Gene Symbol: MCD1
Description: kleisin alpha
Alias: PDS3, RHC21, SCC1, kleisin alpha
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Eng T, Guacci V, Koshland D. ROCC, a conserved region in cohesin's Mcd1 subunit, is essential for the proper regulation of the maintenance of cohesion and establishment of condensation. Mol Biol Cell. 2014;25:2351-64 pubmed publisher
    ..We provide evidence that ROCC controls cohesion maintenance and condensation establishment through differential functional interactions with Pds5p and Wpl1p. ..
  2. Arumugam P, Nishino T, Haering C, Gruber S, Nasmyth K. Cohesin's ATPase activity is stimulated by the C-terminal Winged-Helix domain of its kleisin subunit. Curr Biol. 2006;16:1998-2008 pubmed
    ..Cohesin consists of four core subunits, namely Smc1, Smc3, Scc1, and Scc3...
  3. 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
    ..Our results confirm what has long been suspected but never proven, that cohesion between sister chromatids is essential for efficient double-strand break repair in mitotic cells. ..
  4. Ivanov D, Schleiffer A, Eisenhaber F, Mechtler K, Haering C, Nasmyth K. Eco1 is a novel acetyltransferase that can acetylate proteins involved in cohesion. Curr Biol. 2002;12:323-8 pubmed
    ..In Saccharomyces cerevisiae, a complex consisting of Scc1, Smc1, Smc3, and Scc3 proteins, called "cohesin," mediates the connection between sister chromatids...
  5. Heidinger Pauli J, Unal E, Guacci V, Koshland D. The kleisin subunit of cohesin dictates damage-induced cohesion. Mol Cell. 2008;31:47-56 pubmed publisher
    ..Here, using budding yeast, we show that DSB-induced cohesion occurs when cohesin contains the kleisin subunit, Mcd1 (Scc1), but not when Mcd1 is replaced by its meiotic isoform, Rec8...
  6. Maradeo M, Skibbens R. The Elg1-RFC clamp-loading complex performs a role in sister chromatid cohesion. PLoS ONE. 2009;4:e4707 pubmed publisher
    ..This model is based on findings that CTF18 deletion is lethal in combination with mutations in either CTF7(ECO1) or MCD1 sister chromatid cohesion genes and that ctf18 mutant cells exhibit cohesion defects...
  7. Weitzer S, Lehane C, Uhlmann F. A model for ATP hydrolysis-dependent binding of cohesin to DNA. Curr Biol. 2003;13:1930-40 pubmed
    ..DNA is released in anaphase when separase cleaves cohesin's Scc1 subunit...
  8. 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
    ..Cohesin contains four core subunits: Smc1, Smc3, Scc1, and Scc3...
  9. D Amours D, Stegmeier F, Amon A. Cdc14 and condensin control the dissolution of cohesin-independent chromosome linkages at repeated DNA. Cell. 2004;117:455-69 pubmed
    ..This dual role of the FEAR network in initiating mitotic exit and promoting chromosome segregation ensures that exit from mitosis is coupled to the completion of chromosome segregation. ..

More Information


  1. Haering C, L we J, Hochwagen A, Nasmyth K. Molecular architecture of SMC proteins and the yeast cohesin complex. Mol Cell. 2002;9:773-88 pubmed
    ..are held together by the multisubunit cohesin complex, which contains two SMC (Smc1 and Smc3) and two non-SMC (Scc1 and Scc3) proteins...
  2. Mc Intyre J, Muller E, Weitzer S, Snydsman B, Davis T, Uhlmann F. In vivo analysis of cohesin architecture using FRET in the budding yeast Saccharomyces cerevisiae. EMBO J. 2007;26:3783-93 pubmed
    ..The circumference consists of the Smc1 and Smc3 subunits, while Scc1 is thought to close the ring by bridging the Smc (structural maintenance of chromosomes) ATPase head domains...
  3. Alexandru G, Uhlmann F, Mechtler K, Poupart M, Nasmyth K. Phosphorylation of the cohesin subunit Scc1 by Polo/Cdc5 kinase regulates sister chromatid separation in yeast. Cell. 2001;105:459-72 pubmed
    ..a caspase-related protease (separase) destroys the link between sister chromatids by cleaving the cohesin subunit Scc1. During most of the cell cycle, separase is kept inactive by binding to an inhibitory protein called securin...
  4. Heidinger Pauli J, Onn I, Koshland D. Genetic evidence that the acetylation of the Smc3p subunit of cohesin modulates its ATP-bound state to promote cohesion establishment in Saccharomyces cerevisiae. Genetics. 2010;185:1249-56 pubmed publisher
    ..Finally, we provide evidence that Smc3p acetylation, which has an essential role in cohesion establishment, modulates the Smc3p ATP-bound state. ..
  5. Guacci V, Koshland D, Strunnikov A. A direct link between sister chromatid cohesion and chromosome condensation revealed through the analysis of MCD1 in S. cerevisiae. Cell. 1997;91:47-57 pubmed
    The S. cerevisiae MCD1 (mitotic chromosome determinant) gene was identified in genetic screens for genes important for chromosome structure. MCD1 is essential for viability and homologs are found from yeast to humans...
  6. 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. ..
  7. Sutani T, Kawaguchi T, Kanno R, Itoh T, Shirahige K. Budding yeast Wpl1(Rad61)-Pds5 complex counteracts sister chromatid cohesion-establishing reaction. Curr Biol. 2009;19:492-7 pubmed publisher
    ..Taken together, the presented data suggest that Wpl1-Pds5 complex is inhibitory for cohesion establishment and that Eco1 establishes cohesion by hindering the function of Wpl1-Pds5 temporally in S phase...
  8. Mishra A, Hu B, Kurze A, Beckouët F, Farcas A, Dixon S, et al. Both interaction surfaces within cohesin's hinge domain are essential for its stable chromosomal association. Curr Biol. 2010;20:279-89 pubmed publisher
    The cohesin complex that mediates sister chromatid cohesion contains three core subunits: Smc1, Smc3, and Scc1. Heterotypic interactions between Smc1 and Smc3 dimerization domains create stable V-shaped Smc1/Smc3 heterodimers with a hinge ..
  9. Chan K, Gligoris T, Upcher W, Kato Y, Shirahige K, Nasmyth K, et al. Pds5 promotes and protects cohesin acetylation. Proc Natl Acad Sci U S A. 2013;110:13020-5 pubmed publisher
  10. Lavoie B, Hogan E, Koshland D. In vivo dissection of the chromosome condensation machinery: reversibility of condensation distinguishes contributions of condensin and cohesin. J Cell Biol. 2002;156:805-15 pubmed
    ..Finally, we provide the first evidence of a mechanistic link between condensin and cohesin function. A model discussing the functional interplay between cohesin and condensin is presented. ..
  11. Heidinger Pauli J, Unal E, Koshland D. Distinct targets of the Eco1 acetyltransferase modulate cohesion in S phase and in response to DNA damage. Mol Cell. 2009;34:311-21 pubmed publisher
    ..We show that Mcd1p and Smc3p acetylation antagonize Wpl1p only in their proper context. Thus, Eco1p antagonizes Wpl1p in distinct ways to modulate cohesion generation during the cell cycle and after DNA damage. ..
  12. Severin F, Hyman A, Piatti S. Correct spindle elongation at the metaphase/anaphase transition is an APC-dependent event in budding yeast. J Cell Biol. 2001;155:711-8 pubmed
    ..Our data suggest that stabilization of the elongating spindle at the metaphase to anaphase transition involves Pds1-dependent targets other than Esp1. ..
  13. Hornig N, Uhlmann F. Preferential cleavage of chromatin-bound cohesin after targeted phosphorylation by Polo-like kinase. EMBO J. 2004;23:3144-53 pubmed
    ..This is triggered by the protease separase that cleaves the Scc1 subunit of 'cohesin', the protein complex responsible for holding sister chromatids together in metaphase...
  14. 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
  15. Haering C, Schoffnegger D, Nishino T, Helmhart W, Nasmyth K, Lowe J. Structure and stability of cohesin's Smc1-kleisin interaction. Mol Cell. 2004;15:951-64 pubmed
    ..Cohesin's kleisin subunit Scc1 completes the ring, connecting the ABC-like ATPase heads of a V-shaped Smc1/3 heterodimer...
  16. Uhlmann F, Wernic D, Poupart M, Koonin E, Nasmyth K. Cleavage of cohesin by the CD clan protease separin triggers anaphase in yeast. Cell. 2000;103:375-86 pubmed
    ..Cleavage of Sccl in metaphase arrested cells is sufficient to trigger the separation of sister chromatids and their segregation to opposite cell poles. ..
  17. D Ambrosio L, Lavoie B. Pds5 prevents the PolySUMO-dependent separation of sister chromatids. Curr Biol. 2014;24:361-71 pubmed publisher
    ..distinct posttranslational modifications of cohesin, a protein complex consisting of the Smc1-Smc3 ATPase, the Mcd1/Scc1 ?-kleisin, and Scc3...
  18. Lengronne A, McIntyre J, Katou Y, Kanoh Y, Hopfner K, Shirahige K, et al. Establishment of sister chromatid cohesion at the S. cerevisiae replication fork. Mol Cell. 2006;23:787-99 pubmed
    ..This provides evidence for cohesion establishment in the context of replication forks and imposes constraints on the mechanism involved. ..
  19. Farcas A, Uluocak P, Helmhart W, Nasmyth K. Cohesin's concatenation of sister DNAs maintains their intertwining. Mol Cell. 2011;44:97-107 pubmed publisher
    ..We conclude that by retarding Topo II-driven decatenation, cohesin mediates sister chromatid cohesion by an indirect mechanism as well as one involving entrapment of sister DNAs inside its tripartite ring. ..
  20. Lu S, Goering M, Gard S, Xiong B, McNairn A, Jaspersen S, et al. Eco1 is important for DNA damage repair in S. cerevisiae. Cell Cycle. 2010;9:3315-27 pubmed publisher
    ..Our results suggest that Eco1 promotes the reciprocal exchange of chromosome arms and maintenance of heterozygosity during mitosis. ..
  21. Clift D, Bizzari F, Marston A. Shugoshin prevents cohesin cleavage by PP2A(Cdc55)-dependent inhibition of separase. Genes Dev. 2009;23:766-80 pubmed publisher
    ..We propose that Cdc55 is a separase inhibitor that acts downstream from Shugoshin under conditions where sister chromatids are not under tension. ..
  22. Gruber S, Haering C, Nasmyth K. Chromosomal cohesin forms a ring. Cell. 2003;112:765-77 pubmed
    ..Their ATPase heads are thought to be bridged by a third subunit, Scc1, creating a huge triangular ring that could trap sister DNA molecules...
  23. 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
    ..Rather, our findings suggest that budding yeast cohesins are targeted to CARs largely by Scc2/Scc4 loader association at these locations. ..
  24. Rao H, Uhlmann F, Nasmyth K, Varshavsky A. Degradation of a cohesin subunit by the N-end rule pathway is essential for chromosome stability. Nature. 2001;410:955-9 pubmed
    ..the metaphase-anaphase transition in the yeast Saccharomyces cerevisiae, the ESP1-encoded protease separin cleaves SCC1, a subunit of cohesin with a relative molecular mass of 63,000 (Mr 63K)...
  25. Lengronne A, Katou Y, Mori S, Yokobayashi S, Kelly G, Itoh T, et al. Cohesin relocation from sites of chromosomal loading to places of convergent transcription. Nature. 2004;430:573-8 pubmed
    ..Cohesin localization to places of convergent transcription is conserved in fission yeast, suggesting that it is a common feature of eukaryotic chromosomes. ..
  26. Liang F, Richmond D, Wang Y. Coordination of chromatid separation and spindle elongation by antagonistic activities of mitotic and S-phase CDKs. PLoS Genet. 2013;9:e1003319 pubmed publisher
    ..Therefore, the tight temporal control of spindle elongation and cohesin cleavage assure orchestrated chromosome separation and spindle elongation. ..
  27. Mendoza M, Norden C, Durrer K, Rauter H, Uhlmann F, Barral Y. A mechanism for chromosome segregation sensing by the NoCut checkpoint. Nat Cell Biol. 2009;11:477-83 pubmed publisher
    ..These findings provide the first evidence that NoCut is triggered by the interaction of acetylated chromatin with the passenger complex at the spindle midzone. ..
  28. Edwards S, Li C, Levy D, Brown J, Snow P, Campbell J. Saccharomyces cerevisiae DNA polymerase epsilon and polymerase sigma interact physically and functionally, suggesting a role for polymerase epsilon in sister chromatid cohesion. Mol Cell Biol. 2003;23:2733-48 pubmed
    ..required for establishing sister chromatid cohesion; and pol2 mutations act synergistically with smc1 and scc1. We also show that trf5 Delta mutants, like trf4 Delta mutants, are defective in DNA repair and sister chromatid ..
  29. Snead J, Sullivan M, Lowery D, Cohen M, Zhang C, Randle D, et al. A coupled chemical-genetic and bioinformatic approach to Polo-like kinase pathway exploration. Chem Biol. 2007;14:1261-72 pubmed
    ..Finally, an examination of Cdc5 binding by SPB-localized proteins expanded our knowledge of Cdc5 function at the SPB. ..
  30. Yaakov G, THORN K, Morgan D. Separase biosensor reveals that cohesin cleavage timing depends on phosphatase PP2A(Cdc55) regulation. Dev Cell. 2012;23:124-36 pubmed publisher
    ..The protease separase triggers sister separation by cleaving the Scc1/Mcd1 subunit of the cohesin ring that holds sisters together...
  31. Sundin B, Chiu C, Riffle M, Davis T, Muller E. Localization of proteins that are coordinately expressed with Cln2 during the cell cycle. Yeast. 2004;21:793-800 pubmed
    ..A complete list of localizations, along with images, can be found at our website ( ..
  32. Liu C, van Dyk D, Li Y, Andrews B, Rao H. A genome-wide synthetic dosage lethality screen reveals multiple pathways that require the functioning of ubiquitin-binding proteins Rad23 and Dsk2. BMC Biol. 2009;7:75 pubmed publisher
  33. 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...
  34. Elbatsh A, Haarhuis J, Petela N, Chapard C, Fish A, Celie P, et al. Cohesin Releases DNA through Asymmetric ATPase-Driven Ring Opening. Mol Cell. 2016;61:575-588 pubmed publisher
    ..We propose that Smc3 acetylation locks cohesin rings around the sister chromatids by counteracting an activity associated with one of cohesin's two ATPase sites. ..
  35. Kulemzina I, Schumacher M, Verma V, Reiter J, Metzler J, Failla A, et al. Cohesin rings devoid of Scc3 and Pds5 maintain their stable association with the DNA. PLoS Genet. 2012;8:e1002856 pubmed publisher
    ..The ring is composed of three proteins: Smc1, Smc3 and Scc1. The roles of three additional proteins that associate with the ring, Scc3, Pds5 and Wpl1, are not well understood...
  36. Tittel Elmer M, Lengronne A, Davidson M, Bacal J, Francois P, Hohl M, et al. Cohesin association to replication sites depends on rad50 and promotes fork restart. Mol Cell. 2012;48:98-108 pubmed publisher
    ..Together, these data reveal a function for cohesin in the maintenance of genome integrity during S phase. ..
  37. 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
    ..We show that, unexpectedly, Scc2 association with centromeres depends on cohesin itself. The absence of the Scc1/Mcd1/Rad21 cohesin subunit precludes Scc2 association with centromeres from anaphase until late G1...
  38. Xia Z, Webster A, Du F, Piatkov K, Ghislain M, Varshavsky A. Substrate-binding sites of UBR1, the ubiquitin ligase of the N-end rule pathway. J Biol Chem. 2008;283:24011-28 pubmed publisher
  39. Pakchuen S, Ishibashi M, Takakusagi E, Shirahige K, Sutani T. Physical Association of Saccharomyces cerevisiae Polo-like Kinase Cdc5 with Chromosomal Cohesin Facilitates DNA Damage Response. J Biol Chem. 2016;291:17228-46 pubmed publisher
    ..of anaphase, a protease called separase breaks the link between sister chromatids by cleaving the cohesin subunit Scc1. This irreversible step in the cell cycle is promoted by degradation of the separase inhibitor, securin, and polo-..
  40. Cena A, Kozłowska E, Płochocka D, Grynberg M, Ishikawa T, Fronk J, et al. The F658G substitution in Saccharomyces cerevisiae cohesin Irr1/Scc3 is semi-dominant in the diploid and disturbs mitosis, meiosis and the cell cycle. Eur J Cell Biol. 2008;87:831-44 pubmed publisher
    ..Nevertheless, the mutant protein retains its ability to interact with Scc1p, another component of the cohesin complex, as shown by coimmunoprecipitation. ..
  41. Orgil O, Matityahu A, Eng T, Guacci V, Koshland D, Onn I. A conserved domain in the scc3 subunit of cohesin mediates the interaction with both mcd1 and the cohesin loader complex. PLoS Genet. 2015;11:e1005036 pubmed publisher
    ..Cohesin is comprised of Scc3, Mcd1, Smc1, and Smc3...
  42. Mokranjac D, Sichting M, Neupert W, Hell K. Tim14, a novel key component of the import motor of the TIM23 protein translocase of mitochondria. EMBO J. 2003;22:4945-56 pubmed
    ..We propose a model in which Tim14 is required for the activation of mtHsp70 and enables this chaperone to act in a rapid and regulated manner in the Tim44-mediated trapping of unfolded preproteins entering the matrix. ..
  43. Maradeo M, Skibbens R. Epitope tag-induced synthetic lethality between cohesin subunits and Ctf7/Eco1 acetyltransferase. FEBS Lett. 2010;584:4037-40 pubmed publisher
    ..Tagging either Smc1 or Mcd1/Scc1, but not Scc3/Irr1, appears to abolish access to Smc3 in ctf7/eco1 mutant cells, suggesting that Smc1 and Smc3 head ..
  44. Visintin R, Stegmeier F, Amon A. The role of the polo kinase Cdc5 in controlling Cdc14 localization. Mol Biol Cell. 2003;14:4486-98 pubmed
    ..Cdc5 promotes Cdc14 phosphorylation and, by stimulating the MEN, Cfi1/Net1 phosphorylation. Furthermore, we suggest that Cdc14 release from the nucleolus only occurs when Cdc14 and Cfi1/Net1 are both phosphorylated. ..
  45. 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 establishment pathways in that rfc5-1 greatly accentuates growth defects when expressed in scc2 (deposition), mcd1/scc1 or smc3 (cohesion maintenance) mutated cells...
  46. 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. ..
  47. 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
    ..Here, we identify Irc5 as a novel interactor of the cohesin complex. Irc5 associates with Scc1 cohesin subunit and contributes to cohesin binding to chromatin...
  48. Katis V, Lipp J, Imre R, Bogdanova A, Okaz E, Habermann B, et al. Rec8 phosphorylation by casein kinase 1 and Cdc7-Dbf4 kinase regulates cohesin cleavage by separase during meiosis. Dev Cell. 2010;18:397-409 pubmed publisher
    ..Our data suggest that PP2A protects centromeric cohesion by opposing CK1delta/epsilon- and DDK-dependent phosphorylation of Rec8. ..
  49. Samora C, Saksouk J, Goswami P, Wade B, Singleton M, Bates P, et al. Ctf4 Links DNA Replication with Sister Chromatid Cohesion Establishment by Recruiting the Chl1 Helicase to the Replisome. Mol Cell. 2016;63:371-84 pubmed publisher
    ..Our results reveal how Ctf4 forms a replisomal interaction hub that coordinates replication fork progression and sister chromatid cohesion establishment. ..
  50. Beckouët F, Srinivasan M, Roig M, Chan K, Scheinost J, Batty P, et al. Releasing Activity Disengages Cohesin's Smc3/Scc1 Interface in a Process Blocked by Acetylation. Mol Cell. 2016;61:563-574 pubmed publisher
    Sister chromatid cohesion conferred by entrapment of sister DNAs within a tripartite ring formed between cohesin's Scc1, Smc1, and Smc3 subunits is created during S and destroyed at anaphase through Scc1 cleavage by separase...
  51. Bokros M, Gravenmier C, Jin F, Richmond D, Wang Y. Fin1-PP1 Helps Clear Spindle Assembly Checkpoint Protein Bub1 from Kinetochores in Anaphase. Cell Rep. 2016;14:1074-1085 pubmed publisher
    ..This mechanism is required for efficient SAC silencing after SAC is challenged, and untimely Fin1-kinetochore association causes premature SAC silencing and chromosome missegregation. ..
  52. Heo S, Tatebayashi K, Ikeda H. The budding yeast cohesin gene SCC1/MCD1/RHC21 genetically interacts with PKA, CDK and APC. Curr Genet. 1999;36:329-38 pubmed
    ..cohesion, we have isolated multicopy suppressors of a temperature-sensitive (ts) mutation in the SCC1/MCD1/RHC21 gene which encodes a component of the cohesin complex in budding yeast...
  53. Shimogawa M, Wargacki M, Muller E, Davis T. Laterally attached kinetochores recruit the checkpoint protein Bub1, but satisfy the spindle checkpoint. Cell Cycle. 2010;9:3619-28 pubmed
  54. Kobayashi T, Horiuchi T, Tongaonkar P, Vu L, Nomura M. SIR2 regulates recombination between different rDNA repeats, but not recombination within individual rRNA genes in yeast. Cell. 2004;117:441-53 pubmed
    ..From these and other experiments, we conclude that SIR2 prevents unequal sister-chromatid recombination, probably by forming special cohesin structures, without significant effects on recombinational events within individual rRNA genes. ..
  55. Kateneva A, Konovchenko A, Guacci V, Dresser M. Recombination protein Tid1p controls resolution of cohesin-dependent linkages in meiosis in Saccharomyces cerevisiae. J Cell Biol. 2005;171:241-53 pubmed
    ..We propose that Tid1p remodels Mcd1p-mediated cohesion early in meiotic prophase to facilitate interhomologue recombination and the subsequent segregation of homologous chromosomes. ..
  56. Weiss C, Niv A, Azem A. Two-step purification of mitochondrial Hsp70, Ssc1p, using Mge1(His)(6) immobilized on Ni-agarose. Protein Expr Purif. 2002;24:268-73 pubmed
    ..The CD spectrum of Ssc1p had a pattern and temperature dependence similar to those shown for other hsp70 homologs, with a midpoint of the major transition at approximately 70 degrees C. ..
  57. Rodriguez Navarro S, Igual J, Pérez Ortín J. SRC1: an intron-containing yeast gene involved in sister chromatid segregation. Yeast. 2002;19:43-54 pubmed
    ..Deletion of the gene induces a shortening of the anaphase and aggravates the phenotype of scc1 and esp1 conditional mutants, which suggests a direct role of the protein in sister chromatid separation.
  58. Pinsky B, Nelson C, Biggins S. Protein phosphatase 1 regulates exit from the spindle checkpoint in budding yeast. Curr Biol. 2009;19:1182-7 pubmed publisher
    ..We therefore propose that PP1 activity silences the checkpoint by reversing key phosphorylation events. ..
  59. 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
    ..molecular 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...
  60. Yang H, Ren Q, Zhang Z. Cleavage of Mcd1 by caspase-like protease Esp1 promotes apoptosis in budding yeast. Mol Biol Cell. 2008;19:2127-34 pubmed publisher
    ..Here, we report that Mcd1, the yeast homology of human cohesin Rad21, plays an important role in hydrogen peroxide-induced apoptosis in yeast...
  61. Lee M, Yoon S, Kim K. Mitotic cohesin subunit Mcd1 regulates the progression of meiotic recombination in budding yeast. J Microbiol Biotechnol. 2015;25:598-605 pubmed
    ..Mcd1 (also known as Scc1), the α-kleisin subunit of cohesin, is a key regulatory subunit of the mitotic cohesin complex and is required ..
  62. McAleenan A, Clemente Blanco A, Cordon Preciado V, Sen N, Esteras M, Jarmuz A, et al. Post-replicative repair involves separase-dependent removal of the kleisin subunit of cohesin. Nature. 2013;493:250-4 pubmed publisher
    ..Moreover, a separase-resistant allele of the gene coding for the ?-kleisin subunit of cohesin, Mcd1 (also known as Scc1), reduces double-strand break resection and compromises the efficiency of repair even when loaded during DNA damage...
  63. 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
    ..We suggest that the function of the CC is to transfer ATP binding/hydrolysis signals between the head and the hinge domains. The results provide a new insight into the mechanism of cohesin activity. ..
  64. Sullivan M, Hornig N, Porstmann T, Uhlmann F. Studies on substrate recognition by the budding yeast separase. J Biol Chem. 2004;279:1191-6 pubmed
    Sister chromatid cohesion is resolved at anaphase onset when separase, a site-specific protease, cleaves the Scc1 subunit of the chromosomal cohesin complex that is responsible for holding sister chromatids together...
  65. Huber R, Kulemzina I, Ang K, Chavda A, Suranthran S, Teh J, et al. Impairing Cohesin Smc1/3 Head Engagement Compensates for the Lack of Eco1 Function. Structure. 2016;24:1991-1999 pubmed publisher
    The cohesin ring, which is composed of the Smc1, Smc3, and Scc1 subunits, topologically embraces two sister chromatids from S phase until anaphase to ensure their precise segregation to the daughter cells...
  66. Ghosh S, Gardner J, Smoyer C, Friederichs J, Unruh J, Slaughter B, et al. Acetylation of the SUN protein Mps3 by Eco1 regulates its function in nuclear organization. Mol Biol Cell. 2012;23:2546-59 pubmed publisher
    ..Acetylation of Mps3 by Eco1 is one of the few regulatory mechanisms known to control nuclear organization...
  67. Onn I, Guacci V, Koshland D. The zinc finger of Eco1 enhances its acetyltransferase activity during sister chromatid cohesion. Nucleic Acids Res. 2009;37:6126-34 pubmed publisher