Gene Symbol: KAR9
Description: Kar9p
Alias: Kar9p
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Lee W, Oberle J, Cooper J. The role of the lissencephaly protein Pac1 during nuclear migration in budding yeast. J Cell Biol. 2003;160:355-64 pubmed
    ..Dynein must remain inactive until microtubule ends interact with the bud cortex, at which time dynein and Pac1 appear to be offloaded from the microtubule to the cortex. ..
  2. Korinek W, Copeland M, Chaudhuri A, Chant J. Molecular linkage underlying microtubule orientation toward cortical sites in yeast. Science. 2000;287:2257-9 pubmed
    ..The cortical marker protein Kar9 localizes to the bud tip and is required for the orientation of microtubules toward this region...
  3. Liakopoulos D, Kusch J, Grava S, Vogel J, Barral Y. Asymmetric loading of Kar9 onto spindle poles and microtubules ensures proper spindle alignment. Cell. 2003;112:561-74 pubmed
    ..In budding yeast, the APC-related molecule Kar9 is required for proper alignment of the spindle with the mother-bud axis...
  4. Pereira G, Höfken T, Grindlay J, Manson C, Schiebel E. The Bub2p spindle checkpoint links nuclear migration with mitotic exit. Mol Cell. 2000;6:1-10 pubmed
    ..Since components of the Bub2p checkpoint are conserved in other organisms, we propose that the position of the SPB or mammalian centrosome controls the timing of mitotic exit. ..
  5. Farkasovsky M, Küntzel H. Cortical Num1p interacts with the dynein intermediate chain Pac11p and cytoplasmic microtubules in budding yeast. J Cell Biol. 2001;152:251-62 pubmed
    ..The bud tip location of Num1p depends on Bni1p but does not require Kar9p, Dyn1p, or cMTs, whereas cMT contacts with polar Num1p dots are reduced in cells lacking Dyn1p...
  6. Pereira G, Schiebel E. Kin4 kinase delays mitotic exit in response to spindle alignment defects. Mol Cell. 2005;19:209-21 pubmed
    ..In response to spindle misalignment, Kin4 and Bub2-Bfa1 are brought together at both SPBs. Kin4 now maintains Bub2-Bfa1 activity by counteracting Cdc5, thereby inhibiting mitotic exit. ..
  7. Meednu N, Hoops H, D Silva S, Pogorzala L, Wood S, Farkas D, et al. The spindle positioning protein Kar9p interacts with the sumoylation machinery in Saccharomyces cerevisiae. Genetics. 2008;180:2033-55 pubmed publisher
    ..By two-hybrid analysis, Kar9p and Bim1p interact with the yeast SUMO Smt3p, the E2 enzyme Ubc9p, an E3 Nfi1p, as well as Wss1p, a weak ..
  8. Yin H, Pruyne D, Huffaker T, Bretscher A. Myosin V orientates the mitotic spindle in yeast. Nature. 2000;406:1013-5 pubmed
    ..bud, orientates the spindle early in the cell cycle by binding and polarizing the microtubule-associated protein Kar9 (refs 7-9)...
  9. König C, Maekawa H, Schiebel E. Mutual regulation of cyclin-dependent kinase and the mitotic exit network. J Cell Biol. 2010;188:351-68 pubmed publisher
    ..Consistent with this model, only triple mutants that lack BUB2 and the Cdk1 phosphorylation sites in Mob1 and Cdc15 show mitotic exit defects. ..

More Information


  1. Miller R, Heller K, Frisén L, Wallack D, Loayza D, Gammie A, et al. The kinesin-related proteins, Kip2p and Kip3p, function differently in nuclear migration in yeast. Mol Biol Cell. 1998;9:2051-68 pubmed
    ..Furthermore, we propose that Kip3p functions, in part, by depolymerizing microtubules and is required for the Kar9p-dependent orientation of the cytoplasmic microtubules.
  2. Moore J, Miller R. The cyclin-dependent kinase Cdc28p regulates multiple aspects of Kar9p function in yeast. Mol Biol Cell. 2007;18:1187-202 pubmed
    During mitosis in the yeast Saccharomyces cerevisiae, Kar9p directs one spindle pole body (SPB) toward the incipient daughter cell by linking the associated set of cytoplasmic microtubules (cMTs) to the polarized actin network on the bud ..
  3. Hwang E, Kusch J, Barral Y, Huffaker T. Spindle orientation in Saccharomyces cerevisiae depends on the transport of microtubule ends along polarized actin cables. J Cell Biol. 2003;161:483-8 pubmed
    ..In addition, this process requires the type V myosin protein Myo2, the microtubule end-binding protein Bim1, and Kar9. Here, we show that fusing Bim1 to the tail of the Myo2 is sufficient to orient spindles in the absence of Kar9, ..
  4. Cuschieri L, Miller R, Vogel J. Gamma-tubulin is required for proper recruitment and assembly of Kar9-Bim1 complexes in budding yeast. Mol Biol Cell. 2006;17:4420-34 pubmed
    ..Paradoxically, +TIPs such as the EB1 orthologue Bim1 and Kar9 also associate with spindle pole bodies (SPBs), the centrosome equivalent in budding yeast...
  5. Segal M, Bloom K, Reed S. Bud6 directs sequential microtubule interactions with the bud tip and bud neck during spindle morphogenesis in Saccharomyces cerevisiae. Mol Biol Cell. 2000;11:3689-702 pubmed
    ..These data support a model whereby Bud6 sequentially cues microtubule capture events at the bud tip followed by capture events at the bud neck, necessary for correct spindle morphogenesis and polarity. ..
  6. Miller R, Cheng S, Rose M. Bim1p/Yeb1p mediates the Kar9p-dependent cortical attachment of cytoplasmic microtubules. Mol Biol Cell. 2000;11:2949-59 pubmed
    ..In this process, cortical Kar9p in the bud acts as a link between the actin and microtubule cytoskeletons...
  7. Lee L, Tirnauer J, Li J, Schuyler S, Liu J, Pellman D. Positioning of the mitotic spindle by a cortical-microtubule capture mechanism. Science. 2000;287:2260-2 pubmed
    ..Bim1p, located at microtubule distal ends, bound Kar9p, a protein associated with the daughter cell cortex...
  8. Huisman S, Bales O, Bertrand M, Smeets M, Reed S, Segal M. Differential contribution of Bud6p and Kar9p to microtubule capture and spindle orientation in S. cerevisiae. J Cell Biol. 2004;167:231-44 pubmed
    ..Bud6p function has been linked to Kar9p, a protein guiding MTs along actin cables...
  9. Miller R, Rose M. Kar9p is a novel cortical protein required for cytoplasmic microtubule orientation in yeast. J Cell Biol. 1998;140:377-90 pubmed
    ..A functional GFP-Kar9p fusion protein localized to a single dot at the tip of the shmoo projection...
  10. Maekawa H, Usui T, Knop M, Schiebel E. Yeast Cdk1 translocates to the plus end of cytoplasmic microtubules to regulate bud cortex interactions. EMBO J. 2003;22:438-49 pubmed
    ..b>Kar9, in complex with the EB1-related protein Bim1, mediates contacts of CMs with the cortex of the daughter cell, the ..
  11. Delgehyr N, Lopes C, Moir C, Huisman S, Segal M. Dissecting the involvement of formins in Bud6p-mediated cortical capture of microtubules in S. cerevisiae. J Cell Sci. 2008;121:3803-14 pubmed publisher
    ..A relationship with the second formin, Bnr1p, is unclear. Another player is Kar9p, a protein that guides microtubules along actin cables organised by formins...
  12. Leisner C, Kammerer D, Denoth A, Britschi M, Barral Y, Liakopoulos D. Regulation of mitotic spindle asymmetry by SUMO and the spindle-assembly checkpoint in yeast. Curr Biol. 2008;18:1249-55 pubmed publisher
    ..In budding yeast, the spindle-orientation proteins Kar9 and dynein decorate only one aster of the metaphase spindle and direct it toward the bud [2, 3]...
  13. Stuchell Brereton M, Siglin A, Li J, Moore J, Ahmed S, Williams J, et al. Functional interaction between dynein light chain and intermediate chain is required for mitotic spindle positioning. Mol Biol Cell. 2011;22:2690-701 pubmed publisher
    ..In addition, these data provide the first direct evidence that LC occupancy in the dynein motor complex is important for function. ..
  14. Kammerer D, Stevermann L, Liakopoulos D. Ubiquitylation regulates interactions of astral microtubules with the cleavage apparatus. Curr Biol. 2010;20:1233-43 pubmed publisher
    ..In budding yeast, the Adenomatous Polyposis Coli-related protein Kar9, yeast EB1, and Myo2, a type V myosin, mediate positioning of the mitotic spindle close to the septin-anchored ..
  15. Hayakawa A, Babour A, Sengmanivong L, Dargemont C. Ubiquitylation of the nuclear pore complex controls nuclear migration during mitosis in S. cerevisiae. J Cell Biol. 2012;196:19-27 pubmed publisher
    ..This led to defects in nuclear segregation at the onset of mitosis. Thus, defining ubiquitylation of the yeast NPC highlights yet-unexplored functions of this essential organelle in cell division. ..
  16. Kumar A, Manatschal C, Rai A, Grigoriev I, Degen M, Jaussi R, et al. Short Linear Sequence Motif LxxPTPh Targets Diverse Proteins to Growing Microtubule Ends. Structure. 2017;25:924-932.e4 pubmed publisher
    ..They further provide a unique basis to discover novel +TIPs, and to dissect the role of key interaction nodes and their differential regulation for hierarchical +TIP network organization and function in eukaryotic organisms. ..
  17. Bertazzi D, Kurtulmus B, Pereira G. The cortical protein Lte1 promotes mitotic exit by inhibiting the spindle position checkpoint kinase Kin4. J Cell Biol. 2011;193:1033-48 pubmed publisher
    ..This Lte1-mediated exclusion of Kin4 from the dSPB is essential for proper mitotic exit of cells with a correctly aligned spindle. Therefore, Lte1 promotes mitotic exit by inhibiting Kin4 activity at the dSPB. ..
  18. Hoepfner D, Schaerer F, Brachat A, Wach A, Philippsen P. Reorientation of mispositioned spindles in short astral microtubule mutant spc72Delta is dependent on spindle pole body outer plaque and Kar3 motor protein. Mol Biol Cell. 2002;13:1366-80 pubmed
    ..SPB outer plaque and the astral microtubule motor function of Kar3 but not Kip2/Kip3/Dhc1, or the cortex components Kar9/Num1...
  19. Cepeda García C, Delgehyr N, Juanes Ortiz M, ten Hoopen R, Zhiteneva A, Segal M. Actin-mediated delivery of astral microtubules instructs Kar9p asymmetric loading to the bud-ward spindle pole. Mol Biol Cell. 2010;21:2685-95 pubmed publisher
    In Saccharomyces cerevisiae, Kar9p, one player in spindle alignment, guides the bud-ward spindle pole by linking astral microtubule plus ends to Myo2p-based transport along actin cables generated by the formins Bni1p and Bnr1p and the ..
  20. Eves P, Jin Y, Brunner M, Weisman L. Overlap of cargo binding sites on myosin V coordinates the inheritance of diverse cargoes. J Cell Biol. 2012;198:69-85 pubmed publisher
    ..Vac17 and Mmr1 overlap at one site, whereas Ypt11 and Kar9 bind subsets of residues that also bind Ypt31/Ypt32, Sec4, and Inp2...
  21. Juanes M, Twyman H, Tunnacliffe E, Guo Z, ten Hoopen R, Segal M. Spindle pole body history intrinsically links pole identity with asymmetric fate in budding yeast. Curr Biol. 2013;23:1310-9 pubmed publisher
    ..Temporally symmetric aMT organization perturbed Kar9 polarization by randomizing the choice of the pole to be guided toward the bud...
  22. Moore J, D Silva S, Miller R. The CLIP-170 homologue Bik1p promotes the phosphorylation and asymmetric localization of Kar9p. Mol Biol Cell. 2006;17:178-91 pubmed
    ..cerevisiae is coordinated with the asymmetry of the two poles and requires the microtubule-to-actin linker Kar9p. The asymmetric localization of Kar9p to one spindle pole body (SPB) and microtubule (MT) plus ends requires ..
  23. Pashkova N, Catlett N, Novak J, Weisman L. A point mutation in the cargo-binding domain of myosin V affects its interaction with multiple cargoes. Eukaryot Cell. 2005;4:787-98 pubmed
    ..This large proportion of global suppressors strongly suggests that myo2-2(G1248) causes a conformational change in subdomain I that simultaneously affects multiple cargoes. ..
  24. Haarer B, Helfant A, Nelson S, Cooper J, Amberg D. Stable preanaphase spindle positioning requires Bud6p and an apparent interaction between the spindle pole bodies and the neck. Eukaryot Cell. 2007;6:797-807 pubmed
    ..Regulators of this process include Bud6p (also referred to as the actin-interacting protein Aip3p) and Kar9p, which function to promote contacts between cytoplasmic microtubule ends and actin-delimited cortical attachment ..
  25. Hotz M, Leisner C, Chen D, Manatschal C, Wegleiter T, Ouellet J, et al. Spindle pole bodies exploit the mitotic exit network in metaphase to drive their age-dependent segregation. Cell. 2012;148:958-72 pubmed publisher
    ..During metaphase, the spindle positioning protein Kar9 accumulates asymmetrically, localizing specifically to astral microtubules emanating from the old spindle pole body ..
  26. Tang X, Germain B, Lee W. A novel patch assembly domain in Num1 mediates dynein anchoring at the cortex during spindle positioning. J Cell Biol. 2012;196:743-56 pubmed publisher
    ..We also show that the PA domain is an elongated dimer and discuss the mechanism by which it drives patch assembly. ..
  27. Li J, Lee W, Cooper J. NudEL targets dynein to microtubule ends through LIS1. Nat Cell Biol. 2005;7:686-90 pubmed
    ..We propose a model in which Ndl1 binds transiently to the plus end to promote targeting of LIS1, which cooperatively recruits dynein. ..
  28. 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 show that the yeast spindle positioning protein Kar9 localizes not only to astral but also to kinetochore microtubules, where it becomes targeted for proteasomal ..
  29. Hoffman Sommer M, Kucharczyk R, Piekarska I, Kozlowska E, Rytka J. Mutations in the Saccharomyces cerevisiae vacuolar fusion proteins Ccz1, Mon1 and Ypt7 cause defects in cell cycle progression in a num1Delta background. Eur J Cell Biol. 2009;88:639-52 pubmed publisher
    ..strain can be partially rescued by overproduction of either the calcium pump Pmc1 or the nuclear-migration factors Kar9 and Bim1...
  30. Schweiggert J, Panigada D, Tan A, Liakopoulos D. Kar9 controls the nucleocytoplasmic distribution of yeast EB1. Cell Cycle. 2016;15:2860-2866 pubmed
    ..Here, we show that changes in the amounts of the spindle positioning factor Kar9 specifically affect the distribution of yeast EB1 on spindle microtubules, without influencing other microtubule-..
  31. Drechsler H, Tan A, Liakopoulos D. Yeast GSK-3 kinase regulates astral microtubule function through phosphorylation of the microtubule-stabilizing kinesin Kip2. J Cell Sci. 2015;128:3910-21 pubmed publisher
    ..proteins, such as the yeast CLIP-170 homologue Bik1, dynein and the adenomatous-polyposis-coli-related protein Kar9 to the plus ends of astral MTs...
  32. Scarfone I, Venturetti M, Hotz M, Lengefeld J, Barral Y, Piatti S. Asymmetry of the budding yeast Tem1 GTPase at spindle poles is required for spindle positioning but not for mitotic exit. PLoS Genet. 2015;11:e1004938 pubmed publisher
    ..proteins leading to symmetrical localization of Bub2/Bfa1/Tem1 lead to increased symmetry at spindle poles of the Kar9 protein that mediates spindle positioning and cause spindle misalignment...
  33. Hüls D, Storchova Z, Niessing D. Post-translational modifications regulate assembly of early spindle orientation complex in yeast. J Biol Chem. 2012;287:16238-45 pubmed publisher
    ..Early spindle orientation is driven by the actin-bound myosin Myo2p, which interacts with the adapter Kar9p. The latter also binds to microtubule-associated Bim1p, thereby connecting both types of cytoskeleton...
  34. Daum J, Gomez Ospina N, Winey M, Burke D. The spindle checkpoint of Saccharomyces cerevisiae responds to separable microtubule-dependent events. Curr Biol. 2000;10:1375-8 pubmed
    ..We localized a Bub2p-GFP fusion protein to the cytoplasmic side of the spindle pole body and used a kar9 mutant to show that cells with misoriented spindles are arrested in anaphase of mitosis...
  35. Caydasi A, Micoogullari Y, Kurtulmus B, Palani S, Pereira G. The 14-3-3 protein Bmh1 functions in the spindle position checkpoint by breaking Bfa1 asymmetry at yeast centrosomes. Mol Biol Cell. 2014;25:2143-51 pubmed publisher
    ..Consistently, BMH1-null cells are SPOC deficient. Our work thus identifies Bmh1 as a new SPOC component and refines the molecular mechanism that breaks Bfa1 centrosome asymmetry upon SPOC activation. ..
  36. Hüls D, Niessing D. Purification, crystallization and preliminary crystallographic analysis of Kar9p from Saccharomyces cerevisiae. Acta Crystallogr Sect F Struct Biol Cryst Commun. 2012;68:1251-4 pubmed publisher
    b>Kar9p is required for correct positioning of the mitotic spindle in Saccharomyces cerevisiae. The in vivo function of Kar9p is well understood, but no structural information is available...
  37. Caudron F, Denarier E, Thibout Quintana J, Brocard J, Andrieux A, Fourest Lieuvin A. Mutation of Ser172 in yeast ? tubulin induces defects in microtubule dynamics and cell division. PLoS ONE. 2010;5:e13553 pubmed publisher
    ..When cytoplasmic MT effectors Dyn1p or Kar9p were deleted in S172A and S172E mutants, cells were viable but presented increased ploidy...