Gene Symbol: CBF2
Description: Cbf2p
Alias: CEP2, CSL5, CTF14, NDC10, Cbf2p
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Goh P, Kilmartin J. NDC10: a gene involved in chromosome segregation in Saccharomyces cerevisiae. J Cell Biol. 1993;121:503-12 pubmed
    A mutant, ndc10-1, was isolated by anti-tubulin staining of temperature-sensitive mutant banks of budding yeast. ndc10-1 has a defect chromosome segregation since chromosomes remains at one pole of the anaphase spindle...
  2. Montpetit B, Hazbun T, Fields S, Hieter P. Sumoylation of the budding yeast kinetochore protein Ndc10 is required for Ndc10 spindle localization and regulation of anaphase spindle elongation. J Cell Biol. 2006;174:653-63 pubmed
    ..In this study, we show that the kinetochore proteins Ndc10, Bir1, Ndc80, and Cep3, which mediate the attachment of chromosomes to spindle microtubules, are sumoylated ..
  3. Measday V, Hailey D, Pot I, Givan S, Hyland K, Cagney G, et al. Ctf3p, the Mis6 budding yeast homolog, interacts with Mcm22p and Mcm16p at the yeast outer kinetochore. Genes Dev. 2002;16:101-13 pubmed
    ..In contrast, Ctf3p and Ctf19p fail to bind properly to the centromere in a cse4-1 mutant strain. We conclude that the requirements for CENP-A loading onto centromere DNA differ in fission versus budding yeast. ..
  4. Buvelot S, Tatsutani S, Vermaak D, Biggins S. The budding yeast Ipl1/Aurora protein kinase regulates mitotic spindle disassembly. J Cell Biol. 2003;160:329-39 pubmed
    ..We propose that Ipl1p regulates both the kinetochore and interpolar microtubule plus ends to regulate its various mitotic functions. ..
  5. Lechner J, Carbon J. A 240 kd multisubunit protein complex, CBF3, is a major component of the budding yeast centromere. Cell. 1991;64:717-25 pubmed
    ..A 56 bp region including the 26 bp CDEIII consensus is protected from DNAase I cleavage in the CBF3-CEN DNA complex. ..
  6. Wigge P, Kilmartin J. The Ndc80p complex from Saccharomyces cerevisiae contains conserved centromere components and has a function in chromosome segregation. J Cell Biol. 2001;152:349-60 pubmed
    ..Thus the Ndc80p complex contains centromere-associated components conserved between yeasts and vertebrates. ..
  7. Hyland K, Kingsbury J, Koshland D, Hieter P. Ctf19p: A novel kinetochore protein in Saccharomyces cerevisiae and a potential link between the kinetochore and mitotic spindle. J Cell Biol. 1999;145:15-28 pubmed
    ..We propose that Ctf19p is part of a macromolecular kinetochore complex, which may function as a link between the kinetochore and the mitotic spindle. ..
  8. Connelly C, Hieter P. Budding yeast SKP1 encodes an evolutionarily conserved kinetochore protein required for cell cycle progression. Cell. 1996;86:275-85 pubmed
    ..Skp1p therefore represents an intrinsic kinetochore protein conserved throughout eukaryotic evolution and may be directly involved in linking kinetochore function with the cell cycle-regulatory machinery. ..
  9. Gillis A, Thomas S, Hansen S, Kaplan K. A novel role for the CBF3 kinetochore-scaffold complex in regulating septin dynamics and cytokinesis. J Cell Biol. 2005;171:773-84 pubmed
    ..Mutants in Bir1p similarly affect septin organization, leading us to propose that CBF3 and Bir1p act as passenger proteins to coordinate chromosome segregation with cytokinesis. ..

More Information


  1. Sharp J, Franco A, Osley M, Kaufman P. Chromatin assembly factor I and Hir proteins contribute to building functional kinetochores in S. cerevisiae. Genes Dev. 2002;16:85-100 pubmed
    ..Finally, CAF-I subunits and Hir1 are enriched at centromeres, indicating that these proteins make a direct contribution to centromeric chromatin structures. ..
  2. Kopski K, Huffaker T. Suppressors of the ndc10-2 mutation: a role for the ubiquitin system in Saccharomyces cerevisiae kinetochore function. Genetics. 1997;147:409-20 pubmed
    We have isolated a new conditional-lethal mutation, ndc10-2, in the NDC10/CBF2/CTF14 gene that encodes the 110-kD subunit of the Saccharomyces cerevisiae CBF3 kinetochore complex...
  3. Yoon H, Carbon J. Participation of Bir1p, a member of the inhibitor of apoptosis family, in yeast chromosome segregation events. Proc Natl Acad Sci U S A. 1999;96:13208-13 pubmed
    ..Yeast cells deleted of BIR1 have a chromosome-loss phenotype, which can be completely rescued by elevating NDC10 dosage...
  4. Pinto I, Winston F. Histone H2A is required for normal centromere function in Saccharomyces cerevisiae. EMBO J. 2000;19:1598-612 pubmed
    ..Taken together, these results strongly suggest that histone H2A is required for proper centromere-kinetochore function during chromosome segregation. ..
  5. Stemmann O, Lechner J. The Saccharomyces cerevisiae kinetochore contains a cyclin-CDK complexing homologue, as identified by in vitro reconstitution. EMBO J. 1996;15:3611-20 pubmed
    ..Therefore, Cbf3d is the only CBF3 protein that has a known homologue in higher eukaryotes and may provide the anchor that directs cell cycle-regulated proteins to the kinetochore. ..
  6. Widlund P, Lyssand J, Anderson S, Niessen S, Yates J, Davis T. Phosphorylation of the chromosomal passenger protein Bir1 is required for localization of Ndc10 to the spindle during anaphase and full spindle elongation. Mol Biol Cell. 2006;17:1065-74 pubmed
    ..The middle region is not essential but is required to localize the inner kinetochore protein Ndc10 to the spindle during anaphase and to the midzone at telophase...
  7. Yoon H, Carbon J. Genetic and biochemical interactions between an essential kinetochore protein, Cbf2p/Ndc10p, and the CDC34 ubiquitin-conjugating enzyme. Mol Cell Biol. 1995;15:4835-42 pubmed
    CBF2/NDC10/CTF14 encodes the 110-kDa subunit of CBF3, a key component of the yeast centromere/kinetochore. Overexpression of yeast CDC34 specifically suppresses the temperature-sensitive growth phenotype of the ndc10-1 mutation...
  8. Li Y, Bachant J, Alcasabas A, Wang Y, Qin J, Elledge S. The mitotic spindle is required for loading of the DASH complex onto the kinetochore. Genes Dev. 2002;16:183-97 pubmed
    ..We propose that DASH is a microtubule-binding complex that is transferred to the kinetochore prior to mitosis, thereby defining a new step in kinetochore maturation. ..
  9. Rodrigo Brenni M, Thomas S, Bouck D, Kaplan K. Sgt1p and Skp1p modulate the assembly and turnover of CBF3 complexes required for proper kinetochore function. Mol Biol Cell. 2004;15:3366-78 pubmed
    ..The assembly of CBF3 is opposed by its turnover and disruption of this balance compromises kinetochore function without affecting kinetochore formation on centromeric DNA. ..
  10. Meluh P, Koshland D. Evidence that the MIF2 gene of Saccharomyces cerevisiae encodes a centromere protein with homology to the mammalian centromere protein CENP-C. Mol Biol Cell. 1995;6:793-807 pubmed
    ..between mutations in MIF2 and trans-acting mutations in three known yeast centromere protein genes-CEP1/CBF1/CPF1, NDC10/CBF2, and CEP3/CBF3B...
  11. Camahort R, Li B, Florens L, Swanson S, Washburn M, Gerton J. Scm3 is essential to recruit the histone h3 variant cse4 to centromeres and to maintain a functional kinetochore. Mol Cell. 2007;26:853-65 pubmed
    ..Scm3 interacts with both Ndc10 and Cse4 and is essential to establish centromeric chromatin after DNA replication...
  12. Kroll E, Hyland K, Hieter P, Li J. Establishing genetic interactions by a synthetic dosage lethality phenotype. Genetics. 1996;143:95-102 pubmed
    ..CTF13 overexpression caused synthetic dosage lethality in combination with ctf14-42 (cbf2, ndc10), ctf17-61 (chl4), ctf19-58 and ctf19-26...
  13. Kaplan K, Hyman A, Sorger P. Regulating the yeast kinetochore by ubiquitin-dependent degradation and Skp1p-mediated phosphorylation. Cell. 1997;91:491-500 pubmed
    ..We propose that coupled activation and destruction link the assembly of Cbf3p to the duplication of centromeres in S phase. ..
  14. Russell I, Grancell A, Sorger P. The unstable F-box protein p58-Ctf13 forms the structural core of the CBF3 kinetochore complex. J Cell Biol. 1999;145:933-50 pubmed
    ..subunits of CBF3 are multimeric in solution: p23(Skp1) and p58(Ctf13) form a heterodimer, and p64(Cep3) and p110(Ndc10) form homodimers...
  15. Jiang W, Koltin Y. Two-hybrid interaction of a human UBC9 homolog with centromere proteins of Saccharomyces cerevisiae. Mol Gen Genet. 1996;251:153-60 pubmed
    ..We also provide further in vivo evidence that Mck1p, a protein kinase, is specifically associated with the centromere proteins Cbf2p and Cbf5p, which were previously shown to interact in vitro.
  16. Bouck D, Bloom K. The kinetochore protein Ndc10p is required for spindle stability and cytokinesis in yeast. Proc Natl Acad Sci U S A. 2005;102:5408-13 pubmed
    ..This latter phenotype leads to a cell separation defect in ndc10-1 cells...
  17. Ravid T, Kreft S, Hochstrasser M. Membrane and soluble substrates of the Doa10 ubiquitin ligase are degraded by distinct pathways. EMBO J. 2006;25:533-43 pubmed
    ..Thus, while Doa10 ubiquitinates both membrane and soluble proteins, the mechanisms of subsequent proteasome targeting differ. ..
  18. Basrai M, Kingsbury J, Koshland D, Spencer F, Hieter P. Faithful chromosome transmission requires Spt4p, a putative regulator of chromatin structure in Saccharomyces cerevisiae. Mol Cell Biol. 1996;16:2838-47 pubmed
    ..Double mutants carrying spt4 delta 2::HIS3 and kinetochore mutation ndc10-42 or ctf13-30 show a synthetic conditional phenotype...
  19. Biggins S, Severin F, Bhalla N, Sassoon I, Hyman A, Murray A. The conserved protein kinase Ipl1 regulates microtubule binding to kinetochores in budding yeast. Genes Dev. 1999;13:532-44 pubmed
    ..Because aurora2 has been implicated in oncogenesis, defects in kinetochore function may contribute to genetic instability in human tumors. ..
  20. Ortiz J, Stemmann O, Rank S, Lechner J. A putative protein complex consisting of Ctf19, Mcm21, and Okp1 represents a missing link in the budding yeast kinetochore. Genes Dev. 1999;13:1140-55 pubmed
  21. Romao M, Tanaka K, Sibarita J, Ly Hartig N, Tanaka T, Antony C. Three-dimensional electron microscopy analysis of ndc10-1 mutant reveals an aberrant organization of the mitotic spindle and spindle pole body defects in Saccharomyces cerevisiae. J Struct Biol. 2008;163:18-28 pubmed publisher
    ..b>ndc10-1 mutant was shown to display chromosome mis-segregation as well as an aberrant mitotic spindle (Goh and Kilmartin,..
  22. Jones M, He X, Giddings T, Winey M. Yeast Dam1p has a role at the kinetochore in assembly of the mitotic spindle. Proc Natl Acad Sci U S A. 2001;98:13675-80 pubmed
    ..An epitope-tagged version of Dam1p colocalizes with the integral kinetochore component Ndc10p/Cbf2p in immunofluorescence analysis of chromosome spreads...
  23. Meluh P, Koshland D. Budding yeast centromere composition and assembly as revealed by in vivo cross-linking. Genes Dev. 1997;11:3401-12 pubmed
    ..Finally, given the sensitivity of this technique, its application to other sequence-specific protein-DNA complexes within the cell, such as origins of replication and enhancer-promoter regions, could be of significant value. ..
  24. Winkler A, Korstanje R, Zonneveld B, Hooykaas P, Steensma H. Isolation and characterization of KIUBP2, a ubiquitin hydrolase gene of Kluyveromyces lactis that can suppress a ts-mutation in CBF2, a gene encoding a centromeric protein of Saccharomyces cerevisiae. Curr Genet. 2000;38:17-22 pubmed
    The Kluyveromyces lactis UBP2 gene was isolated as a suppressor of a temperature-sensitive mutation in CBF2, a gene coding for a centromere-binding protein of Saccharomyces cerevisiae...
  25. Au W, Crisp M, DeLuca S, Rando O, Basrai M. Altered dosage and mislocalization of histone H3 and Cse4p lead to chromosome loss in Saccharomyces cerevisiae. Genetics. 2008;179:263-75 pubmed publisher
    ..Suppression of these phenotypes by histone delta 16H3 and GALCSE4 supports the conclusion that proper stoichiometry affects the localization of histone H3 and Cse4p and is thus essential for accurate chromosome segregation. ..
  26. Makhnevych T, Wong P, Pogoutse O, Vizeacoumar F, Greenblatt J, Emili A, et al. Hsp110 is required for spindle length control. J Cell Biol. 2012;198:623-36 pubmed publisher
    ..These data suggest a model whereby the Hsp70-Hsp110 chaperone complex antagonizes Cin8 plus-end motility and prevents premature spindle elongation in S phase. ..
  27. Mythreye K, Bloom K. Differential kinetochore protein requirements for establishment versus propagation of centromere activity in Saccharomyces cerevisiae. J Cell Biol. 2003;160:833-43 pubmed
    ..Furthermore, this study identifies Chl4p in the initiation and specification of a heritable chromatin state. ..
  28. Ma L, Ho K, Piggott N, Luo Z, Measday V. Interactions between the kinetochore complex and the protein kinase A pathway in Saccharomyces cerevisiae. G3 (Bethesda). 2012;2:831-41 pubmed publisher
    ..we show that lowering protein kinase A activity can rescue the chromosome loss defect of the inner kinetochore ndc10 mutant...
  29. Pietrasanta L, Thrower D, Hsieh W, Rao S, Stemmann O, Lechner J, et al. Probing the Saccharomyces cerevisiae centromeric DNA (CEN DNA)-binding factor 3 (CBF3) kinetochore complex by using atomic force microscopy. Proc Natl Acad Sci U S A. 1999;96:3757-62 pubmed
    ..These multi-armed complexes indicate that CBF3 can bind two DNA molecules together in vitro and, thus, may be involved in holding together chromatid pairs during mitosis. ..
  30. Shang C, Hazbun T, Cheeseman I, Aranda J, Fields S, Drubin D, et al. Kinetochore protein interactions and their regulation by the Aurora kinase Ipl1p. Mol Biol Cell. 2003;14:3342-55 pubmed
    ..Strikingly, we found that the Dam1p-Ndc80p and Dam1p-Spc34p interactions were weakened by mutations mimicking phosphorylation at Ipl1p sites, allowing us to formulate a model for the effects of phosphoregulation on kinetochore function. ..
  31. Stoepel J, Ottey M, Kurischko C, Hieter P, Luca F. The mitotic exit network Mob1p-Dbf2p kinase complex localizes to the nucleus and regulates passenger protein localization. Mol Biol Cell. 2005;16:5465-79 pubmed
    ..These findings reveal kinetochores as sites for MEN signaling and implicate MEN in coordinating chromosome segregation and/or spindle integrity with mitotic exit and cytokinesis via regulation of chromosome passenger proteins. ..
  32. Collins K, Castillo A, Tatsutani S, Biggins S. De novo kinetochore assembly requires the centromeric histone H3 variant. Mol Biol Cell. 2005;16:5649-60 pubmed
    ..Consistent with this observation, Cse4-depleted cells had severe chromosome segregation defects. We therefore propose that yeast kinetochore assembly requires both centromeric DNA specificity and centromeric chromatin. ..
  33. Thomas S, Kaplan K. A Bir1p Sli15p kinetochore passenger complex regulates septin organization during anaphase. Mol Biol Cell. 2007;18:3820-34 pubmed
    ..We present a model for how discrete passenger complexes coordinate distinct aspects of mitosis. ..
  34. Lin H, Choi J, Hasek J, DeLillo N, Lou W, Vancura A. Phospholipase C is involved in kinetochore function in Saccharomyces cerevisiae. Mol Cell Biol. 2000;20:3597-607 pubmed
    ..These results suggest that Plc1p associates with kinetochores and regulates some aspect of kinetochore function and demonstrate an intranuclear function of phospholipase C in eukaryotic cells. ..
  35. Schweiggert J, Stevermann L, Panigada D, Kammerer D, Liakopoulos D. Regulation of a Spindle Positioning Factor at Kinetochores by SUMO-Targeted Ubiquitin Ligases. Dev Cell. 2016;36:415-27 pubmed publisher
    ..We propose that kinetochores serve as platforms to recruit STUbLs in a SUMO-dependent manner in order to ensure correct spindle function by regulating levels of microtubule-associated proteins. ..
  36. Hemmerich P, Stoyan T, Wieland G, Koch M, Lechner J, Diekmann S. Interaction of yeast kinetochore proteins with centromere-protein/transcription factor Cbf1. Proc Natl Acad Sci U S A. 2000;97:12583-8 pubmed
    ..A functional comparison of the centromere binding proteins with transcription factors binding at MET16 promoters reveals the strong analogy between centromeres and the MET16 promoter. ..
  37. Kanta H, Laprade L, Almutairi A, Pinto I. Suppressor analysis of a histone defect identifies a new function for the hda1 complex in chromosome segregation. Genetics. 2006;173:435-50 pubmed
    ..Moreover, these suppressors provide the basis for future studies regarding histone function in chromosome segregation. ..
  38. Pearson C, Maddox P, Zarzar T, Salmon E, Bloom K. Yeast kinetochores do not stabilize Stu2p-dependent spindle microtubule dynamics. Mol Biol Cell. 2003;14:4181-95 pubmed
    ..Loss of the kinetochore (ndc10-1, ndc10-2, and ctf13-30) does not drastically affect spindle microtubule turnover, indicating that Stu2p, not the ..
  39. DeLillo N, Romero C, Lin H, Vancura A. Genetic evidence for a role of phospholipase C at the budding yeast kinetochore. Mol Genet Genomics. 2003;269:261-70 pubmed
    ..PLC1 displays strong genetic interactions with genes encoding components of the inner kinetochore, including NDC10, SKP1, MIF2, CEP1, CEP3, and CTF13...
  40. Murillo Pineda M, Cabello Lobato M, Clemente Ruiz M, Monje Casas F, Prado F. Defective histone supply causes condensin-dependent chromatin alterations, SAC activation and chromosome decatenation impairment. Nucleic Acids Res. 2014;42:12469-82 pubmed publisher
    ..Therefore, our results reveal the importance of a precise interplay between histone supply and condensin/Top2 for pericentric chromatin structure, precatenanes resolution and centromere biorientation. ..
  41. Bock L, Pagliuca C, Kobayashi N, Grove R, Oku Y, Shrestha K, et al. Cnn1 inhibits the interactions between the KMN complexes of the yeast kinetochore. Nat Cell Biol. 2012;14:614-24 pubmed publisher
    ..Cnn1 regulates KMN activity in a spatiotemporal manner by inhibiting the interaction between its complexes. Cnn1 activity peaks in anaphase and is driven by the Cdc28, Mps1 and Ipl1 kinases. ..
  42. Perriches T, Singleton M. Structure of yeast kinetochore Ndc10 DNA-binding domain reveals unexpected evolutionary relationship to tyrosine recombinases. J Biol Chem. 2012;287:5173-9 pubmed publisher
    We have solved the x-ray structure of the N-terminal half of the yeast kinetochore protein Ndc10 at 1.9 Å resolution...
  43. Baker R, Harris K, Zhang K. Mutations synthetically lethal with cep1 target S. cerevisiae kinetochore components. Genetics. 1998;149:73-85 pubmed
    ..Three of these four CSL genes were subsequently found to be known or imputed kinetochore genes: CEP3, NDC10, and CSE4. The fourth, CSL4, corresponded to ORF YNL232w on chromosome XIV, and was found to be essential...
  44. Scott R, Cairo L, Van de Vosse D, Wozniak R. The nuclear export factor Xpo1p targets Mad1p to kinetochores in yeast. J Cell Biol. 2009;184:21-9 pubmed publisher
    ..These results reveal an important function for Xpo1p in mediating intranuclear transport events and identify a signaling pathway between kinetochores and NPCs. ..
  45. Samel A, Cuomo A, Bonaldi T, Ehrenhofer Murray A. Methylation of CenH3 arginine 37 regulates kinetochore integrity and chromosome segregation. Proc Natl Acad Sci U S A. 2012;109:9029-34 pubmed publisher
    ..Altogether, our data identify a unique regulatory principle on centromeric chromatin by posttranslational modification of the amino terminus of CenH3. ..
  46. Peng Y, Wong C, Nakajima Y, Tyers R, Sarkeshik A, Yates J, et al. Overlapping kinetochore targets of CK2 and Aurora B kinases in mitotic regulation. Mol Biol Cell. 2011;22:2680-9 pubmed publisher
    ..Furthermore, we show Mif2 and the inner kinetochore protein Ndc10 are phosphorylated by CK2, and this phosphorylation plays antagonistic and synergistic roles with Aurora B ..
  47. Furth N, Gertman O, Shiber A, Alfassy O, Cohen I, Rosenberg M, et al. Exposure of bipartite hydrophobic signal triggers nuclear quality control of Ndc10 at the endoplasmic reticulum/nuclear envelope. Mol Biol Cell. 2011;22:4726-39 pubmed publisher
    ..These findings substantiate the ability of the ER quality control system to recognize subtle perturbation(s) in the native structure of a nuclear protein. ..
  48. Cho U, Harrison S. Ndc10 is a platform for inner kinetochore assembly in budding yeast. Nat Struct Mol Biol. 2011;19:48-55 pubmed publisher
    ..In point-centromere yeasts, the CBF3 complex Skp1-Ctf13-(Cep3)(2)-(Ndc10)(2) recognizes a conserved centromeric DNA element through contacts made by Cep3 and Ndc10...
  49. Dodgson S, Santaguida S, Kim S, Sheltzer J, Amon A. The pleiotropic deubiquitinase Ubp3 confers aneuploidy tolerance. Genes Dev. 2016;30:2259-2271 pubmed
    ..We propose that interfering with Ubp3/USP10 function could be a productive avenue in the development of novel cancer therapeutics. ..
  50. 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. ..
  51. Lee B, Kiburz B, Amon A. Spo13 maintains centromeric cohesion and kinetochore coorientation during meiosis I. Curr Biol. 2004;14:2168-82 pubmed
  52. Hsu J, Huang J, Meluh P, Laurent B. The yeast RSC chromatin-remodeling complex is required for kinetochore function in chromosome segregation. Mol Cell Biol. 2003;23:3202-15 pubmed
  53. Pot I, Measday V, Snydsman B, Cagney G, Fields S, Davis T, et al. Chl4p and iml3p are two new members of the budding yeast outer kinetochore. Mol Biol Cell. 2003;14:460-76 pubmed
    ..These physical interaction dependencies provide insights into the molecular architecture and centromere DNA loading requirements of the outer kinetochore complex. ..
  54. Jiang W, Lim M, Yoon H, Thorner J, Martin G, Carbon J. Overexpression of the yeast MCK1 protein kinase suppresses conditional mutations in centromere-binding protein genes CBF2 and CBF5. Mol Gen Genet. 1995;246:360-6 pubmed
    ..support of this model was provided by the finding that purified Mck1p phosphorylates in vitro the 110 kDa subunit (Cbf2p) of the multimeric centromere binding factor CBF3...
  55. Li Z, Vizeacoumar F, Bahr S, Li J, Warringer J, Vizeacoumar F, et al. Systematic exploration of essential yeast gene function with temperature-sensitive mutants. Nat Biotechnol. 2011;29:361-7 pubmed publisher
    ..This mutant collection should facilitate a wide range of systematic studies aimed at understanding the functions of essential genes. ..
  56. Shivaraju M, Camahort R, Mattingly M, Gerton J. Scm3 is a centromeric nucleosome assembly factor. J Biol Chem. 2011;286:12016-23 pubmed publisher
    ..assembly activity depend on an evolutionarily conserved core motif in Scm3, but localization of the CBF3 subunit Ndc10 to centromeres does not depend on this motif...