Gene Symbol: SCC2
Description: cohesin-loading factor complex subunit SCC2
Alias: cohesin-loading factor complex subunit SCC2
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Mayer M, Gygi S, Aebersold R, Hieter P. Identification of RFC(Ctf18p, Ctf8p, Dcc1p): an alternative RFC complex required for sister chromatid cohesion in S. cerevisiae. Mol Cell. 2001;7:959-70 pubmed
  2. D Ambrosio C, Schmidt C, Katou Y, Kelly G, Itoh T, Shirahige K, et al. Identification of cis-acting sites for condensin loading onto budding yeast chromosomes. Genes Dev. 2008;22:2215-27 pubmed publisher
    ..Condensin-binding sites coincide with those of the loading factor Scc2/4 of the related cohesin complex...
  3. Lopez Serra L, Kelly G, Patel H, Stewart A, Uhlmann F. The Scc2-Scc4 complex acts in sister chromatid cohesion and transcriptional regulation by maintaining nucleosome-free regions. Nat Genet. 2014;46:1147-51 pubmed publisher
    ..Cohesin loading onto chromosomes depends on the Scc2-Scc4 cohesin loader complex, but the chromatin features that form cohesin loading sites remain poorly understood...
  4. Gard S, Light W, Xiong B, Bose T, McNairn A, Harris B, et al. Cohesinopathy mutations disrupt the subnuclear organization of chromatin. J Cell Biol. 2009;187:455-62 pubmed publisher
    ..We propose that the cohesin network affects gene regulation by facilitating the subnuclear organization of chromatin. ..
  5. Kogut I, Wang J, Guacci V, Mistry R, Megee P. The Scc2/Scc4 cohesin loader determines the distribution of cohesin on budding yeast chromosomes. Genes Dev. 2009;23:2345-57 pubmed publisher
    ..Chromosomal cohesins are distributed nonrandomly, and their deposition requires the heterodimeric Scc2/Scc4 loader...
  6. Ciosk R, Shirayama M, Tanaka T, Toth A, Shevchenko A, Nasmyth K. Cohesin's binding to chromosomes depends on a separate complex consisting of Scc2 and Scc4 proteins. Mol Cell. 2000;5:243-54 pubmed
    ..In scc2 or scc4 mutants, cohesin complexes form normally but fail to bind both to centromeres and to chromosome arms...
  7. Toth A, Ciosk R, Uhlmann F, Galova M, Schleiffer A, Nasmyth K. Yeast cohesin complex requires a conserved protein, Eco1p(Ctf7), to establish cohesion between sister chromatids during DNA replication. Genes Dev. 1999;13:320-33 pubmed
    ..The sixth protein, Eco1p(Ctf7p), is not a Cohesin subunit. It is necessary for the establishment of cohesion during DNA replication but not for its maintenance during G2 and M phases. ..
  8. Lindroos H, Ström L, Itoh T, Katou Y, Shirahige K, Sjögren C. Chromosomal association of the Smc5/6 complex reveals that it functions in differently regulated pathways. Mol Cell. 2006;22:755-67 pubmed
    ..Altogether this shows that the complex maintains genomic integrity in multiple ways, and evidence is presented that the Smc5/6 complex is needed during replication to prevent the accumulation of branched chromosome structures. ..
  9. Unal E, Arbel Eden A, Sattler U, Shroff R, Lichten M, Haber J, et al. DNA damage response pathway uses histone modification to assemble a double-strand break-specific cohesin domain. Mol Cell. 2004;16:991-1002 pubmed
    ..We also provide evidence that the DSB-induced cohesin domain functions in postreplicative repair. ..

More Information


  1. Fernius J, Nerusheva O, Galander S, Alves F, Rappsilber J, Marston A. Cohesin-dependent association of scc2/4 with the centromere initiates pericentromeric cohesion establishment. Curr Biol. 2013;23:599-606 pubmed publisher
    ..Cohesin loading onto chromosomes universally requires the Scc2/4 "loader" complex (also called NippedBL/Mau2), mutations in which cause the developmental disorder Cornelia de ..
  2. Ström L, Lindroos H, Shirahige K, Sjögren C. Postreplicative recruitment of cohesin to double-strand breaks is required for DNA repair. Mol Cell. 2004;16:1003-15 pubmed
    ..We find that in the absence of functional cohesin-loading proteins (Scc2/Scc4), the accumulation of cohesin at DSBs is abolished and repair is defective, even though sister chromatids are ..
  3. McLellan J, O Neil N, Tarailo S, Stoepel J, Bryan J, Rose A, et al. Synthetic lethal genetic interactions that decrease somatic cell proliferation in Caenorhabditis elegans identify the alternative RFC CTF18 as a candidate cancer drug target. Mol Biol Cell. 2009;20:5306-13 pubmed publisher
    ..elegans. Furthermore, the C. elegans assay system will contribute to our knowledge of genetic interactions in a multicellular animal and is a powerful approach to identify new cancer therapeutic targets. ..
  4. Hinshaw S, Makrantoni V, Kerr A, Marston A, Harrison S. Structural evidence for Scc4-dependent localization of cohesin loading. elife. 2015;4:e06057 pubmed publisher
    ..Initiation of sister chromatid cohesion depends on a separate complex, Scc2(NIPBL)/Scc4(Mau2) (Scc2/4), which loads cohesin onto DNA and determines its localization across the genome...
  5. Sjogren C, Nasmyth K. Sister chromatid cohesion is required for postreplicative double-strand break repair in Saccharomyces cerevisiae. Curr Biol. 2001;11:991-5 pubmed
    ..Proteins needed to load cohesin onto chromosomes (Scc2) and to generate cohesion during S phase (Eco1) are also shown to be required for repair...
  6. 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
    ..Finally, we demonstrate that in the absence of Irc5 both the level of chromatin-bound Scc2, a member of cohesin loading complex, and physical interaction between Scc1 and Scc2 are reduced...
  7. Kirkland J, Kamakaka R. Long-range heterochromatin association is mediated by silencing and double-strand DNA break repair proteins. J Cell Biol. 2013;201:809-26 pubmed publisher
    ..SMC proteins and Scc2 localize to the silenced domain, and Scc2 binding requires the presence of ?-H2A...
  8. Orgil O, Mor H, Matityahu A, Onn I. Identification of a region in the coiled-coil domain of Smc3 that is essential for cohesin activity. Nucleic Acids Res. 2016;44:6309-17 pubmed publisher
    ..However, the mutation does not affect inter-protein interactions in either the core complex or with the Scc2 loader...
  9. Lindgren E, Hägg S, Giordano F, Björkegren J, Ström L. Inactivation of the budding yeast cohesin loader Scc2 alters gene expression both globally and in response to a single DNA double strand break. Cell Cycle. 2014;13:3645-58 pubmed publisher
    ..Cohesin and its DNA loader, the Scc2/4 complex have been implicated in all these cellular actions...
  10. Cortes Ledesma F, Aguilera A. Double-strand breaks arising by replication through a nick are repaired by cohesin-dependent sister-chromatid exchange. EMBO Rep. 2006;7:919-26 pubmed
    ..evidence that cohesins are required for repair of replication-born DSBs by SCE, as determined in smc3, scc1 and scc2 mutants, but not for other recombinational repair events...
  11. Wendt K. Resolving the Genomic Localization of the Kollerin Cohesin-Loader Complex. Methods Mol Biol. 2017;1515:115-123 pubmed
    The kollerin complex, consisting of Scc2/Scc4 in yeast and Nipbl/Mau2 in vertebrates, is crucial for the chromatin-association of the cohesin complex and therefore for the critical functions of cohesin in cell division, transcriptional ..
  12. Shwartz M, Matityahu A, Onn I. Identification of Functional Domains in the Cohesin Loader Subunit Scc4 by a Random Insertion/Dominant Negative Screen. G3 (Bethesda). 2016;6:2655-63 pubmed publisher
    ..Initial association of cohesin with chromosomes requires the loader-a heterodimer composed of Scc4 and Scc2 However, very little is known about the loader's mechanism of action...
  13. Zakari M, Trimble Ross R, Peak A, Blanchette M, Seidel C, Gerton J. The SMC Loader Scc2 Promotes ncRNA Biogenesis and Translational Fidelity. PLoS Genet. 2015;11:e1005308 pubmed publisher
    The Scc2-Scc4 complex is essential for loading the cohesin complex onto DNA. Cohesin has important roles in chromosome segregation, DSB repair, and chromosome condensation...
  14. Woodman J, Hoffman M, Dzieciatkowska M, Hansen K, Megee P. Phosphorylation of the Scc2 cohesin deposition complex subunit regulates chromosome condensation through cohesin integrity. Mol Biol Cell. 2015;26:3754-67 pubmed publisher mediated by cohesin, which is deposited on chromosomes by a separate conserved loading complex composed of Scc2 and Scc4 in Saccharomyces cerevisiae...
  15. Hinshaw S, Makrantoni V, Harrison S, Marston A. The Kinetochore Receptor for the Cohesin Loading Complex. Cell. 2017;171:72-84.e13 pubmed publisher
    ..that phosphorylation of the Ctf19 kinetochore protein by a conserved kinase, DDK, provides a binding site for the Scc2/4 cohesin loading complex, thereby directing cohesin loading to centromeres...
  16. Hamza A, Baetz K. Iron-responsive transcription factor Aft1 interacts with kinetochore protein Iml3 and promotes pericentromeric cohesin. J Biol Chem. 2012;287:4139-47 pubmed publisher
    ..Our work defines a new role for Aft1 in chromosome stability and transmission. ..
  17. Parnas O, Zipin Roitman A, Mazor Y, Liefshitz B, Ben Aroya S, Kupiec M. The ELG1 clamp loader plays a role in sister chromatid cohesion. PLoS ONE. 2009;4:e5497 pubmed publisher
    ..A genetic screen identified the cohesin subunit Mcd1/Scc1 and its loader Scc2 as suppressors of the synthetic lethality between elg1 and ctf4...
  18. Lin W, Jin H, Liu X, Hampton K, Yu H. Scc2 regulates gene expression by recruiting cohesin to the chromosome as a transcriptional activator during yeast meiosis. Mol Biol Cell. 2011;22:1985-96 pubmed publisher
    To tether sister chromatids, a protein-loading complex, including Scc2, recruits cohesin to the chromosome at discrete loci...
  19. Arumugam P, Gruber S, Tanaka K, Haering C, Mechtler K, Nasmyth K. ATP hydrolysis is required for cohesin's association with chromosomes. Curr Biol. 2003;13:1941-53 pubmed
    ..Inactivation of the Scc2/4 complex had a similar if not identical effect; namely, the production of tripartite cohesin rings that cannot ..
  20. Woodman J, Fara T, Dzieciatkowska M, Trejo M, Luong N, Hansen K, et al. Cell cycle-specific cleavage of Scc2 regulates its cohesin deposition activity. Proc Natl Acad Sci U S A. 2014;111:7060-5 pubmed publisher
    ..Cohesins are deposited by a conserved heterodimeric loading complex composed of the Scc2 and Scc4 proteins in Saccharomyces cerevisiae, but how the Scc2/Scc4 deposition complex regulates the ..
  21. Ghosh S, Hajra S, Jayaram M. Faithful segregation of the multicopy yeast plasmid through cohesin-mediated recognition of sisters. Proc Natl Acad Sci U S A. 2007;104:13034-9 pubmed
    ..Through binary fluorescence tagging, we reveal that segregation of replicated plasmids occurs in a sister-to-sister fashion. Thus, cohesin serves the same fundamental purpose in plasmid and chromosome segregation. ..
  22. Maradeo M, Garg A, Skibbens R. Rfc5p regulates alternate RFC complex functions in sister chromatid pairing reactions in budding yeast. Cell Cycle. 2010;9:4370-8 pubmed
    ..rescue is specific to establishment pathways in that rfc5-1 greatly accentuates growth defects when expressed in scc2 (deposition), mcd1/scc1 or smc3 (cohesion maintenance) mutated cells...