Gene Symbol: CDC20
Description: ubiquitin-protein transferase activating protein CDC20
Alias: PAC5, ubiquitin-protein transferase activating protein CDC20
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Holt L, Krutchinsky A, Morgan D. Positive feedback sharpens the anaphase switch. Nature. 2008;454:353-7 pubmed publisher
    ..Our results also suggest that coupling securin degradation with changes in Cdk1 and Cdc14 activities helps coordinate the initiation of sister-chromatid separation with changes in spindle dynamics. ..
  2. Poddar A, Stukenberg P, Burke D. Two complexes of spindle checkpoint proteins containing Cdc20 and Mad2 assemble during mitosis independently of the kinetochore in Saccharomyces cerevisiae. Eukaryot Cell. 2005;4:867-78 pubmed
    Favored models of spindle checkpoint signaling propose that two inhibitory complexes (Mad2-Cdc20 and Mad2-Mad3-Bub3-Cdc20) must be assembled at kinetochores in order to inhibit mitosis...
  3. Kraft C, Vodermaier H, Maurer Stroh S, Eisenhaber F, Peters J. The WD40 propeller domain of Cdh1 functions as a destruction box receptor for APC/C substrates. Mol Cell. 2005;18:543-53 pubmed
    Activation of the anaphase-promoting complex/cyclosome (APC/C) by Cdc20 and Cdh1 leads to ubiquitin-dependent degradation of securin and cyclin B and thereby promotes the initiation of anaphase and exit from mitosis...
  4. Nezi L, Rancati G, De Antoni A, Pasqualato S, Piatti S, Musacchio A. Accumulation of Mad2-Cdc20 complex during spindle checkpoint activation requires binding of open and closed conformers of Mad2 in Saccharomyces cerevisiae. J Cell Biol. 2006;174:39-51 pubmed
    ..C-Mad2 forms when Mad2 binds its checkpoint target Cdc20 or its kinetochore receptor Mad1. When unbound to these ligands, Mad2 folds as O-Mad2...
  5. Schwab M, Lutum A, Seufert W. Yeast Hct1 is a regulator of Clb2 cyclin proteolysis. Cell. 1997;90:683-93 pubmed
    ..Hct1 and the related Cdc20 may function as substrate-specific regulators of proteolysis during mitosis.
  6. Tan G, Magurno J, Cooper K. Ama1p-activated anaphase-promoting complex regulates the destruction of Cdc20p during meiosis II. Mol Biol Cell. 2011;22:315-26 pubmed publisher
    ..These findings provide a mechanism to temporally down-regulate APC/C(Cdc20) activity as the cells complete meiosis II and form spores.
  7. Wang Y, Burke D. Cdc55p, the B-type regulatory subunit of protein phosphatase 2A, has multiple functions in mitosis and is required for the kinetochore/spindle checkpoint in Saccharomyces cerevisiae. Mol Cell Biol. 1997;17:620-6 pubmed
    ..Finally, cdc55::LEU2 suppresses the temperature sensitivity of cdc20-1, suggesting additional roles for CDC55 in mitosis.
  8. Pan J, Chen R. Spindle checkpoint regulates Cdc20p stability in Saccharomyces cerevisiae. Genes Dev. 2004;18:1439-51 pubmed
    ..wild-type Saccharomyces cerevisiae contains less Cdc20p than spindle checkpoint mutants do, whereas their CDC20 mRNA levels are similar...
  9. Camasses A, Bogdanova A, Shevchenko A, Zachariae W. The CCT chaperonin promotes activation of the anaphase-promoting complex through the generation of functional Cdc20. Mol Cell. 2003;12:87-100 pubmed
    The WD repeat protein Cdc20 is essential for progression through mitosis because it is required to activate ubiquitin ligation by the anaphase-promoting complex (APC/C)...

More Information


  1. Foster S, Morgan D. The APC/C subunit Mnd2/Apc15 promotes Cdc20 autoubiquitination and spindle assembly checkpoint inactivation. Mol Cell. 2012;47:921-32 pubmed publisher
    ..of unattached kinetochores, anaphase is delayed when three SAC components (Mad2, Mad3/BubR1, and Bub3) inhibit Cdc20, the activating subunit of the anaphase-promoting complex (APC/C)...
  2. King E, van der Sar S, Hardwick K. Mad3 KEN boxes mediate both Cdc20 and Mad3 turnover, and are critical for the spindle checkpoint. PLoS ONE. 2007;2:e342 pubmed
    ..In the presence of defective kinetochore-microtubule interactions, APC/C(Cdc20) is inhibited by the spindle checkpoint, thereby delaying anaphase onset and providing more time for spindle ..
  3. Zachariae W, Nasmyth K. Whose end is destruction: cell division and the anaphase-promoting complex. Genes Dev. 1999;13:2039-58 pubmed
  4. Shirayama M, Toth A, Galova M, Nasmyth K. APC(Cdc20) promotes exit from mitosis by destroying the anaphase inhibitor Pds1 and cyclin Clb5. Nature. 1999;402:203-7 pubmed
    ..release of Cdc14 and enables it to promote Cdk1 inactivation, but it is known to be dependent on APC/C bound by Cdc20 (APC(Cdc20)) (ref. 4)...
  5. Rudner A, Hardwick K, Murray A. Cdc28 activates exit from mitosis in budding yeast. J Cell Biol. 2000;149:1361-76 pubmed
    ..CDC28-VF has reduced Cdc20- dependent APC activity in mitosis, but normal Hct1- dependent APC activity in the G1 phase of the cell cycle...
  6. Shirayama M, Zachariae W, Ciosk R, Nasmyth K. The Polo-like kinase Cdc5p and the WD-repeat protein Cdc20p/fizzy are regulators and substrates of the anaphase promoting complex in Saccharomyces cerevisiae. EMBO J. 1998;17:1336-49 pubmed
    ..Accumulation of Cdc20p contributes to activation of Pds1p proteolysis in metaphase, whereas accumulation of Cdc5p facilitates the activation of Clb2p proteolysis. ..
  7. Burton J, Solomon M. Mad3p, a pseudosubstrate inhibitor of APCCdc20 in the spindle assembly checkpoint. Genes Dev. 2007;21:655-67 pubmed
    ..The checkpoint proteins Mad2 and Mad3/BubR1 bind to Cdc20, although how they inhibit APCCdc20 is unclear...
  8. Lee B, Amon A. Role of Polo-like kinase CDC5 in programming meiosis I chromosome segregation. Science. 2003;300:482-6 pubmed
    ..Thus, sister-kinetochore coorientation and chromosome segregation during meiosis I are coupled through their dependence on CDC5. ..
  9. Hilioti Z, Chung Y, Mochizuki Y, Hardy C, COHEN FIX O. The anaphase inhibitor Pds1 binds to the APC/C-associated protein Cdc20 in a destruction box-dependent manner. Curr Biol. 2001;11:1347-52 pubmed
    ..In mitotic cells, two forms of the APC/C exist, APC/C(Cdc20) and APC/C(Cdh1), which differ in their associated WD-repeat proteins (Cdc20 and Cdh1, respectively), time of ..
  10. Passmore L, McCormack E, Au S, Paul A, Willison K, Harper J, et al. Doc1 mediates the activity of the anaphase-promoting complex by contributing to substrate recognition. EMBO J. 2003;22:786-96 pubmed
    ..These results imply that Doc1p/Apc10 may play a role to regulate the binding of specific substrates, similar to that of the coactivators. ..
  11. Salah S, Nasmyth K. Destruction of the securin Pds1p occurs at the onset of anaphase during both meiotic divisions in yeast. Chromosoma. 2000;109:27-34 pubmed
    ..This process is mediated by the anaphase promoting complex and an accessory protein Cdc20. In meiosis a single round of DNA replication is followed by two successive rounds of segregation...
  12. Enquist Newman M, Sullivan M, Morgan D. Modulation of the mitotic regulatory network by APC-dependent destruction of the Cdh1 inhibitor Acm1. Mol Cell. 2008;30:437-46 pubmed publisher
    ..The APC is activated by association with Cdc20 in midmitosis and Cdh1 in late mitosis and G1...
  13. Fraschini R, Beretta A, Sironi L, Musacchio A, Lucchini G, Piatti S. Bub3 interaction with Mad2, Mad3 and Cdc20 is mediated by WD40 repeats and does not require intact kinetochores. EMBO J. 2001;20:6648-59 pubmed
    ..and Mps1 proteins, prevents anaphase entry and mitotic exit by inhibiting the anaphase promoting complex activator Cdc20 in response to monopolar attachment of sister kinetochores to spindle fibres...
  14. Matyskiela M, Morgan D. Analysis of activator-binding sites on the APC/C supports a cooperative substrate-binding mechanism. Mol Cell. 2009;34:68-80 pubmed publisher
    ..Substrates are recruited to the APC/C by activator proteins (Cdc20 or Cdh1), but it is not known where substrates are bound during catalysis...
  15. Tinker Kulberg R, Morgan D. Pds1 and Esp1 control both anaphase and mitotic exit in normal cells and after DNA damage. Genes Dev. 1999;13:1936-49 pubmed
    ..The effects of Esp1 are dependent on Cdc20 (an activating subunit of the APC) and on several additional proteins (Cdc5, Cdc14, Cdc15, Tem1) that form a ..
  16. Mallory M, Cooper K, Strich R. Meiosis-specific destruction of the Ume6p repressor by the Cdc20-directed APC/C. Mol Cell. 2007;27:951-61 pubmed
    ..This control appears direct as Cdc20p and Ume6p associate in vivo and APC/C(Cdc20) ubiquitylates Ume6p in vitro...
  17. Foe I, Foster S, Cheung S, DeLuca S, Morgan D, Toczyski D. Ubiquitination of Cdc20 by the APC occurs through an intramolecular mechanism. Curr Biol. 2011;21:1870-7 pubmed publisher
    Cells control progression through late mitosis by regulating Cdc20 and Cdh1, the two mitotic activators of the anaphase-promoting complex (APC). The control of Cdc20 protein levels during the cell cycle is not well understood...
  18. Burton J, Solomon M. D box and KEN box motifs in budding yeast Hsl1p are required for APC-mediated degradation and direct binding to Cdc20p and Cdh1p. Genes Dev. 2001;15:2381-95 pubmed
    ..contains both a destruction box (D box) and a KEN box motif that are necessary for Hsl1p turnover with either APC(Cdc20) or APC(Cdh1)...
  19. Visintin R, Craig K, Hwang E, Prinz S, Tyers M, Amon A. The phosphatase Cdc14 triggers mitotic exit by reversal of Cdk-dependent phosphorylation. Mol Cell. 1998;2:709-18 pubmed
    ..Feedback between these pathways may lead to precipitous collapse of mitotic CDK activity and help coordinate exit from mitosis. ..
  20. Schwab M, Neutzner M, Mocker D, Seufert W. Yeast Hct1 recognizes the mitotic cyclin Clb2 and other substrates of the ubiquitin ligase APC. EMBO J. 2001;20:5165-75 pubmed
    ..Previous studies in the yeast Saccharomyces cerevisiae identified the conserved WD40 proteins Cdc20 and Hct1 (Cdh1) as substrate-specific activators of the APC, but their precise mechanism of action has remained ..
  21. Hardwick K, Johnston R, Smith D, Murray A. MAD3 encodes a novel component of the spindle checkpoint which interacts with Bub3p, Cdc20p, and Mad2p. J Cell Biol. 2000;148:871-82 pubmed
    ..We discuss roles for Mad3p and its interactions with other spindle checkpoint proteins and with Cdc20p, the target of the checkpoint. ..
  22. Searle J, Schollaert K, Wilkins B, Sanchez Y. The DNA damage checkpoint and PKA pathways converge on APC substrates and Cdc20 to regulate mitotic progression. Nat Cell Biol. 2004;6:138-45 pubmed
    ..protein kinase (PKA) pathway supports Chk1 in the regulation of mitosis by targeting the mitotic inducer Cdc20. Cdc20 is phosphorylated on PKA consensus sites after DNA damage, and this phosphorylation requires the Atr ..
  23. Hwang L, Lau L, Smith D, Mistrot C, Hardwick K, Hwang E, et al. Budding yeast Cdc20: a target of the spindle checkpoint. Science. 1998;279:1041-4 pubmed
    ..system, three proteins that are components of the checkpoint, Mad1, Mad2, and Mad3, were shown to interact with Cdc20, a protein required for exit from mitosis. Mad2 and Mad3 coprecipitated with Cdc20 at all stages of the cell cycle...
  24. Lim H, Goh P, Surana U. Cdc20 is essential for the cyclosome-mediated proteolysis of both Pds1 and Clb2 during M phase in budding yeast. Curr Biol. 1998;8:231-4 pubmed
    ..In Saccharomyces cerevisiae, chromosome segregation also requires the CDC20 gene, whose product contains WD40 repeats [11,12]...
  25. Visintin R, Prinz S, Amon A. CDC20 and CDH1: a family of substrate-specific activators of APC-dependent proteolysis. Science. 1997;278:460-3 pubmed
    ..The conserved Cdc20 and Cdh1 proteins were identified as limiting, substrate-specific activators of APC-dependent proteolysis...
  26. Zachariae W, Schwab M, Nasmyth K, Seufert W. Control of cyclin ubiquitination by CDK-regulated binding of Hct1 to the anaphase promoting complex. Science. 1998;282:1721-4 pubmed
    ..The mutual inhibition between APC and CDKs explains how cells suppress mitotic CDK activity during G1 and then establish a period with elevated kinase activity from S phase until anaphase. ..
  27. Yamamoto A, Guacci V, Koshland D. Pds1p, an inhibitor of anaphase in budding yeast, plays a critical role in the APC and checkpoint pathway(s). J Cell Biol. 1996;133:99-110 pubmed
    ..Therefore, in the DNA-damage checkpoint, Pds1p is required for the inhibition of cytokinesis and DNA replication as well as anaphase. The role of Pds1 protein in anaphase inhibition and general cell cycle regulation is discussed. ..
  28. Oelschlaegel T, Schwickart M, Matos J, Bogdanova A, Camasses A, Havlis J, et al. The yeast APC/C subunit Mnd2 prevents premature sister chromatid separation triggered by the meiosis-specific APC/C-Ama1. Cell. 2005;120:773-88 pubmed
    ..In vitro, Mnd2 inhibits ubiquitination of Pds1 by APC/C-Ama1 but not by other APC/C holo-enzymes. We conclude that chromosome segregation in meiosis depends on the selective inhibition of a meiosis-specific form of the APC/C. ..
  29. Simonetta M, Manzoni R, Mosca R, Mapelli M, Massimiliano L, Vink M, et al. The influence of catalysis on mad2 activation dynamics. PLoS Biol. 2009;7:e10 pubmed publisher
    ..The target of Mad2 is Cdc20, an activator of the anaphase-promoting complex/cyclosome (APC/C)...
  30. Rossio V, Kazatskaya A, Hirabayashi M, Yoshida S. Comparative genetic analysis of PP2A-Cdc55 regulators in budding yeast. Cell Cycle. 2014;13:2073-83 pubmed publisher
  31. Burton J, Solomon M. Hsl1p, a Swe1p inhibitor, is degraded via the anaphase-promoting complex. Mol Cell Biol. 2000;20:4614-25 pubmed
  32. Schibler A, Koutelou E, Tomida J, Wilson Pham M, Wang L, Lu Y, et al. Histone H3K4 methylation regulates deactivation of the spindle assembly checkpoint through direct binding of Mad2. Genes Dev. 2016;30:1187-97 pubmed publisher
    ..These proteins inhibit Cdc20, an activator of the anaphase-promoting complex/cyclosome (APC/C)...
  33. 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. ..
  34. Katis V, Matos J, Mori S, Shirahige K, Zachariae W, Nasmyth K. Spo13 facilitates monopolin recruitment to kinetochores and regulates maintenance of centromeric cohesion during yeast meiosis. Curr Biol. 2004;14:2183-96 pubmed
    ..Crucially, cohesin at centromeres is retained to allow sister centromeres to biorient at the second division. Little is known about how these meiosis I-specific events are regulated...
  35. Agarwal R, Tang Z, Yu H, COHEN FIX O. Two distinct pathways for inhibiting pds1 ubiquitination in response to DNA damage. J Biol Chem. 2003;278:45027-33 pubmed
    ..on its ubiquitination by the anaphase-promoting complex/cyclosome ubiquitin ligase, acting in conjunction with the Cdc20 protein (APC/CCdc20)...
  36. Liang H, Lim H, Venkitaraman A, Surana U. Cdk1 promotes kinetochore bi-orientation and regulates Cdc20 expression during recovery from spindle checkpoint arrest. EMBO J. 2012;31:403-16 pubmed publisher
    ..Moreover, Cdk1 is essential for sustaining the expression of Cdc20, an activator of the anaphase promoting complex/cyclosome (APC/C) required for anaphase progression...
  37. Tsuchiya D, Gonzalez C, Lacefield S. The spindle checkpoint protein Mad2 regulates APC/C activity during prometaphase and metaphase of meiosis I in Saccharomyces cerevisiae. Mol Biol Cell. 2011;22:2848-61 pubmed publisher
    ..We find that Mad2 sets the duration of meiosis I by regulating the activity of APC(Cdc20)...
  38. Tavormina P, Burke D. Cell cycle arrest in cdc20 mutants of Saccharomyces cerevisiae is independent of Ndc10p and kinetochore function but requires a subset of spindle checkpoint genes. Genetics. 1998;148:1701-13 pubmed
    ..We examined the role of the spindle checkpoint in the arrest in cdc20 mutants that arrest prior to anaphase with an aberrant spindle...
  39. 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. ..
  40. Lau D, Murray A. Mad2 and Mad3 cooperate to arrest budding yeast in mitosis. Curr Biol. 2012;22:180-90 pubmed publisher
    ..tension and arrests cells at metaphase by inhibiting the anaphase-promoting complex (APC) and its coactivator Cdc20. Despite numerous studies, we still do not understand how the checkpoint proteins coordinate with each other to ..
  41. Taberner F, Quilis I, Sendra J, Bañó M, Igual J. Regulation of cell cycle transcription factor Swi5 by karyopherin Msn5. Biochim Biophys Acta. 2012;1823:959-70 pubmed publisher
    ..This deleterious effect was eliminated when Swi5 nuclear import was abrogated, suggesting that nuclear export by Msn5 is important for cell physiology, because it prevents toxic Swi5 nuclear accumulation. ..
  42. Arnold L, Höckner S, Seufert W. Insights into the cellular mechanism of the yeast ubiquitin ligase APC/C-Cdh1 from the analysis of in vivo degrons. Mol Biol Cell. 2015;26:843-58 pubmed publisher
    ..Substrate selection by this ubiquitin ligase depends on related activator proteins, Cdc20 and Cdh1, which bind and activate the APC/C at distinct cell cycle stages...
  43. Schuyler S, Murray A. An in vitro assay for Cdc20-dependent mitotic anaphase-promoting complex activity from budding yeast. Methods Mol Biol. 2009;545:271-85 pubmed publisher
    ..ligase called the cyclosome or Anaphase-Promoting Complex (APC), and a WD40-repeat protein co-factor called Cdc20. In vitro Cdc20-dependent APC (APC(Cdc20)) assays have been useful in the identification and validation of target ..
  44. Indjeian V, Stern B, Murray A. The centromeric protein Sgo1 is required to sense lack of tension on mitotic chromosomes. Science. 2005;307:130-3 pubmed
  45. Tan G, Lewandowski R, Mallory M, Strich R, Cooper K. Mutually dependent degradation of Ama1p and Cdc20p terminates APC/C ubiquitin ligase activity at the completion of meiotic development in yeast. Cell Div. 2013;8:9 pubmed publisher
    ..In addition, these results support a model in which Ama1p and Cdc20p relocate to the substrate address within the APC/C cavity prior to degradation. ..
  46. Chiroli E, Rancati G, Catusi I, Lucchini G, Piatti S. Cdc14 inhibition by the spindle assembly checkpoint prevents unscheduled centrosome separation in budding yeast. Mol Biol Cell. 2009;20:2626-37 pubmed publisher
    ..The target of the SAC is the E3 ubiquitin ligase anaphase-promoting complex (APC) bound to its Cdc20 activator...
  47. Connors B, Rochelle L, Roberts A, Howard G. A Synthetic Interaction between CDC20 and RAD4 in Saccharomyces cerevisiae upon UV Irradiation. Mol Biol Int. 2014;2014:519290 pubmed publisher
    ..We identified a synthetic interaction upon UV exposure between Rad4 and Cdc20, a protein that modulates the activity of the anaphase promoting complex (APC/C), a multisubunit E3 ubiquitin ..
  48. Lim H, Surana U. Cdc20, a beta-transducin homologue, links RAD9-mediated G2/M checkpoint control to mitosis in Saccharomyces cerevisiae. Mol Gen Genet. 1996;253:138-48 pubmed
    ..Here we show that a cdc13 mutant overexpressing Cdc20, a beta-transducin homologue, no longer arrests in G2 at the restrictive temperature but instead undergoes nuclear ..
  49. 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
    ..Finally, SCC1/MCD1/RHC21 was found to interact genetically with CDC20, an activator of APC...
  50. Lu D, Hsiao J, Davey N, Van Voorhis V, Foster S, Tang C, et al. Multiple mechanisms determine the order of APC/C substrate degradation in mitosis. J Cell Biol. 2014;207:23-39 pubmed publisher
    ..We explored these mechanisms using quantitative fluorescence microscopy of GFP-tagged APC/C(Cdc20) substrates in living budding yeast cells...
  51. Hein J, Nilsson J. Stable MCC binding to the APC/C is required for a functional spindle assembly checkpoint. EMBO Rep. 2014;15:264-72 pubmed publisher
    ..SAC) delays progression into anaphase until all chromosomes have aligned on the metaphase plate by inhibiting Cdc20, the mitotic co-activator of the APC/C...
  52. Wang R, Burton J, Solomon M. Transcriptional and post-transcriptional regulation of Cdc20 during the spindle assembly checkpoint in S. cerevisiae. Cell Signal. 2017;33:41-48 pubmed publisher
    ..One of the activators and substrate-binding proteins for the APC is Cdc20. It has been shown previously that Cdc20 can promote its own degradation by the APC in normal cycling cells mainly ..
  53. Girard J, Tenthorey J, Morgan D. An E2 accessory domain increases affinity for the anaphase-promoting complex and ensures E2 competition. J Biol Chem. 2015;290:24614-25 pubmed publisher
    ..The UBA domain is required for normal Ubc1 function and E2 competition in vivo. Thus, the UBA domain of Ubc1 ensures efficient polyubiquitination of substrate by balancing Ubc1 affinity with that of Ubc4. ..
  54. Rossio V, Michimoto T, Sasaki T, Ohbayashi I, Kikuchi Y, Yoshida S. Nuclear PP2A-Cdc55 prevents APC-Cdc20 activation during the spindle assembly checkpoint. J Cell Sci. 2013;126:4396-405 pubmed publisher
    ..Here, we show that an important target of Cdc55 in the SAC is the anaphase-promoting complex (APC) coupled with Cdc20 and that APC-Cdc20 is kept inactive by dephosphorylation by nuclear PP2A-Cdc55 when spindle is damaged...
  55. Yamaguchi Y, Ota K, Ito T. A novel Cdc42-interacting domain of the yeast polarity establishment protein Bem1. Implications for modulation of mating pheromone signaling. J Biol Chem. 2007;282:29-38 pubmed
    ..Intriguingly, Cdc42 can mediate an indirect interaction of the SH3b-CI to the CRIB domain of Ste20. These results suggest that the SH3b and the CI collaborate in tethering of Ste20 to Bem1 to ensure efficient mating pheromone signaling. ..
  56. Chien C, Chen R. Cdc48 chaperone and adaptor Ubx4 distribute the proteasome in the nucleus for anaphase proteolysis. J Biol Chem. 2013;288:37180-91 pubmed publisher
    ..we found that double mutations in Cdc48 and its adaptor Ubx4 cause mitotic arrest with sustained Clb2 and Cdc20 proteins that are normally degraded in anaphase...
  57. Visintin R, Amon A. Regulation of the mitotic exit protein kinases Cdc15 and Dbf2. Mol Biol Cell. 2001;12:2961-74 pubmed
    ..BUB2 negatively regulates all 3 events. Loading of Cdc15 on SPBs depends on TEM1, whereas loading of Dbf2 on SPBs and activation of Dbf2 kinase depend on TEM1 and CDC15. ..
  58. Mariani L, Chiroli E, Nezi L, Muller H, Piatti S, Musacchio A, et al. Role of the Mad2 dimerization interface in the spindle assembly checkpoint independent of kinetochores. Curr Biol. 2012;22:1900-8 pubmed publisher
    ..Surprisingly, downstream of kinetochores the surface does not mediate Mad2 dimerization. Instead, our results are consistent with a role of Mad3 as the main interactor of Mad2 via the dimerization surface. ..
  59. Eshleman H, Morgan D. Sgo1 recruits PP2A to chromosomes to ensure sister chromatid bi-orientation during mitosis. J Cell Sci. 2014;127:4974-83 pubmed publisher
    ..We conclude that in early mitosis, Sgo1 associates transiently with pericentromeric chromatin to promote bi-orientation, in large part by recruiting the Rts1 isoform of PP2A. ..
  60. Sullivan M, Holt L, Morgan D. Cyclin-specific control of ribosomal DNA segregation. Mol Cell Biol. 2008;28:5328-36 pubmed publisher
    ..The coincident destruction of securin, Clb5, and Dbf4 coordinates bulk chromosome segregation with segregation of rDNA. ..
  61. Melloy P, Holloway S. Changes in the localization of the Saccharomyces cerevisiae anaphase-promoting complex upon microtubule depolymerization and spindle checkpoint activation. Genetics. 2004;167:1079-94 pubmed
    ..Localization of Cdc23-GFP was disrupted upon nocodazole treatment in the kinetochore mutant okp1-5 and in the cdc20-1 mutant...
  62. Zachariae W, Shin T, Galova M, Obermaier B, Nasmyth K. Identification of subunits of the anaphase-promoting complex of Saccharomyces cerevisiae. Science. 1996;274:1201-4 pubmed
    ..A small heat-inducible subunit is encoded by the CDC26 gene. The yeast APC is a 36S particle that contains at least seven different proteins. ..