 - ORFeome cloning and global analysis of protein localization in the fission yeast Schizosaccharomyces pombe.

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PMID:23028377 - Factors that promote H3 chromatin integrity during transcription prevent promiscuous deposition of CENP-A(Cnp1) in fission yeast.

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PMID:20178743 - Dicer-independent primal RNAs trigger RNAi and heterochromatin formation.

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PMID:22895252 - A novel RNAi protein, Dsh1, assembles RNAi machinery on chromatin to amplify heterochromatic siRNA.

Kawakami K et al. Genes Dev 2012 Aug 15;26(16):1811-24

PMID:29996109 - Quantitative Phosphoproteomics Reveals the Signaling Dynamics of Cell-Cycle Kinases in the Fission Yeast Schizosaccharomyces pombe.

Swaffer MP et al. Cell Rep 2018 Jul 10;24(2):503-514

PMID:27697832 - Rad51 and Rad54 promote noncrossover recombination between centromere repeats on the same chromatid to prevent isochromosome formation.

Onaka AT et al. Nucleic Acids Res 2016 Dec 15;44(22):10744-10757

PMID:26582768 - Abo1, a conserved bromodomain AAA-ATPase, maintains global nucleosome occupancy and organisation.

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PMID:34375584 - A composite DNA element that functions as a maintainer required for epigenetic inheritance of heterochromatin.

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GO_REF:0000002 - Comments

PMID:12389037 - Structure of the SET domain histone lysine methyltransferase Clr4.

Min J et al. Nat Struct Biol 2002 Nov;9(11):828-32

PMID:29167439 - Eroded telomeres are rearranged in quiescent fission yeast cells through duplications of subtelomeric sequences.

Maestroni L et al. Nat Commun 2017 Nov 22;8(1):1684

PMID:36408846 - Differential phosphorylation of Clr4 SUV39H by Cdk1 accompanies a histone H3 methylation switch that is essential for gametogenesis.

Kuzdere T et al. EMBO Rep 2023 Jan 09;24(1):e55928

PMID:38285941 - Specialized replication of heterochromatin domains ensures self-templated chromatin assembly and epigenetic inheritance.

Nathanailidou P et al. Proc Natl Acad Sci U S A 2024 Feb 06;121(6):e2315596121

PMID:37189341 - Regulation of the SUV39H Family Methyltransferases: Insights from Fission Yeast.

Nakamura R et al. Biomolecules 2023 Mar 25;13(4)

PMID:24013502 - Epe1 recruits BET family bromodomain protein Bdf2 to establish heterochromatin boundaries.

Wang J et al. Genes Dev 2013 Sep 01;27(17):1886-902

PMID:18505873 - Schizosaccharomyces pombe histone acetyltransferase Mst1 (KAT5) is an essential protein required for damage response and chromosome segregation.

Gómez EB et al. Genetics 2008 Jun;179(2):757-71

PMID:20211136 - Stc1: a critical link between RNAi and chromatin modification required for heterochromatin integrity.

Bayne EH et al. Cell 2010 Mar 05;140(5):666-77

PMID:28682306 - Unique roles for histone H3K9me states in RNAi and heritable silencing of transcription.

Jih G et al. Nature 2017 Jul 27;547(7664):463-467

PMID:16762840 - Swi6/HP1 recruits a JmjC domain protein to facilitate transcription of heterochromatic repeats.

Zofall M et al. Mol Cell 2006 Jun 09;22(5):681-92

PMID:12867036 - Sir2 regulates histone H3 lysine 9 methylation and heterochromatin assembly in fission yeast.

Shankaranarayana GD et al. Curr Biol 2003 Jul 15;13(14):1240-6

PMID:23050226 - A genetic screen to discover pathways affecting cohesin function in Schizosaccharomyces pombe identifies chromatin effectors.

Chen Z et al. G3 (Bethesda) 2012 Oct;2(10):1161-8

PMID:28318821 - SNF2 Family Protein Fft3 Suppresses Nucleosome Turnover to Promote Epigenetic Inheritance and Proper Replication.

Taneja N et al. Mol Cell 2017 Apr 06;66(1):50-62.e6

PMID:16127433 - Ubiquitin ligase component Cul4 associates with Clr4 histone methyltransferase to assemble heterochromatin.

Jia S et al. Nat Cell Biol 2005 Oct;7(10):1007-13

PMID:30726745 - Fission Yeast NDR/LATS Kinase Orb6 Regulates Exocytosis via Phosphorylation of the Exocyst Complex.

Tay YD et al. Cell Rep 2019 Feb 05;26(6):1654-1667.e7

PMID:24920274 - Meiotic long non-coding meiRNA accumulates as a dot at its genetic locus facilitated by Mmi1 and plays as a decoy to lure Mmi1.

Shichino Y et al. Open Biol 2014 Jun;4(6):140022

PMID:11242054 - Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain.

Bannister AJ et al. Nature 2001 Mar 01;410(6824):120-4

PMID:10949293 - Regulation of chromatin structure by site-specific histone H3 methyltransferases.

Rea S et al. Nature 2000 Aug 10;406(6796):593-9

PMID:20537132 - Global fitness profiling of fission yeast deletion strains by barcode sequencing.

Han TX et al. Genome Biol 2010;11(6):R60

PMID:16168376 - SUMO modification is involved in the maintenance of heterochromatin stability in fission yeast.

Shin JA et al. Mol Cell 2005 Sep 16;19(6):817-28

PMID:20062003 - Hairpin RNA induces secondary small interfering RNA synthesis and silencing in trans in fission yeast.

Simmer F et al. EMBO Rep 2010 Feb;11(2):112-8

PMID:24763107 - Absolute proteome and phosphoproteome dynamics during the cell cycle of Schizosaccharomyces pombe (Fission Yeast).

Carpy A et al. Mol Cell Proteomics 2014 Aug;13(8):1925-36

PMID:25483073 - Chromosome segregation and organization are targets of 5'-Fluorouracil in eukaryotic cells.

Mojardín L et al. Cell Cycle 2015;14(2):206-18

PMID:29294138 - Editor's Highlight: A Genome-wide Screening of Target Genes Against Silver Nanoparticles in Fission Yeast.

Lee AR et al. Toxicol Sci 2018 Jan 01;161(1):171-185

PMID:29632066 - TOR complex 2 in fission yeast is required for chromatin-mediated gene silencing and assembly of heterochromatic domains at subtelomeres.

Cohen A et al. J Biol Chem 2018 May 25;293(21):8138-8150

PMID:39094565 - Mapping the dynamics of epigenetic adaptation in S. pombe during heterochromatin misregulation.

Larkin A et al. Dev Cell 2024 Jul 26;

PMID:39747188 - PhpC NF-Y transcription factor infiltrates heterochromatin to generate cryptic intron-containing transcripts crucial for small RNA production.

Srivastav MK et al. Nat Commun 2025 Jan 02;16(1):268

PMID:24013500 - A conserved ncRNA-binding protein recruits silencing factors to heterochromatin through an RNAi-independent mechanism.

Marina DB et al. Genes Dev 2013 Sep 01;27(17):1851-6

PMID:25002536 - A novel histone deacetylase complex in the control of transcription and genome stability.

Zilio N et al. Mol Cell Biol 2014 Sep 15;34(18):3500-14

PMID:26804021 - Shugoshin forms a specialized chromatin domain at subtelomeres that regulates transcription and replication timing.

Tashiro S et al. Nat Commun 2016 Jan 25;7:10393

PMID:18809570 - Balance between distinct HP1 family proteins controls heterochromatin assembly in fission yeast.

Sadaie M et al. Mol Cell Biol 2008 Dec;28(23):6973-88

PMID:21224386 - Role of Swi6/HP1 self-association-mediated recruitment of Clr4/Suv39 in establishment and maintenance of heterochromatin in fission yeast.

Haldar S et al. J Biol Chem 2011 Mar 18;286(11):9308-20

PMID:32269268 - Abo1 is required for the H3K9me2 to H3K9me3 transition in heterochromatin.

Dong W et al. Sci Rep 2020 Apr 08;10(1):6055

PMID:29852001 - New insights into donor directionality of mating-type switching in Schizosaccharomyces pombe.

Maki T et al. PLoS Genet 2018 May;14(5):e1007424

PMID:11283354 - Role of histone H3 lysine 9 methylation in epigenetic control of heterochromatin assembly.

Nakayama J et al. Science 2001 Apr 06;292(5514):110-3

PMID:29784772 - A Heterochromatin Domain Forms Gradually at a New Telomere and Is Dynamic at Stable Telomeres.

Wang J et al. Mol Cell Biol 2018 Aug 01;38(15)

PMID:36830746 - Opposing Roles of FACT for Euchromatin and Heterochromatin in Yeast.

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