Gene Symbol: MRC1
Description: chromatin-modulating protein MRC1
Alias: YCL060C, chromatin-modulating protein MRC1
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Robert T, Dervins D, Fabre F, Gangloff S. Mrc1 and Srs2 are major actors in the regulation of spontaneous crossover. EMBO J. 2006;25:2837-46 pubmed
    ..Our results support the view that Mrc1 plays a specific role in DNA replication, promoting the Srs2 recruitment to PCNA independently of checkpoint ..
  2. Grandin N, Charbonneau M. Mrc1, a non-essential DNA replication protein, is required for telomere end protection following loss of capping by Cdc13, Yku or telomerase. Mol Genet Genomics. 2007;277:685-99 pubmed
    ..Here, we report that in the absence of Mrc1, a component of the replication forks, telomeres of cdc13 or yku70 mutants exhibited increased degradation, while ..
  3. Hu F, Alcasabas A, Elledge S. Asf1 links Rad53 to control of chromatin assembly. Genes Dev. 2001;15:1061-6 pubmed
    ..Thus, checkpoint pathways directly regulate chromatin assembly to promote survival in response to DNA damage and replication blocks. ..
  4. Hodgson B, Calzada A, Labib K. Mrc1 and Tof1 regulate DNA replication forks in different ways during normal S phase. Mol Biol Cell. 2007;18:3894-902 pubmed
    The Mrc1 and Tof1 proteins are conserved throughout evolution, and in budding yeast they are known to associate with the MCM helicase and regulate the progression of DNA replication forks...
  5. Bjergbaek L, Cobb J, Tsai Pflugfelder M, Gasser S. Mechanistically distinct roles for Sgs1p in checkpoint activation and replication fork maintenance. EMBO J. 2005;24:405-17 pubmed
    ..function, the Sgs1p/Top3p complex acts in parallel to the Claspin-related adaptor, Mrc1p, although the sgs1 and mrc1 mutations are epistatic for Rad53p activation...
  6. Komata M, Bando M, Araki H, Shirahige K. The direct binding of Mrc1, a checkpoint mediator, to Mcm6, a replication helicase, is essential for the replication checkpoint against methyl methanesulfonate-induced stress. Mol Cell Biol. 2009;29:5008-19 pubmed publisher
    b>Mrc1 plays a role in mediating the DNA replication checkpoint. We surveyed replication elongation proteins that interact directly with Mrc1 and identified a replicative helicase, Mcm6, as a specific Mrc1-binding protein...
  7. Razidlo D, Lahue R. Mrc1, Tof1 and Csm3 inhibit CAG.CTG repeat instability by at least two mechanisms. DNA Repair (Amst). 2008;7:633-40 pubmed publisher
    ..CTG repeat contraction rates were sought using a disruption library. This screen identified Mrc1, the homolog of human Claspin, which mediates the replication and DNA damage checkpoints, and also couples the ..
  8. Mohanty B, Bairwa N, Bastia D. The Tof1p-Csm3p protein complex counteracts the Rrm3p helicase to control replication termination of Saccharomyces cerevisiae. Proc Natl Acad Sci U S A. 2006;103:897-902 pubmed
    ..Other genes such as MRC1, WSS1, and PSY2 that are also involved in the MRC1 checkpoint pathway were not involved in this global control...
  9. Alabert C, Bianco J, Pasero P. Differential regulation of homologous recombination at DNA breaks and replication forks by the Mrc1 branch of the S-phase checkpoint. EMBO J. 2009;28:1131-41 pubmed publisher replication stress also depends on the Mec1 kinase, which activates the DNA replication checkpoint in an Mrc1-dependent manner in response to fork arrest...

More Information


  1. Osborn A, Elledge S. Mrc1 is a replication fork component whose phosphorylation in response to DNA replication stress activates Rad53. Genes Dev. 2003;17:1755-67 pubmed
    ..In budding yeast, this pathway includes the kinases Mec1 and Rad53. Here we report that the Mediator protein Mrc1, which is required for normal DNA replication and for activation of Rad53, is present at replication forks...
  2. Schmidt K, Kolodner R. Suppression of spontaneous genome rearrangements in yeast DNA helicase mutants. Proc Natl Acad Sci U S A. 2006;103:18196-201 pubmed
    ..DNA damage sensors and results from activation of the Mec1/Rad9-dependent DNA damage response rather than the Mrc1-dependent replication stress response...
  3. Lou H, Komata M, Katou Y, Guan Z, Reis C, Budd M, et al. Mrc1 and DNA polymerase epsilon function together in linking DNA replication and the S phase checkpoint. Mol Cell. 2008;32:106-17 pubmed publisher
    Yeast Mrc1, ortholog of metazoan Claspin, is both a central component of normal DNA replication forks and a mediator of the S phase checkpoint...
  4. Calzada A, Hodgson B, Kanemaki M, Bueno A, Labib K. Molecular anatomy and regulation of a stable replisome at a paused eukaryotic DNA replication fork. Genes Dev. 2005;19:1905-19 pubmed
    ..The budding yeast proteins Mrc1 and Tof1 associate with the putative MCM-Cdc45 helicase and limit progression of the replisome when nucleotides are ..
  5. Chen S, Zhou H. Reconstitution of Rad53 activation by Mec1 through adaptor protein Mrc1. J Biol Chem. 2009;284:18593-604 pubmed publisher
    ..Using an activity-based assay for Rad53, we found that Mrc1, a replication fork-associated protein, cooperates with Mec1 to activate Rad53 directly...
  6. Pike B, Tenis N, Heierhorst J. Rad53 kinase activation-independent replication checkpoint function of the N-terminal forkhead-associated (FHA1) domain. J Biol Chem. 2004;279:39636-44 pubmed important mechanistic difference to the homologous Schizosaccharomyces pombe FHA domain that is required for Mrc1-dependent activation of the corresponding Cds1 kinase...
  7. Psakhye I, Jentsch S. Protein group modification and synergy in the SUMO pathway as exemplified in DNA repair. Cell. 2012;151:807-820 pubmed publisher
    ..We propose that SUMOylation may thus often target a protein group rather than individual proteins, whereas localized modification enzymes and highly specific triggers ensure specificity. ..
  8. Xu H, Boone C, Brown G. Genetic dissection of parallel sister-chromatid cohesion pathways. Genetics. 2007;176:1417-29 pubmed
    ..These data defined two cohesion pathways, one containing CSM3, TOF1, CTF4, and CHL1, and the second containing MRC1, CTF18, CTF8, and DCC1...
  9. Mimura S, Komata M, Kishi T, Shirahige K, Kamura T. SCF(Dia2) regulates DNA replication forks during S-phase in budding yeast. EMBO J. 2009;28:3693-705 pubmed publisher
    ..Using modified yeast two-hybrid screening, we have identified components of the replisome (Mrc1, Ctf4 and Mcm2), as Dia2-binding proteins. Mrc1 and Ctf4 were ubiquitinated by SCF(Dia2) both in vivo and in vitro...
  10. Katou Y, Kanoh Y, Bando M, Noguchi H, Tanaka H, Ashikari T, et al. S-phase checkpoint proteins Tof1 and Mrc1 form a stable replication-pausing complex. Nature. 2003;424:1078-83 pubmed
    ..Here we demonstrate that the checkpoint regulatory proteins Tof1 and Mrc1 interact directly with the DNA replication machinery in Saccharomyces cerevisiae...
  11. Bando M, Katou Y, Komata M, Tanaka H, Itoh T, Sutani T, et al. Csm3, Tof1, and Mrc1 form a heterotrimeric mediator complex that associates with DNA replication forks. J Biol Chem. 2009;284:34355-65 pubmed publisher
    b>Mrc1 (mediator of replication checkpoint), Tof1 (topoisomerase I interacting factor), and Csm3 (chromosome segregation in meiosis) are checkpoint-mediator proteins that function during DNA replication and activate the effector kinase ..
  12. Alcasabas A, Osborn A, Bachant J, Hu F, Werler P, Bousset K, et al. Mrc1 transduces signals of DNA replication stress to activate Rad53. Nat Cell Biol. 2001;3:958-65 pubmed
    ..We describe a conserved protein Mrc1, mediator of the replication checkpoint, required for activation of ScRad53 and SpCds1 during replication stress...
  13. Tourrière H, Versini G, Cordon Preciado V, Alabert C, Pasero P. Mrc1 and Tof1 promote replication fork progression and recovery independently of Rad53. Mol Cell. 2005;19:699-706 pubmed
    ..The critical role of Mrc1p in HU is therefore to promote fork recovery in a Rad53p-independent manner, presumably through the formation of a stable fork-pausing complex. ..
  14. Szyjka S, Viggiani C, Aparicio O. Mrc1 is required for normal progression of replication forks throughout chromatin in S. cerevisiae. Mol Cell. 2005;19:691-7 pubmed
    b>Mrc1 associates with replication forks, where it transmits replication stress signals and is required for normal replisome pausing in response to nucleotide depletion...
  15. Morohashi H, Maculins T, Labib K. The amino-terminal TPR domain of Dia2 tethers SCF(Dia2) to the replisome progression complex. Curr Biol. 2009;19:1943-9 pubmed publisher
    ..This interaction requires the RPC components Mrc1 and Ctf4, both of which associate with a tetratricopeptide repeat (TPR) domain located at the amino terminus of ..
  16. Putnam C, Hayes T, Kolodner R. Post-replication repair suppresses duplication-mediated genome instability. PLoS Genet. 2010;6:e1000933 pubmed publisher
    ..Our analysis is consistent with models in which PRR prevents replication damage from becoming double strand breaks (DSBs) and/or regulates the activity of HR on DSBs. ..
  17. Hoch N, Chen E, Buckland R, Wang S, Fazio A, Hammet A, et al. Molecular basis of the essential s phase function of the rad53 checkpoint kinase. Mol Cell Biol. 2013;33:3202-13 pubmed publisher
    ..Our findings indicate that the essential S phase function of Rad53 is comprised by the combination of its role in regulating basal dNTP levels and its compensatory kinase function if dNTP levels are perturbed. ..
  18. Keogh M, Kim J, Downey M, Fillingham J, Chowdhury D, Harrison J, et al. A phosphatase complex that dephosphorylates gammaH2AX regulates DNA damage checkpoint recovery. Nature. 2006;439:497-501 pubmed
    ..The dephosphorylation of gammaH2AX by the HTP-C is necessary for efficient recovery from the DNA damage checkpoint. ..
  19. De Piccoli G, Katou Y, Itoh T, Nakato R, Shirahige K, Labib K. Replisome stability at defective DNA replication forks is independent of S phase checkpoint kinases. Mol Cell. 2012;45:696-704 pubmed publisher
  20. Torres J, Bessler J, Zakian V. Local chromatin structure at the ribosomal DNA causes replication fork pausing and genome instability in the absence of the S. cerevisiae DNA helicase Rrm3p. Genes Dev. 2004;18:498-503 pubmed
  21. Duch A, Felipe Abrio I, Barroso S, Yaakov G, Garcia Rubio M, Aguilera A, et al. Coordinated control of replication and transcription by a SAPK protects genomic integrity. Nature. 2013;493:116-9 pubmed publisher
    ..Hog1 interacts with and phosphorylates Mrc1, a component of the replication complex...
  22. Ohouo P, Bastos de Oliveira F, Liu Y, Ma C, Smolka M. DNA-repair scaffolds dampen checkpoint signalling by counteracting the adaptor Rad9. Nature. 2013;493:120-4 pubmed publisher
    ..Our findings show that DDC signalling is monitored and modulated through the direct action of DNA-repair factors. ..
  23. Reha Krantz L, Siddique M, Murphy K, Tam A, O Carroll M, Lou S, et al. Drug-sensitive DNA polymerase ? reveals a role for mismatch repair in checkpoint activation in yeast. Genetics. 2011;189:1211-24 pubmed publisher
    ..Mec1, Rad9, the DNA damage clamp Ddc1-Rad17-Mec3, and the Rad24-dependent clamp loader, but does not depend on Mrc1, which acts as the signaling adapter for the replication checkpoint...
  24. Wang G, Tong X, Weng S, Zhou H. Multiple phosphorylation of Rad9 by CDK is required for DNA damage checkpoint activation. Cell Cycle. 2012;11:3792-800 pubmed publisher
    ..We propose that multiple phosphorylation of Rad9 by CDK may provide a more robust system to allow Rad9 to control cell cycle-dependent DNA damage checkpoint activation. ..
  25. Mohanty B, Bairwa N, Bastia D. Contrasting roles of checkpoint proteins as recombination modulators at Fob1-Ter complexes with or without fork arrest. Eukaryot Cell. 2009;8:487-95 pubmed publisher
    ..Other checkpoint proteins of the checkpoint adapter/mediator class such as Mrc1 and Rad9, which channel signals from the sensor to the effector kinase, tended to suppress recombination at Fob1-..
  26. Nedelcheva M, Roguev A, Dolapchiev L, Shevchenko A, Taskov H, Shevchenko A, et al. Uncoupling of unwinding from DNA synthesis implies regulation of MCM helicase by Tof1/Mrc1/Csm3 checkpoint complex. J Mol Biol. 2005;347:509-21 pubmed
    ..In concordance with this suggestion, we found that the Tof1/Csm3/Mrc1 checkpoint complex interacts directly with the MCM helicase during both replication fork progression and when the ..
  27. Szakal B, Branzei D. Premature Cdk1/Cdc5/Mus81 pathway activation induces aberrant replication and deleterious crossover. EMBO J. 2013;32:1155-67 pubmed publisher
  28. Simoneau A, Ricard Ã, Weber S, Hammond Martel I, Wong L, Sellam A, et al. Chromosome-wide histone deacetylation by sirtuins prevents hyperactivation of DNA damage-induced signaling upon replicative stress. Nucleic Acids Res. 2016;44:2706-26 pubmed publisher
    ..Overall, our data support the concept that chromosome-wide histone deacetylation by sirtuins is critical to mitigate growth defects caused by endogenous genotoxins. ..
  29. Ge Z, Nair D, Guan X, Rastogi N, Freitas M, Parthun M. Sites of acetylation on newly synthesized histone H4 are required for chromatin assembly and DNA damage response signaling. Mol Cell Biol. 2013;33:3286-98 pubmed publisher
    ..These results indicate that the sites of acetylation on newly synthesized histones H3 and H4 can function in nonoverlapping ways that are required for chromatin assembly, viability, and DNA damage response signaling. ..
  30. Lin C, Wu M, Gay S, Marjavaara L, Lai M, Hsiao W, et al. H2B mono-ubiquitylation facilitates fork stalling and recovery during replication stress by coordinating Rad53 activation and chromatin assembly. PLoS Genet. 2014;10:e1004667 pubmed publisher
  31. Faucher D, Wellinger R. Methylated H3K4, a transcription-associated histone modification, is involved in the DNA damage response pathway. PLoS Genet. 2010;6: pubmed publisher
    ..Given the high degree of conservation for the methyltransferase and the histone mark in a broad variety of organisms, these results could have similar implications for genome stability mechanisms in vertebrate and mammalian cells. ..
  32. Gómez González B, Felipe Abrio I, Aguilera A. The S-phase checkpoint is required to respond to R-loops accumulated in THO mutants. Mol Cell Biol. 2009;29:5203-13 pubmed publisher
    ..In light of these results, we propose a model in which R-loop-mediated recombination is explained by template switching. ..
  33. Maculins T, Nkosi P, Nishikawa H, Labib K. Tethering of SCF(Dia2) to the Replisome Promotes Efficient Ubiquitylation and Disassembly of the CMG Helicase. Curr Biol. 2015;25:2254-9 pubmed publisher
    ..leucine-rich repeats, but Dia2 also has a TPR domain at its amino terminus that interacts with the Ctf4 and Mrc1 subunits of the replisome progression complex, which assembles around the CMG helicase at replication forks...
  34. Niño C, Guet D, Gay A, Brutus S, Jourquin F, Mendiratta S, et al. Posttranslational marks control architectural and functional plasticity of the nuclear pore complex basket. J Cell Biol. 2016;212:167-80 pubmed publisher
    ..Together, these data reveal an original mechanism contributing to the plasticity of the NPC at a molecular-organization and functional level. ..
  35. Tsolou A, Lydall D. Mrc1 protects uncapped budding yeast telomeres from exonuclease EXO1. DNA Repair (Amst). 2007;6:1607-17 pubmed
    b>Mrc1 (Mediator of Replication Checkpoint 1) is a component of the DNA replication fork machinery and is necessary for checkpoint activation after replication stress. In this study, we addressed the role of Mrc1 at uncapped telomeres...
  36. Lin Z, Dong M, Zhang Y, Lee E, Lin H. Cbr1 is a Dph3 reductase required for the tRNA wobble uridine modification. Nat Chem Biol. 2016;12:995-997 pubmed publisher
    ..The NADH- and Cbr1-dependent reduction of Dph3 may provide a regulatory linkage between cellular metabolic state and protein translation. ..
  37. Borges V, Smith D, Whitehouse I, Uhlmann F. An Eco1-independent sister chromatid cohesion establishment pathway in S. cerevisiae. Chromosoma. 2013;122:121-34 pubmed publisher Eco1, several other factors contribute to cohesion establishment, including Ctf4, Ctf18, Tof1, Csm3, Chl1 and Mrc1, but little is known about their roles. Here, we show that each of these factors facilitates cohesin acetylation...
  38. Koren A, Soifer I, Barkai N. MRC1-dependent scaling of the budding yeast DNA replication timing program. Genome Res. 2010;20:781-90 pubmed publisher
    ..In sharp contrast, cells deleted of MRC1, a gene implicated in replication fork stabilization and in the replication checkpoint pathway, maintained wild-..
  39. Gellon L, Razidlo D, Gleeson O, Verra L, Schulz D, Lahue R, et al. New functions of Ctf18-RFC in preserving genome stability outside its role in sister chromatid cohesion. PLoS Genet. 2011;7:e1001298 pubmed publisher
    ..First, genetic instability in mutants of Ctf18-RFC was exacerbated by simultaneous deletion of the fork stabilizer Mrc1, but suppressed by deletion of the repair protein Rad52...
  40. García Rodríguez L, De Piccoli G, Marchesi V, Jones R, Edmondson R, Labib K. A conserved Polϵ binding module in Ctf18-RFC is required for S-phase checkpoint activation downstream of Mec1. Nucleic Acids Res. 2015;43:8830-8 pubmed publisher
    ..These findings indicate that the association of Ctf18-RFC with Pol ϵ at defective replication forks is a key step in activation of the S-phase checkpoint. ..
  41. Liang J, Suhandynata R, Zhou H. Phosphorylation of Sae2 Mediates Forkhead-associated (FHA) Domain-specific Interaction and Regulates Its DNA Repair Function. J Biol Chem. 2015;290:10751-63 pubmed publisher
    ..These findings suggest that threonine-specific phosphorylation of Sae2 by Mec1 and Tel1 contributes to DNA repair and genome maintenance via its interactions with Rad53 and Dun1. ..
  42. Caldwell J, Chen Y, Schollaert K, Theis J, Babcock G, Newlon C, et al. Orchestration of the S-phase and DNA damage checkpoint pathways by replication forks from early origins. J Cell Biol. 2008;180:1073-86 pubmed publisher
    ..Thus, oncogene-mediated deregulation of cyclins in the early stages of cancer development could contribute to genomic instability through a deficiency in the forks required to establish the S-phase checkpoint. ..
  43. Green B, Li J. Loss of rereplication control in Saccharomyces cerevisiae results in extensive DNA damage. Mol Biol Cell. 2005;16:421-32 pubmed
    ..In contrast, Mrc1, a checkpoint protein required for recognition of replication stress, does not play a role in the response to ..
  44. Tripathi K, Matmati N, Zheng W, Hannun Y, Mohanty B. Cellular morphogenesis under stress is influenced by the sphingolipid pathway gene ISC1 and DNA integrity checkpoint genes in Saccharomyces cerevisiae. Genetics. 2011;189:533-47 pubmed publisher
    ..That is, yeast carrying deletions of both ISC1 and a replication checkpoint mediator gene including MRC1, TOF1, or CSM3 display basal morphological defects, which increase following HU treatment...
  45. Chang M, Bellaoui M, Zhang C, Desai R, Morozov P, Delgado Cruzata L, et al. RMI1/NCE4, a suppressor of genome instability, encodes a member of the RecQ helicase/Topo III complex. EMBO J. 2005;24:2024-33 pubmed
    ..In addition, rmi1Delta strains fail to fully activate Rad53 upon exposure to DNA-damaging agents, suggesting that Rmi1 is also an important part of the Rad53-dependent DNA damage response. ..
  46. Das Bradoo S, Nguyen H, Wood J, Ricke R, Haworth J, Bielinsky A. Defects in DNA ligase I trigger PCNA ubiquitylation at Lys 107. Nat Cell Biol. 2010;12:74-9; sup pp 1-20 pubmed publisher
    ..We propose that PCNA ubiquitylation provides a 'DNA damage code' that allows cells to categorize different types of defects that arise during DNA replication. ..
  47. Alzu A, Bermejo R, Begnis M, Lucca C, Piccini D, Carotenuto W, et al. Senataxin associates with replication forks to protect fork integrity across RNA-polymerase-II-transcribed genes. Cell. 2012;151:835-846 pubmed publisher
  48. Jay K, Smith D, Blackburn E. Early Loss of Telomerase Action in Yeast Creates a Dependence on the DNA Damage Response Adaptor Proteins. Mol Cell Biol. 2016;36:1908-19 pubmed publisher
    ..We propose that normally occurring telomeric DNA replication stress is resolved by telomerase activity and the DDR in two parallel pathways and that deletion of Sml1 prevents this stress. ..
  49. Putnam C, Pallis K, Hayes T, Kolodner R. DNA repair pathway selection caused by defects in TEL1, SAE2, and de novo telomere addition generates specific chromosomal rearrangement signatures. PLoS Genet. 2014;10:e1004277 pubmed publisher
    ..on chrV L while retaining a chrV L telomeric hph marker was significantly higher in tel1?, sae2?, rad53? sml1?, and mrc1? tof1? mutants...
  50. Redon C, Pilch D, Bonner W. Genetic analysis of Saccharomyces cerevisiae H2A serine 129 mutant suggests a functional relationship between H2A and the sister-chromatid cohesion partners Csm3-Tof1 for the repair of topoisomerase I-induced DNA damage. Genetics. 2006;172:67-76 pubmed
    ..We now report that efficient repair also requires proteins involved in chromatid cohesion: Csm3; Tof1; Mrc1, and Dcc1. Epistasis analysis defined several pathways involving these proteins...
  51. Esta A, Ma E, Dupaigne P, Maloisel L, Guerois R, Le Cam E, et al. Rad52 sumoylation prevents the toxicity of unproductive Rad51 filaments independently of the anti-recombinase Srs2. PLoS Genet. 2013;9:e1003833 pubmed publisher
    ..This conclusion is strengthened by the finding that Rad52 is often associated with complete Rad51 filaments in vitro...
  52. Li X, Tye B. Ploidy dictates repair pathway choice under DNA replication stress. Genetics. 2011;187:1031-40 pubmed publisher
    ..In response to replication stress, early events associated with ploidy dictate the repair pathway choice. This study uncovers a fundamental difference between haplophase and diplophase in the maintenance of genome integrity. ..
  53. Karumbati A, Wilson T. Abrogation of the Chk1-Pds1 checkpoint leads to tolerance of persistent single-strand breaks in Saccharomyces cerevisiae. Genetics. 2005;169:1833-44 pubmed
    ..We propose a model in which recombinational repair during S phase coupled with failure of the metaphase-anaphase checkpoint allows for tolerance of persistent single-strand breaks at the expense of genome stability. ..
  54. Cussiol J, Jablonowski C, Yimit A, Brown G, Smolka M. Dampening DNA damage checkpoint signalling via coordinated BRCT domain interactions. EMBO J. 2015;34:1704-17 pubmed publisher
    ..MBD also interacts with the Mus81 nuclease following checkpoint dampening, suggesting a spatio-temporal coordination of checkpoint signalling and DNA repair via a combinatorial mode of BRCT-domains interactions. ..
  55. Traven A, Hammet A, Tenis N, Denis C, Heierhorst J. Ccr4-not complex mRNA deadenylase activity contributes to DNA damage responses in Saccharomyces cerevisiae. Genetics. 2005;169:65-75 pubmed
    ..the major cytoplasmic mRNA deadenylase complex, have complex genetic interactions with the checkpoint genes DUN1, MRC1, RAD9, and RAD17 in response to DNA-damaging agents hydroxyurea (HU) and methylmethane sulfonate (MMS)...
  56. Vijeh Motlagh N, Seki M, Branzei D, Enomoto T. Mgs1 and Rad18/Rad5/Mms2 are required for survival of Saccharomyces cerevisiae mutants with novel temperature/cold sensitive alleles of the DNA polymerase delta subunit, Pol31. DNA Repair (Amst). 2006;5:1459-74 pubmed
    ..Deletion of SGS1, RAD52, SRS2, MRC1 or RAD24 had a deleterious effect only in combination with those pol31 alleles that had a phenotype as single ..
  57. Voineagu I, Narayanan V, Lobachev K, Mirkin S. Replication stalling at unstable inverted repeats: interplay between DNA hairpins and fork stabilizing proteins. Proc Natl Acad Sci U S A. 2008;105:9936-41 pubmed publisher
    ..Finally, we report that yeast fork-stabilizing proteins, Tof1 and Mrc1, are required to counteract repeat-mediated replication stalling...
  58. Saka K, Takahashi A, Sasaki M, Kobayashi T. More than 10% of yeast genes are related to genome stability and influence cellular senescence via rDNA maintenance. Nucleic Acids Res. 2016;44:4211-21 pubmed publisher
  59. Che J, Smith S, Kim Y, Shim E, Myung K, Lee S. Hyper-Acetylation of Histone H3K56 Limits Break-Induced Replication by Inhibiting Extensive Repair Synthesis. PLoS Genet. 2015;11:e1004990 pubmed publisher
    ..RTT109 or ASF1, or in combination with the H3K56R mutation, which also restores tolerance to replication stress in mrc1 mutants...
  60. Budd M, Antoshechkin I, Reis C, Wold B, Campbell J. Inviability of a DNA2 deletion mutant is due to the DNA damage checkpoint. Cell Cycle. 2011;10:1690-8 pubmed
    ..of a novel, spontaneously arising suppressor of dna2? now reveals that mutation of rad9 and double mutation of rad9 mrc1 can also suppress the lethality of dna2? mutants...
  61. Ohya T, Arai H, Kubota Y, Shinagawa H, Hishida T. A SUMO-like domain protein, Esc2, is required for genome integrity and sister chromatid cohesion in Saccharomyces cerevisiae. Genetics. 2008;180:41-50 pubmed publisher
  62. Tsai F, Vijayraghavan S, Prinz J, MacAlpine H, MacAlpine D, Schwacha A. Mcm2-7 Is an Active Player in the DNA Replication Checkpoint Signaling Cascade via Proposed Modulation of Its DNA Gate. Mol Cell Biol. 2015;35:2131-43 pubmed publisher
    ..This allele exhibits normal downstream mediator (Mrc1) phosphorylation, implying DRC sensor kinase activation...
  63. Böhm S, Szakal B, Herken B, Sullivan M, Mihalevic M, Kabbinavar F, et al. The Budding Yeast Ubiquitin Protease Ubp7 Is a Novel Component Involved in S Phase Progression. J Biol Chem. 2016;291:4442-52 pubmed publisher
    ..In summary, our results suggest that Ubp7 contributes to S phase progression by affecting the chromatin state at replication forks, and we propose histone H2B ubiquitination as a potential substrate of Ubp7. ..
  64. Leclere A, Yang J, Kirkpatrick D. The role of CSM3, MRC1, and TOF1 in minisatellite stability and large loop DNA repair during meiosis in yeast. Fungal Genet Biol. 2013;50:33-43 pubmed publisher
    ..length expansions are controlled by the products of the CSM3 and TOF1 genes, while contractions are controlled by MRC1. By examining meiotic segregation patterns in yeast strains heterozygous for the 26bp his4-lopd insert, we found ..
  65. Torres J, Schnakenberg S, Zakian V. Saccharomyces cerevisiae Rrm3p DNA helicase promotes genome integrity by preventing replication fork stalling: viability of rrm3 cells requires the intra-S-phase checkpoint and fork restart activities. Mol Cell Biol. 2004;24:3198-212 pubmed
    ..The rrm3 system provides a unique opportunity to learn the fate of forks whose progress is impaired by natural impediments rather than by exogenous DNA damage. ..
  66. Koepp D, Kile A, Swaminathan S, Rodriguez Rivera V. The F-box protein Dia2 regulates DNA replication. Mol Biol Cell. 2006;17:1540-8 pubmed
    ..Interestingly, prolonging G1-phase in dia2 cells prevents the accumulation of DNA damage in S-phase. We propose that Dia2 is an origin-binding protein that plays a role in regulating DNA replication. ..
  67. Duch A, Canal B, Barroso S, García Rubio M, Seisenbacher G, Aguilera A, et al. Multiple signaling kinases target Mrc1 to prevent genomic instability triggered by transcription-replication conflicts. Nat Commun. 2018;9:379 pubmed publisher
    ..mechanism for the protection of genomic integrity upon transcriptional outbursts in S phase that is mediated by Mrc1. The N-terminal phosphorylation of Mrc1 blocked replication and prevented transcription-associated recombination (..
  68. Naylor M, Li J, Osborn A, Elledge S. Mrc1 phosphorylation in response to DNA replication stress is required for Mec1 accumulation at the stalled fork. Proc Natl Acad Sci U S A. 2009;106:12765-70 pubmed publisher
    ..The budding yeast Mec1 sensor kinase, Mrc1 mediator, and Rad53 effector kinase are central to this signal transduction cascade in S phase...
  69. Baldo V, Testoni V, Lucchini G, Longhese M. Dominant TEL1-hy mutations compensate for Mec1 lack of functions in the DNA damage response. Mol Cell Biol. 2008;28:358-75 pubmed
    ..Finally, Tel1-hy544 can activate the checkpoint more efficiently than wild-type Tel1, while it causes telomere shortening, indicating that the checkpoint and telomeric functions of Tel1 can be separable. ..
  70. Nitani N, Nakamura K, Nakagawa C, Masukata H, Nakagawa T. Regulation of DNA replication machinery by Mrc1 in fission yeast. Genetics. 2006;174:155-65 pubmed
    ..b>Mrc1 is required to activate Cds1 and prevent the replication machinery from uncoupling from DNA synthesis...
  71. Berens T, Toczyski D. Colocalization of Mec1 and Mrc1 is sufficient for Rad53 phosphorylation in vivo. Mol Biol Cell. 2012;23:1058-67 pubmed publisher
    ..In the endogenous replication checkpoint, Mec1 phosphorylation of Rad53 requires Mrc1, a replisome component...
  72. Litwin I, Bakowski T, Maciaszczyk Dziubinska E, Wysocki R. The LSH/HELLS homolog Irc5 contributes to cohesin association with chromatin in yeast. Nucleic Acids Res. 2017;45:6404-6416 pubmed publisher
    ..Our results suggest that Irc5 is an auxiliary factor that is involved in cohesin association with chromatin. ..