protein coding gene - rad9 (SPAC664.07c) - checkpoint clamp complex protein Rad9

Gene summary

Standard name

rad9

Systematic ID

SPAC664.07c

Product

checkpoint clamp complex protein Rad9

Organism

Schizosaccharomyces pombe (fission yeast)

UniProt ID

P26306

ORFeome ID

23/23C09

Characterisation status

biological role published

Feature type

mRNA gene

Genomic location

chromosome I: 1713920..1715716 reverse strand

Annotation

Protein features

Orthologs

• RAD9B (HGNC:21700) Homo sapiens
• RAD9A (HGNC:9827) Homo sapiens
• rad9 (SJAG_01605) Schizosaccharomyces japonicus
• DDC1 (YPL194W) Saccharomyces cerevisiae

References / Literature

PMID:21095590 - DDK phosphorylates checkpoint clamp component Rad9 and promotes its release from damaged chromatin.

Furuya K et al. Mol Cell 2010 Nov 24;40(4):606-18

PMID:26499799 - Probing the Mec1ATR Checkpoint Activation Mechanism with Small Peptides.

Wanrooij PH et al. J Biol Chem 2016 Jan 01;291(1):393-401

PMID:24943839 - A role for nuclear envelope-bridging complexes in homology-directed repair.

Swartz RK et al. Mol Biol Cell 2014 Aug 15;25(16):2461-71

PMID:8846774 - Fission yeast rad17: a homologue of budding yeast RAD24 that shares regions of sequence similarity with DNA polymerase accessory proteins.

Griffiths DJ et al. EMBO J 1995 Dec 01;14(23):5812-23

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: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:10521402 - Meiotic DNA replication checkpoint control in fission yeast.

Murakami H et al. Genes Dev 1999 Oct 01;13(19):2581-93

PMID:18414064 - Schizosaccharomyces pombe Orc5 plays multiple roles in the maintenance of genome stability throughout the cell cycle.

Kato H et al. Cell Cycle 2008 Apr 15;7(8):1085-96

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:16371652 - Production of reactive oxygen species in response to replication stress and inappropriate mitosis in fission yeast.

Marchetti MA et al. J Cell Sci 2006 Jan 01;119(Pt 1):124-31

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: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:26746798 - Inner nuclear membrane protein Lem2 facilitates Rad3-mediated checkpoint signaling under replication stress induced by nucleotide depletion in fission yeast.

Xu YJ Cell Signal 2016 Apr;28(4):235-45

PMID:10683155 - A novel mutant allele of the chromatin-bound fission yeast checkpoint protein Rad17 separates the DNA structure checkpoints.

Griffiths D et al. J Cell Sci 2000 Mar;113 ( Pt 6):1075-88

PMID:28441348 - Deoxynucleoside Salvage in Fission Yeast Allows Rescue of Ribonucleotide Reductase Deficiency but Not Spd1-Mediated Inhibition of Replication.

Fleck O et al. Genes (Basel) 2017 Apr 25;8(5)

PMID:31332096 - A tel2 Mutation That Destabilizes the Tel2-Tti1-Tti2 Complex Eliminates Rad3 ATR Kinase Signaling in the DNA Replication Checkpoint and Leads to Telomere Shortening in Fission Yeast.

Xu YJ et al. Mol Cell Biol 2019 Oct 15;39(20)

PMID:7254221 - Spontaneous and UV-induced recombination in radiation-sensitive mutants of Schizosaccharomyces pombe.

Grossenbacher-Grunder AM et al. Mutat Res 1981 Mar;81(1):37-48

PMID:166019 - Genetic control of radiation sensitivity in Schizosaccharomyces pombe.

Nasim A et al. Genetics 1975 Apr;79(4):573-82

PMID:12062100 - Cid13 is a cytoplasmic poly(A) polymerase that regulates ribonucleotide reductase mRNA.

Saitoh S et al. Cell 2002 May 31;109(5):563-73

PMID:9566891 - Fission yeast rad12+ regulates cell cycle checkpoint control and is homologous to the Bloom's syndrome disease gene.

Davey S et al. Mol Cell Biol 1998 May;18(5):2721-8

PMID:33260998 - High-Throughput Flow Cytometry Combined with Genetic Analysis Brings New Insights into the Understanding of Chromatin Regulation of Cellular Quiescence.

Zahedi Y et al. Int J Mol Sci 2020 Nov 27;21(23)

PMID:9739083 - Cdc18p can block mitosis by two independent mechanisms.

Greenwood E et al. J Cell Sci 1998 Oct;111 ( Pt 20):3101-8

PMID:1852603 - Cloning and characterisation of the rad9 DNA repair gene from Schizosaccharomyces pombe.

Murray JM et al. Nucleic Acids Res 1991 Jul 11;19(13):3525-31

PMID:11027257 - Analysis of fission yeast primase defines the checkpoint responses to aberrant S phase initiation.

Tan S et al. Mol Cell Biol 2000 Nov;20(21):7853-66

PMID:1563349 - Checkpoint controls in Schizosaccharomyces pombe: rad1.

Rowley R et al. EMBO J 1992 Apr;11(4):1335-42

PMID:31538680 - The fission yeast FHIT homolog affects checkpoint control of proliferation and is regulated by mitochondrial electron transport.

Sjölander JJ et al. Cell Biol Int 2020 Feb;44(2):412-423

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:19205745 - Redundant roles of Srs2 helicase and replication checkpoint in survival and rDNA maintenance in Schizosaccharomyces pombe.

Yasuhira S Mol Genet Genomics 2009 May;281(5):497-509

PMID:29121084 - Fission yeast strains with circular chromosomes require the 9-1-1 checkpoint complex for the viability in response to the anti-cancer drug 5-fluorodeoxyuridine.

Shamim HM et al. PLoS One 2017;12(11):e0187775

PMID:9563836 - Mutant PCNA alleles are associated with cdc phenotypes and sensitivity to DNA damage in fission yeast.

Arroyo MP et al. Mol Gen Genet 1998 Mar;257(5):505-18

PMID:21561865 - The phosphorylation network for efficient activation of the DNA replication checkpoint in fission yeast.

Yue M et al. J Biol Chem 2011 Jul 01;286(26):22864-74

PMID:22761595 - The Rad4(TopBP1) ATR-activation domain functions in G1/S phase in a chromatin-dependent manner.

Lin SJ et al. PLoS Genet 2012 Jun;8(6):e1002801

PMID:24074952 - Phosphorylation-dependent assembly and coordination of the DNA damage checkpoint apparatus by Rad4(TopBP1).

Qu M et al. Mol Cell 2013 Sep 26;51(6):723-736

PMID:28410370 - A systematic screen for morphological abnormalities during fission yeast sexual reproduction identifies a mechanism of actin aster formation for cell fusion.

Dudin O et al. PLoS Genet 2017 Apr;13(4):e1006721

PMID:9524127 - Hus1p, a conserved fission yeast checkpoint protein, interacts with Rad1p and is phosphorylated in response to DNA damage.

Kostrub CF et al. EMBO J 1998 Apr 01;17(7):2055-66

PMID:10648611 - Characterization of Schizosaccharomyces pombe Hus1: a PCNA-related protein that associates with Rad1 and Rad9.

Caspari T et al. Mol Cell Biol 2000 Feb;20(4):1254-62

PMID:18931302 - Significant conservation of synthetic lethal genetic interaction networks between distantly related eukaryotes.

Dixon SJ et al. Proc Natl Acad Sci U S A 2008 Oct 28;105(43):16653-8

PMID:12514100 - Checkpoint activation regulates mutagenic translesion synthesis.

Kai M et al. Genes Dev 2003 Jan 01;17(1):64-76

PMID:12930957 - Nuclear factories for signalling and repairing DNA double strand breaks in living fission yeast.

Meister P et al. Nucleic Acids Res 2003 Sep 01;31(17):5064-73

PMID:12455694 - mcl1+, the Schizosaccharomyces pombe homologue of CTF4, is important for chromosome replication, cohesion, and segregation.

Williams DR et al. Eukaryot Cell 2002 Oct;1(5):758-73

PMID:19264558 - Screening a genome-wide S. pombe deletion library identifies novel genes and pathways involved in genome stability maintenance.

Deshpande GP et al. DNA Repair (Amst) 2009 May 01;8(5):672-9

PMID:15533944 - Interaction of checkpoint proteins Hus1/Rad1/Rad9 with DNA base excision repair enzyme MutY homolog in fission yeast, Schizosaccharomyces pombe.

Chang DY et al. J Biol Chem 2005 Jan 07;280(1):408-17

GO_REF:0000033 - Annotation inferences using phylogenetic trees

PMID:10996309 - Schizosaccharomyces pombe Rad9 contains a BH3-like region and interacts with the anti-apoptotic protein Bcl-2.

Komatsu K et al. FEBS Lett 2000 Sep 15;481(2):122-6

PMID:8290359 - Cloning and characterisation of the Schizosaccharomyces pombe rad8 gene, a member of the SNF2 helicase family.

Doe CL et al. Nucleic Acids Res 1993 Dec 25;21(25):5964-71

PMID:24192486 - The conserved Fanconi anemia nuclease Fan1 and the SUMO E3 ligase Pli1 act in two novel Pso2-independent pathways of DNA interstrand crosslink repair in yeast.

Fontebasso Y et al. DNA Repair (Amst) 2013 Dec;12(12):1011-23

PMID:15643072 - Mcl1p is a polymerase alpha replication accessory factor important for S-phase DNA damage survival.

Williams DR et al. Eukaryot Cell 2005 Jan;4(1):166-77

PMID:15972456 - Brc1-mediated DNA repair and damage tolerance.

Sheedy DM et al. Genetics 2005 Oct;171(2):457-68

PMID:22645654 - Opposing role of condensin hinge against replication protein A in mitosis and interphase through promoting DNA annealing.

Akai Y et al. Open Biol 2011 Dec;1(4):110023

PMID:10757807 - Cid1, a fission yeast protein required for S-M checkpoint control when DNA polymerase delta or epsilon is inactivated.

Wang SW et al. Mol Cell Biol 2000 May;20(9):3234-44

PMID:18667534 - Activation of the DNA damage checkpoint in mutants defective in DNA replication initiation.

Yin L et al. Mol Biol Cell 2008 Oct;19(10):4374-82

PMID:22797921 - Heat induction of a novel Rad9 variant from a cryptic translation initiation site reduces mitotic commitment.

Janes S et al. J Cell Sci 2012 Oct 01;125(Pt 19):4487-97

PMID:11073995 - Rdp1, a novel zinc finger protein, regulates the DNA damage response of rhp51(+) from Schizosaccharomyces pombe.

Shim YS et al. Mol Cell Biol 2000 Dec;20(23):8958-68

PMID:12196391 - Telomere binding of checkpoint sensor and DNA repair proteins contributes to maintenance of functional fission yeast telomeres.

Nakamura TM et al. Genetics 2002 Aug;161(4):1437-52

PMID:16723501 - Rad4TopBP1, a scaffold protein, plays separate roles in DNA damage and replication checkpoints and DNA replication.

Taricani L et al. Mol Biol Cell 2006 Aug;17(8):3456-68

PMID:17515930 - Rad3-dependent phosphorylation of the checkpoint clamp regulates repair-pathway choice.

Kai M et al. Nat Cell Biol 2007 Jun;9(6):691-7

PMID:21945095 - Mcm10 interacts with Rad4/Cut5(TopBP1) and its association with origins of DNA replication is dependent on Rad4/Cut5(TopBP1).

Taylor M et al. DNA Repair (Amst) 2011 Nov 10;10(11):1154-63

PMID:24587136 - The oxidative stress responsive transcription factor Pap1 confers DNA damage resistance on checkpoint-deficient fission yeast cells.

Belfield C et al. PLoS One 2014;9(2):e89936

PMID:8536960 - Extragenic suppressors of Schizosaccharomyces pombe rad9 mutations uncouple radioresistance and hydroxyurea sensitivity from cell cycle checkpoint control.

Lieberman HB Genetics 1995 Sep;141(1):107-17

PMID:12589755 - The fission yeast spSet1p is a histone H3-K4 methyltransferase that functions in telomere maintenance and DNA repair in an ATM kinase Rad3-dependent pathway.

Kanoh J et al. J Mol Biol 2003 Feb 28;326(4):1081-94

PMID:11058134 - Mutant alleles of Schizosaccharomyces pombe rad9(+) alter hydroxyurea resistance, radioresistance and checkpoint control.

Hang H et al. Nucleic Acids Res 2000 Nov 01;28(21):4340-9

PMID:24663817 - Rad4 mainly functions in Chk1-mediated DNA damage checkpoint pathway as a scaffold protein in the fission yeast Schizosaccharomyces pombe.

Yue M et al. PLoS One 2014;9(3):e92936

PMID:9693370 - Mutational effect of fission yeast polalpha on cell cycle events.

Bhaumik D et al. Mol Biol Cell 1998 Aug;9(8):2107-23

PMID:18818364 - Conservation and rewiring of functional modules revealed by an epistasis map in fission yeast.

Roguev A et al. Science 2008 Oct 17;322(5900):405-10

PMID:24806966 - The extent of error-prone replication restart by homologous recombination is controlled by Exo1 and checkpoint proteins.

Tsang E et al. J Cell Sci 2014 Jul 01;127(Pt 13):2983-94

PMID:15155581 - Chk1 activation requires Rad9 S/TQ-site phosphorylation to promote association with C-terminal BRCT domains of Rad4TOPBP1.

Furuya K et al. Genes Dev 2004 May 15;18(10):1154-64

PMID:17426133 - The checkpoint clamp, Rad9-Rad1-Hus1 complex, preferentially stimulates the activity of apurinic/apyrimidinic endonuclease 1 and DNA polymerase beta in long patch base excision repair.

Gembka A et al. Nucleic Acids Res 2007;35(8):2596-608

PMID:10954610 - The stress-activated MAP kinase Sty1/Spc1 and a 3'-regulatory element mediate UV-induced expression of the uvi15(+) gene at the post-transcriptional level.

Kim M et al. Nucleic Acids Res 2000 Sep 01;28(17):3392-402

PMID:21493688 - Cdt1 proteolysis is promoted by dual PIP degrons and is modulated by PCNA ubiquitylation.

Guarino E et al. Nucleic Acids Res 2011 Aug;39(14):5978-90

PMID:37200372 - Comprehensive mutational analysis of the checkpoint signaling function of Rpa1/Ssb1 in fission yeast.

Xu YJ et al. PLoS Genet 2023 May;19(5):e1010691

PMID:31262821 - Replication fork stalling elicits chromatin compaction for the stability of stalling replication forks.

Feng G et al. Proc Natl Acad Sci U S A 2019 Jul 16;116(29):14563-14572

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:8943031 - A human homolog of the Schizosaccharomyces pombe rad9+ checkpoint control gene.

Lieberman HB et al. Proc Natl Acad Sci U S A 1996 Nov 26;93(24):13890-5

PMID:12839619 - High dosage Rhp51 suppression of the MMS sensitivity of DNA structure checkpoint mutants reveals a relationship between Crb2 and Rhp51.

Smeets MF et al. Genes Cells 2003 Jul;8(7):573-86

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:9092625 - Characterisation of Schizosaccharomyces pombe rad31, a UBA-related gene required for DNA damage tolerance.

Shayeghi M et al. Nucleic Acids Res 1997 Mar 15;25(6):1162-9

PMID:17725619 - Protective role and regulation of Rad9 from the fission yeast Schizosaccharomyces pombe.

Kang MH et al. FEMS Microbiol Lett 2007 Oct;275(2):270-7

PMID:9398669 - A WD repeat protein controls the cell cycle and differentiation by negatively regulating Cdc2/B-type cyclin complexes.

Yamaguchi S et al. Mol Biol Cell 1997 Dec;8(12):2475-86

PMID:9487130 - Regulation of telomere length by checkpoint genes in Schizosaccharomyces pombe.

Dahlen M et al. Mol Biol Cell 1998 Mar;9(3):611-21

PMID:38718864 - H2B oncohistones cause homologous recombination defect and genomic instability through reducing H2B monoubiquitination in Schizosaccharomyces pombe.

Qin B et al. J Biol Chem 2024 May 06;:107345

PMID:9766521 - Molecular cloning and tissue-specific expression of Mrad9, a murine orthologue of the Schizosaccharomyces pombe rad9+ checkpoint control gene.

Hang H et al. J Cell Physiol 1998 Nov;177(2):241-7

PMID:22792081 - Phosphorylation-dependent interactions between Crb2 and Chk1 are essential for DNA damage checkpoint.

Qu M et al. PLoS Genet 2012 Jul;8(7):e1002817

PMID:9658208 - Sensitivity to cisplatin and platinum-containing compounds of Schizosaccharomyces pombe rad mutants.

Perego P et al. Mol Pharmacol 1998 Jul;54(1):213-9

PMID:21511999 - Comparative functional genomics of the fission yeasts.

Rhind N et al. Science 2011 May 20;332(6032):930-6

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

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

PMID:23677513 - A role for Myh1 in DNA repair after treatment with strand-breaking and crosslinking chemotherapeutic agents.

Jansson K et al. Environ Mol Mutagen 2013 Jun;54(5):327-37

PMID:12032307 - A single unbranched S-phase DNA damage and replication fork blockage checkpoint pathway.

Marchetti MA et al. Proc Natl Acad Sci U S A 2002 May 28;99(11):7472-7

PMID:1588907 - Molecular cloning and analysis of Schizosaccharomyces pombe rad9, a gene involved in DNA repair and mutagenesis.

Lieberman HB et al. Mol Gen Genet 1992 Apr;232(3):367-76

PMID:18378696 - Molecular characterization of the role of the Schizosaccharomyces pombe nip1+/ctp1+ gene in DNA double-strand break repair in association with the Mre11-Rad50-Nbs1 complex.

Akamatsu Y et al. Mol Cell Biol 2008 Jun;28(11):3639-51

PMID:25795664 - Genetic Interaction Landscape Reveals Critical Requirements for Schizosaccharomyces pombe Brc1 in DNA Damage Response Mutants.

Sánchez A et al. G3 (Bethesda) 2015 Mar 19;5(5):953-62

PMID:15548596 - Analysis of mutant phenotypes and splicing defects demonstrates functional collaboration between the large and small subunits of the essential splicing factor U2AF in vivo.

Webb CJ et al. Mol Biol Cell 2005 Feb;16(2):584-96

PMID:15498101 - Germinating fission yeast spores delay in G1 in response to UV irradiation.

Nilssen EA et al. BMC Cell Biol 2004 Oct 21;5(1):40

PMID:18653539 - Caspase-dependent and -independent lipotoxic cell-death pathways in fission yeast.

Low CP et al. J Cell Sci 2008 Aug 15;121(Pt 16):2671-84

PMID:18675827 - The tumor suppressor homolog in fission yeast, myh1(+), displays a strong interaction with the checkpoint gene rad1(+).

Jansson K et al. Mutat Res 2008 Sep 26;644(1-2):48-55

PMID:10559981 - A Rad3-Rad26 complex responds to DNA damage independently of other checkpoint proteins.

Edwards RJ et al. Nat Cell Biol 1999 Nov;1(7):393-8

PMID:29259000 - Genes Important for Schizosaccharomyces pombe Meiosis Identified Through a Functional Genomics Screen.

Blyth J et al. Genetics 2018 Feb;208(2):589-603

PMID:30824696 - Systematic analysis reveals the prevalence and principles of bypassable gene essentiality.

Li J et al. Nat Commun 2019 Mar 01;10(1):1002

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:23173672 - Identification of novel genes involved in DNA damage response by screening a genome-wide Schizosaccharomyces pombe deletion library.

Pan X et al. BMC Genomics 2012 Nov 23;13:662

PMID:30148840 - Tdp1 processes chromate-induced single-strand DNA breaks that collapse replication forks.

Ganguly A et al. PLoS Genet 2018 Aug;14(8):e1007595