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protein coding gene - gar2 (SPAC140.02) - nucleolin, nucleolar histone chaperone required for rRNA metabolism

Gene summary

Standard name
gar2
Systematic ID
SPAC140.02
Product
nucleolin, nucleolar histone chaperone required for rRNA metabolism
Organism
Schizosaccharomyces pombe (fission yeast)
UniProt ID
P41891
ORFeome ID
24/24D02
Characterisation status
biological role published
Feature type
mRNA gene
Genomic location
chromosome I: 1897462..1899491 forward strand

Annotation

Comment

PBO:0016891 - longer transcript expressed during cold shock due to altered 3'-end processing

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Complementation

PBO:0016927 - functionally complements S. cerevisiae NSR1

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GO biological process

GO:0006338 - chromatin remodeling

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GO:0042254 - ribosome biogenesis

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GO:0016072 - rRNA metabolic process

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GO cellular component

GO:0001651 - dense fibrillar component

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GO:0005730 - nucleolus

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GO molecular function

GO:0140713 - histone chaperone activity

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GO:0003723 - RNA binding

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Genome organisation

PBO:0002125 - alternative transcripts

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Modification

MOD:00046 - O-phospho-L-serine

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MOD:00047 - O-phospho-L-threonine

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MOD:00696 - phosphorylated residue

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MOD:01149 - sumoylated lysine

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MOD:01148 - ubiquitinylated lysine

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Multi-locus phenotype

FYPO:0007376 - abolished epigenetic heterochromatin inheritance

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Genotypes:

FYPO:0007480 - decreased spatial extent of subtelomeric heterochromatin assembly

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Genotypes:

FYPO:0004948 - increased spatial extent of centromeric heterochromatin assembly

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Genotypes:

Protein features

PBO:0111764 - rrm RNA recognition motif

Qualitative gene expression

PomGeneEx:0000019 - protein level decreased

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PomGeneEx:0000027 - ribosomal density decreased

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PomGeneEx:0000012 - RNA level decreased

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Quantitative gene expression

PBO:0006310 - protein level

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PBO:0011963 - RNA level

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Single locus phenotype

FYPO:0003698 - abnormal dense body present in nucleolus

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Genotypes:

FYPO:0002736 - abnormal mitotic cell cycle phase

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Genotypes:

FYPO:0000361 - abnormal nucleolar morphology

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Genotypes:

FYPO:0003697 - cell wall invagination into cytoplasm

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Genotypes:

FYPO:0000080 - decreased cell population growth at low temperature

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Genotypes:

FYPO:0003743 - decreased cell population growth during glucose starvation

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Genotypes:

FYPO:0009053 - decreased cell population growth on glutamate nitrogen source

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Genotypes:

FYPO:0000684 - decreased cell population growth on glycerol carbon source

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Genotypes:

FYPO:0009091 - decreased cell population growth on lysine and proline nitrogen source

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Genotypes:

FYPO:0009092 - decreased cell population growth on lysine and serine nitrogen source

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Genotypes:

FYPO:0009073 - decreased cell population growth on lysine nitrogen source

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Genotypes:

FYPO:0002924 - decreased cell population growth on maltose carbon source

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Genotypes:

FYPO:0000250 - decreased cell population growth on proline nitrogen source

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Genotypes:

FYPO:0007562 - decreased cell population growth on serine nitrogen source

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Genotypes:

FYPO:0003412 - decreased chromatin silencing at centromere outer repeat

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Genotypes:

FYPO:0003695 - decreased cytosolic small ribosomal subunit level

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Genotypes:

FYPO:0007478 - decreased epigenetic heterochromatin inheritance

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Genotypes:

FYPO:0000470 - decreased mating type switching

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Genotypes:

FYPO:0003694 - decreased mature 18S rRNA level

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Genotypes:

FYPO:0001838 - decreased protein phosphorylation during vegetative growth

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Genotypes:

FYPO:0001117 - decreased RNA level during vegetative growth

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Genotypes:

FYPO:0001355 - decreased vegetative cell population growth

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Genotypes:

FYPO:0001122 - elongated vegetative cell

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Genotypes:

FYPO:0003696 - increased cytosolic large ribosomal subunit level

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Genotypes:

FYPO:0000825 - increased RNA level during vegetative growth

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Genotypes:

FYPO:0000245 - loss of viability in stationary phase

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Genotypes:

FYPO:0002619 - normal growth on sodium butyrate

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Genotypes:

FYPO:0002620 - normal growth on trichostatin A

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Genotypes:

FYPO:0001134 - normal mature 18S rRNA level

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Genotypes:

FYPO:0007234 - normal protein localization to centromere central core

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Genotypes:

FYPO:0000763 - resistance to cadmium

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Genotypes:

FYPO:0001034 - resistance to tunicamycin

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Genotypes:

FYPO:0007933 - sensitive to 2,2′-dipyridyl

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Genotypes:

FYPO:0001097 - sensitive to amitrole

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Genotypes:

FYPO:0009067 - sensitive to amorolfine

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Genotypes:

FYPO:0001701 - sensitive to bortezomib

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Genotypes:

FYPO:0001501 - sensitive to brefeldin A

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Genotypes:

FYPO:0000097 - sensitive to caffeine during vegetative growth

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Genotypes:

FYPO:0009080 - sensitive to calcofluor and sodium dodecyl sulfate

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Genotypes:

FYPO:0001188 - sensitive to Calcofluor White

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Genotypes:

FYPO:0009069 - sensitive to ciclopirox olamine

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Genotypes:

FYPO:0000104 - sensitive to cycloheximide

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Genotypes:

FYPO:0000799 - sensitive to diamide

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Genotypes:

FYPO:0007931 - sensitive to egtazic acid

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Genotypes:

FYPO:0000842 - sensitive to ethanol during vegetative growth

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Genotypes:

FYPO:0007928 - sensitive to ethylenediaminetetraacetic acid

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Genotypes:

FYPO:0000785 - sensitive to formamide

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Genotypes:

FYPO:0000087 - sensitive to hydrogen peroxide

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Genotypes:

FYPO:0000088 - sensitive to hydroxyurea

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Genotypes:

FYPO:0009071 - sensitive to itraconazole

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Genotypes:

FYPO:0001719 - sensitive to lithium

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Genotypes:

FYPO:0006836 - sensitive to magnesium chloride

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Genotypes:

FYPO:0009088 - sensitive to magnesium chloride and sodium dodecyl sulfate

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Genotypes:

FYPO:0000089 - sensitive to methyl methanesulfonate

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Genotypes:

FYPO:0001214 - sensitive to potassium chloride

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Genotypes:

FYPO:0009082 - sensitive to potassium chloride and methyl methanesulfonate

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Genotypes:

FYPO:0007924 - sensitive to potassium chloride and sodium dodecyl sulfate

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Genotypes:

FYPO:0002328 - sensitive to terbinafine

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Genotypes:

FYPO:0000797 - sensitive to tert-butyl hydroperoxide

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Genotypes:

FYPO:0000091 - sensitive to thiabendazole

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Genotypes:

FYPO:0002701 - sensitive to torin1

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Genotypes:

FYPO:0001457 - sensitive to tunicamycin

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Genotypes:

FYPO:0000115 - sensitive to valproic acid

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Genotypes:

FYPO:0003656 - sensitive to vanadate

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Genotypes:

FYPO:0001234 - slow vegetative cell population growth

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Genotypes:

FYPO:0002060 - viable vegetative cell population

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Genotypes:

FYPO:0002177 - viable vegetative cell with normal cell morphology

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Genotypes:

Taxonomic conservation

PBO:0011065 - conserved in eukaryotes

PBO:0011071 - conserved in eukaryotes only

PBO:0011064 - conserved in fungi

PBO:0011069 - conserved in metazoa

PBO:0011070 - conserved in vertebrates

PBO:0006222 - predominantly single copy (one to one)

Protein features

IDNameInterPro nameDB name
PF00076RRM_1RRM_domPFAM
cd12448RRM2_gar2Gar2_RRM2CDD
PS50102RRMRRM_domPROSITE_PROFILES
SM00360rrm1_1RRM_domSMART
SSF54928RNA-binding domain, RBDRBD_domain_sfSUPERFAMILY
G3DSA:3.30.70.330Nucleotide-bd_a/b_plait_sfGENE3D
PTHR48024GEO13361P1-RELATEDRNA-binding_regPANTHER
mobidb-lite-Disorderdisorder_predictionMOBIDB-Disorder
mobidb-lite-Glycine-richdisorder_predictionMOBIDB-Glycine-rich
mobidb-lite-Low-complexitydisorder_predictionMOBIDB-Low-complexity
mobidb-lite-Negative-Polyelectrolytedisorder_predictionMOBIDB-Negative-Polyelectrolyte
mobidb-lite-Polyampholytedisorder_predictionMOBIDB-Polyampholyte
mobidb-lite-Positive-Polyelectrolytedisorder_predictionMOBIDB-Positive-Polyelectrolyte

Orthologs

References / Literature

PMID:24957674 - Yeast X-chromosome-associated protein 5 (Xap5) functions with H2A.Z to suppress aberrant transcripts.
Anver S et al. EMBO Rep 2014 Aug;15(8):894-902
PMID:24713849 - Post-transcriptional regulation of meiotic genes by a nuclear RNA silencing complex.
Egan ED et al. RNA 2014 Jun;20(6):867-81
PMID:34250083 - Barcode sequencing and a high-throughput assay for chronological lifespan uncover ageing-associated genes in fission yeast.
Romila CA et al. Microb Cell 2021 Jul 05;8(7):146-160
PMID:36361590 - Defining the Functional Interactome of Spliceosome-Associated G-Patch Protein Gpl1 in the Fission Yeast Schizosaccharomyces pombe .
Selicky T et al. Int J Mol Sci 2022 Oct 24;23(21)
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:18257517 - Phosphoproteome analysis of fission yeast.
Wilson-Grady JT et al. J Proteome Res 2008 Mar;7(3):1088-97
PMID:25373780 - A genomic Multiprocess survey of machineries that control and link cell shape, microtubule organization, and cell-cycle progression.
Graml V et al. Dev Cell 2014 Oct 27;31(2):227-239
PMID:32454109 - Transcription-facilitating histone chaperons interact with genomic and synthetic G4 structures.
Pavlova II et al. Int J Biol Macromol 2020 Oct 01;160:1144-1157
PMID:40406582 - The Schizosaccharomyces pombe nucleolar protein Nsk1 modulates rDNA silencing during interphase.
Chen JS et al. MicroPubl Biol 2025;2025
PMID:8493104 - Study of multiple fibrillarin mRNAs reveals that 3' end formation in Schizosaccharomyces pombe is sensitive to cold shock.
Girard JP et al. Nucleic Acids Res 1993 Apr 25;21(8):1881-7
PMID:23861937 - Genome-wide screening for genes associated with valproic acid sensitivity in fission yeast.
Zhang L et al. PLoS One 2013;8(7):e68738
PMID:22403715 - Regulation of fission yeast morphogenesis by PP2A activator pta2.
Bernal M et al. PLoS One 2012;7(3):e32823
PMID:16823372 - ORFeome cloning and global analysis of protein localization in the fission yeast Schizosaccharomyces pombe.
Matsuyama A et al. Nat Biotechnol 2006 Jul;24(7):841-7
PMID:32101745 - Dense Transposon Integration Reveals Essential Cleavage and Polyadenylation Factors Promote Heterochromatin Formation.
Lee SY et al. Cell Rep 2020 Feb 25;30(8):2686-2698.e8
PMID:21652630 - Characterization of Mug33 reveals complementary roles for actin cable-dependent transport and exocyst regulators in fission yeast exocytosis.
Snaith HA et al. J Cell Sci 2011 Jul 01;124(Pt 13):2187-99
PMID:28945192 - Regulated Ire1-dependent mRNA decay requires no-go mRNA degradation to maintain endoplasmic reticulum homeostasis in S. pombe .
Guydosh NR et al. Elife 2017 Sep 25;6
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:17690116 - Cdc18/CDC6 activates the Rad3-dependent checkpoint in the fission yeast.
Fersht N et al. Nucleic Acids Res 2007;35(16):5323-37
PMID:9211982 - Ultrastructural changes in the Schizosaccharomyces pombe nucleolus following the disruption of the gar2+ gene, which encodes a nucleolar protein structurally related to nucleolin.
Léger-Silvestre I et al. Chromosoma 1997 Jun;105(7-8):542-52
PMID:31064814 - Proximity-dependent biotinylation mediated by TurboID to identify protein-protein interaction networks in yeast.
Larochelle M et al. J Cell Sci 2019 May 31;132(11)
PMID:33313903 - Ribosome profiling reveals ribosome stalling on tryptophan codons and ribosome queuing upon oxidative stress in fission yeast.
Rubio A et al. Nucleic Acids Res 2021 Jan 11;49(1):383-399
PMID:34496258 - Transcription and chromatin-based surveillance mechanism controls suppression of cryptic antisense transcription.
Heo DH et al. Cell Rep 2021 Sep 07;36(10):109671
PMID:39476757 - Characterization of Ksg1 protein kinase-dependent phosphoproteome in the fission yeast S. pombe.
Cipak L et al. Biochem Biophys Res Commun 2024 Oct 25;736:150895
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:37970674 - SUMOylation regulates Lem2 function in centromere clustering and silencing.
Strachan J et al. J Cell Sci 2023 Dec 01;136(23)
PMID:34296454 - The TOR-dependent phosphoproteome and regulation of cellular protein synthesis.
Mak T et al. EMBO J 2021 Aug 16;40(16):e107911
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: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:27298342 - Identification of S-phase DNA damage-response targets in fission yeast reveals conservation of damage-response networks.
Willis NA et al. Proc Natl Acad Sci U S A 2016 Jun 28;113(26):E3676-85
PMID:23101633 - Quantitative analysis of fission yeast transcriptomes and proteomes in proliferating and quiescent cells.
Marguerat S et al. Cell 2012 Oct 26;151(3):671-83
PMID:28218250 - Chromatin remodeller Fun30 Fft3 induces nucleosome disassembly to facilitate RNA polymerase II elongation.
Lee J et al. Nat Commun 2017 Feb 20;8:14527
PMID:23658229 - Red5 and three nuclear pore components are essential for efficient suppression of specific mRNAs during vegetative growth of fission yeast.
Sugiyama T et al. Nucleic Acids Res 2013 Jul;41(13):6674-86
PMID:37787768 - Broad functional profiling of fission yeast proteins using phenomics and machine learning.
Rodríguez-López M et al. Elife 2023 Oct 03;12
PMID:21712547 - Mitotic substrates of the kinase aurora with roles in chromatin regulation identified through quantitative phosphoproteomics of fission yeast.
Koch A et al. Sci Signal 2011 Jun 28;4(179):rs6
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:17213188 - Regulation of the nuclear poly(A)-binding protein by arginine methylation in fission yeast.
Perreault A et al. J Biol Chem 2007 Mar 09;282(10):7552-62
PMID:20403971 - Proteomic and functional analysis of the noncanonical poly(A) polymerase Cid14.
Keller C et al. RNA 2010 Jun;16(6):1124-9
PMID:25375137 - Systematic analysis of the role of RNA-binding proteins in the regulation of RNA stability.
Hasan A et al. PLoS Genet 2014 Nov;10(11):e1004684
PMID:21511999 - Comparative functional genomics of the fission yeasts.
Rhind N et al. Science 2011 May 20;332(6032):930-6
PMID:20473289 - Analysis of a genome-wide set of gene deletions in the fission yeast Schizosaccharomyces pombe.
Kim DU et al. Nat Biotechnol 2010 Jun;28(6):617-623
PMID:9211981 - Mitosis-specific phosphorylation of gar2, a fission yeast nucleolar protein structurally related to nucleolin.
Gulli MP et al. Chromosoma 1997 Jun;105(7-8):532-41
PMID:38051102 - Fission Yeast TORC1 Promotes Cell Proliferation through Sfp1, a Transcription Factor Involved in Ribosome Biogenesis.
Tai YT et al. Mol Cell Biol 2023 Dec 05;:1-18
PMID:19430462 - Decapping is preceded by 3' uridylation in a novel pathway of bulk mRNA turnover.
Rissland OS et al. Nat Struct Mol Biol 2009 Jun;16(6):616-23
PMID:21760946 - Identification of genes affecting the toxicity of anti-cancer drug bortezomib by genome-wide screening in S. pombe.
Takeda K et al. PLoS One 2011;6(7):e22021
PMID:27401558 - The Nrd1-like protein Seb1 coordinates cotranscriptional 3' end processing and polyadenylation site selection.
Lemay JF et al. Genes Dev 2016 Jul 01;30(13):1558-72
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:29899117 - Ccp1 modulates epigenetic stability at centromeres and affects heterochromatin distribution in Schizosaccharomyces pombe .
Lu M et al. J Biol Chem 2018 Aug 03;293(31):12068-12080
PMID:22681890 - Hierarchical modularity and the evolution of genetic interactomes across species.
Ryan CJ et al. Mol Cell 2012 Jun 08;46(5):691-704
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:26167880 - SR protein kinases promote splicing of nonconsensus introns.
Lipp JJ et al. Nat Struct Mol Biol 2015 Aug;22(8):611-7
PMID:39705284 - Proteomic and phosphoproteomic analyses reveal that TORC1 is reactivated by pheromone signaling during sexual reproduction in fission yeast.
Bérard M et al. PLoS Biol 2024 Dec 20;22(12):e3002963
PMID:30321377 - Proteomic profiling and functional characterization of post-translational modifications of the fission yeast RNA exosome.
Telekawa C et al. Nucleic Acids Res 2018 Nov 30;46(21):11169-11183
PMID:22540037 - Predicting the fission yeast protein interaction network.
Pancaldi V et al. G3 (Bethesda) 2012 Apr;2(4):453-67
PMID:24634168 - Proteome-wide search for PP2A substrates in fission yeast.
Bernal M et al. Proteomics 2014 Jun;14(11):1367-80
PMID:9693363 - The role of the Schizosaccharomyces pombe gar2 protein in nucleolar structure and function depends on the concerted action of its highly charged N terminus and its RNA-binding domains.
Sicard H et al. Mol Biol Cell 1998 Aug;9(8):2011-23
PMID:7596817 - gar2 is a nucleolar protein from Schizosaccharomyces pombe required for 18S rRNA and 40S ribosomal subunit accumulation.
Gulli MP et al. Nucleic Acids Res 1995 Jun 11;23(11):1912-8
PMID:37913773 - TOR inactivation triggers heterochromatin formation in rDNA during glucose starvation.
Hirai H et al. Cell Rep 2023 Nov 28;42(11):113320