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| | SPC34 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; also localized to nuclear side of spindle pole body. (295 aa) |
| | CLB2 | G2/mitotic-specific cyclin-2; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the transition from G2 to M phase; accumulates during G2 and M, then targeted via a destruction box motif for ubiquitin-mediated degradation by the proteasome; CLB2 has a paralog, CLB1, that arose from the whole genome duplication. (491 aa) |
| | MCM16 | Inner kinetochore subunit MCM16; Component of the Ctf19 complex and the COMA subcomplex; involved in kinetochore-microtubule mediated chromosome segregation; binds to centromere DNA; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-H and fission yeast fta3. (181 aa) |
| | ACM1 | APC/C-CDH1 modulator 1; Pseudosubstrate inhibitor of the APC/C; suppresses APC/C [Cdh1]-mediated proteolysis of mitotic cyclins; associates with Cdh1p, Bmh1p and Bmh2p; cell cycle regulated protein; the anaphase-promoting complex/cyclosome is also known as APC/C. (209 aa) |
| | NSL1 | Kinetochore-associated protein NSL1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; required for accurate chromosome segregation; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (216 aa) |
| | KIP2 | Kinesin-like protein KIP2; Kinesin-related motor protein involved in mitotic spindle positioning; stabilizes microtubules by targeting Bik1p to the plus end; functions as a microtubule polymerase and catastrophe inhibitor in vitro; Kip2p levels are controlled during the cell cycle. (706 aa) |
| | CTF19 | Inner kinetochore subunit CTF19; Outer kinetochore protein, needed for accurate chromosome segregation; component of kinetochore sub-complex COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) that functions as platform for kinetochore assembly; required for spindle assembly checkpoint; minimizes potentially deleterious centromere-proximal crossovers by preventing meiotic DNA break formation proximal to centromere; homolog of human centromere constitutive-associated network (CCAN) subunit CENP-P and fission yeast fta2. (369 aa) |
| | NUD1 | Protein NUD1; Component of the spindle pole body outer plaque; acts through the mitotic exit network to specify asymmetric spindle pole body inheritance. (851 aa) |
| | PAC1 | Nuclear distribution protein PAC1; Involved in nuclear migration, part of the dynein/dynactin pathway; targets dynein to microtubule tips, which is necessary for sliding of microtubules along bud cortex; serves at interface between dynein's ATPase site and its microtubule binding stalk, causing individual dynein motors to remain attached to microtubules for long periods; synthetic lethal with bni1; homolog of human LIS1, mutations in which cause the severe brain disorder lissencephaly. (494 aa) |
| | CDC31 | Cell division control protein 31; Calcium-binding component of the spindle pole body (SPB) half-bridge; required for SPB duplication in mitosis and meiosis II; homolog of mammalian centrin; binds multiubiquitinated proteins and is involved in proteasomal protein degradation. (161 aa) |
| | APC5 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the platform domain of the APC/C, based on structural analysis; relative distribution to nuclear foci decreases upon DNA replication stress; Belongs to the APC5 family. (685 aa) |
| | SLK19 | Kinetochore-associated protein; required for chromosome segregation and kinetochore clustering; required for normal segregation of chromosomes in meiosis and mitosis; component of the FEAR regulatory network, which promotes Cdc14p release from the nucleolus during anaphase; potential Cdc28p substrate. (821 aa) |
| | RIO1 | Serine/threonine-protein kinase RIO1; Serine kinase involved in cell cycle regulation and rDNA integrity; associated with late pre-40S particles via its conserved C-terminal domain and participates in late 40S biogenesis; association with pre-40S particles regulated by its catalytic ATPase site and likely occurs after the release of Rio2p from these particles; involved in cell cycle progression and processing of the 20S pre-rRNA into mature 18S rRNA; phosphorylates Rpa43p in anaphase to remove Pol I from rDNA. (484 aa) |
| | SGO1 | Shugoshin; Component of the spindle checkpoint; involved in sensing lack of tension on mitotic chromosomes; protects centromeric Rec8p at meiosis I; required for accurate chromosomal segregation at meiosis II and for mitotic chromosome stability; recruits condensin to the pericentric region of chromosomes during meiosis; dissociates from pericentromeres when sister kinetochores are under tension; Belongs to the shugoshin family. (590 aa) |
| | PAP2 | Non-canonical poly(A) polymerase; involved in nuclear RNA degradation as a component of TRAMP; catalyzes polyadenylation of hypomodified tRNAs, and snoRNA and rRNA precursors; required for mRNA surveillance and maintenance of genome integrity, serving as a link between RNA and DNA metabolism; overlapping but non-redundant functions with Trf5p; relocalizes to cytosol in response to hypoxia. (584 aa) |
| | NUF2 | Kinetochore protein NUF2; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (451 aa) |
| | EGT2 | Protein EGT2; Glycosylphosphatidylinositol (GPI)-anchored cell wall endoglucanase; required for proper cell separation after cytokinesis; expression is activated by Swi5p and tightly regulated in a cell cycle-dependent manner. (1041 aa) |
| | TRF5 | Non-canonical poly(A) polymerase; involved in nuclear RNA degradation as a component of the TRAMP complex; catalyzes polyadenylation of hypomodified tRNAs, and snoRNA and rRNA precursors; overlapping but non-redundant functions with Pap2p. (642 aa) |
| | KAR1 | Cell division control protein KAR1; Protein involved in karyogamy and spindle pole body duplication; involved in karyogamy during mating; involved in spindle pole body duplication during mitosis; localizes to the half-bridge of the spindle pole body; interacts with Spc72p during karyogamy; also interacts with Cdc31p; essential gene. (433 aa) |
| | APC1 | Largest subunit of the Anaphase-Promoting Complex/Cyclosome; APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the platform domain of the APC/C, based on structural analysis; localizes to nuclear foci that become diffuse upon DNA replication stress; Belongs to the APC1 family. (1748 aa) |
| | IBD2 | Protein IBD2; Component of the BUB2-dependent spindle checkpoint pathway; interacts with Bfa1p and functions upstream of Bub2p and Bfa1p. (351 aa) |
| | INN1 | Ingression protein 1; Essential protein that associates with contractile actomyosin ring; required for ingression of the plasma membrane into the bud neck during cytokinesis; C2 domain, a membrane targeting module, is required for function; activates chitin synthase activity of Chs2p during cytokinesis. (409 aa) |
| | DMA2 | Ubiquitin-protein ligase (E3); controls septin dynamics and spindle position checkpoint (SPOC) with ligase Dma1p by regulating recruitment of Elm1p to bud neck; regulates levels of eIF2 subunit Gcd11p, as well as abundance, localization, and ubiquitination of Cdk inhibitory kinase Swe1p; ortholog of human RNF8, similar to human Chfr; contains FHA and RING finger domains; DMA2 has a paralog, DMA1, that arose from the whole genome duplication. (522 aa) |
| | DOM34 | Protein that facilitates ribosomal subunit dissociation; Dom34-Hbs1 complex and Rli1p have roles in dissociating inactive ribosomes to facilitate translation restart, particularly ribosomes stalled in 3' UTRs; required for RNA cleavage in no-go decay, but reports conflict on endonuclease activity; Pelota ortholog; protein abundance increases in response to DNA replication stress; DOM34 has a paralog, YCL001W-B, that arose from the whole genome duplication. (386 aa) |
| | NUP53 | FG-nucleoporin component of central core of nuclear pore complex (NPC); also part of the NPC nuclear basket; contributes directly to nucleocytoplasmic transport; involved in regulation of transcription and mitosis; induces membrane tubulation, which may contribute to nuclear pore assembly; NUP53 has a paralog, ASM4, that arose from the whole genome duplication. (475 aa) |
| | SPC24 | Kinetochore protein SPC24; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (213 aa) |
| | PDS5 | Cohesion maintenance factor; involved in sister chromatid condensation and cohesion; colocalizes with cohesin on chromosomes; performs its cohesin maintenance function in pre-anaphase cells by protecting the integrity of the cohesion complex; also required during meiosis; relocalizes to the cytosol in response to hypoxia. (1277 aa) |
| | BUB2 | Mitotic check point protein BUB2; Mitotic exit network regulator; forms GTPase-activating Bfa1p-Bub2p complex that binds Tem1p and spindle pole bodies, blocks cell cycle progression before anaphase in response to spindle and kinetochore damage; Belongs to the BUB2 family. (306 aa) |
| | MIH1 | M-phase inducer phosphatase; Protein tyrosine phosphatase involved in cell cycle control; regulates the phosphorylation state of Cdc28p; homolog of S. pombe cdc25; Belongs to the MPI phosphatase family. (554 aa) |
| | HOF1 | Cytokinesis protein 2; Protein that regulates actin cytoskeleton organization; required for cytokinesis, actin cable organization, and secretory vesicle trafficking; localized to bud neck; phosphorylated by Dbf2p; regulates actomyosin ring dynamics and septin localization; contains an SH3 domain; N terminus controls cell size and levels of actin cables, while C terminus controls actin cable organization via direct regulation of the formin Bnr1p. (669 aa) |
| | CDC5 | Cell cycle serine/threonine-protein kinase CDC5/MSD2; Polo-like kinase; controls targeting and activation of Rho1p at cell division site via Rho1p guanine nucleotide exchange factors; regulates Spc72p; also functions in adaptation to DNA damage during meiosis; regulates the shape of the nucleus and expansion of the nuclear envelope during mitosis; similar to Xenopus Plx1 and S. pombe Plo1p; human homologs PLK1, PLK3 can each complement yeast cdc5 thermosensitive mutants; Belongs to the protein kinase superfamily. Ser/Thr protein kinase family. CDC5/Polo subfamily. (705 aa) |
| | TEM1 | GTP-binding protein of the Ras superfamily; involved in termination of M-phase; controls actomyosin and septin dynamics during cytokinesis. (245 aa) |
| | CTF3 | Inner kinetochore subunit CTF3; Outer kinetochore protein that forms a complex with Mcm16p and Mcm22p; may bind the kinetochore to spindle microtubules; required for the spindle assembly checkpoint; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-I and fission yeast mis6; Belongs to the CENP-I/CTF3 family. (733 aa) |
| | NKP2 | Central kinetochore protein and subunit of the Ctf19 complex; mutants have elevated rates of chromosome loss; orthologous to fission yeast kinetochore protein cnl2. (153 aa) |
| | CDC25 | Cell division control protein 25; Membrane bound guanine nucleotide exchange factor; also known as a GEF or GDP-release factor; indirectly regulates adenylate cyclase through activation of Ras1p and Ras2p by stimulating the exchange of GDP for GTP; required for progression through G1; thermosensitivity of the cdc25-5 mutant is functionally complemented by human RASGRF1 or by a fragment of human SOS1 comprising the CDC25-related catalytic domain. (1589 aa) |
| | YCS4 | Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation, chromatin binding of condensin, tRNA gene clustering at the nucleolus, and silencing at the mating type locus; required for replication slow zone (RSZ) breakage following Mec1p inactivation; Belongs to the CND1 (condensin subunit 1) family. (1176 aa) |
| | NDL1 | Homolog of nuclear distribution factor NudE; NUDEL; interacts with Pac1p and regulates dynein targeting to microtubule plus ends. (189 aa) |
| | CLB4 | G2/mitotic-specific cyclin-4; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the G2/M transition; may be involved in DNA replication and spindle assembly; accumulates during S phase and G2, then targeted for ubiquitin-mediated degradation; CLB4 has a paralog, CLB3, that arose from the whole genome duplication. (460 aa) |
| | CBF5 | H/ACA ribonucleoprotein complex subunit CBF5; Pseudouridine synthase catalytic subunit of box H/ACA snoRNPs; acts on large and small rRNAs, on snRNA U2, and on some mRNAs; mutations in human ortholog dyskerin cause the disorder dyskeratosis congenita; small nucleolar ribonucleoprotein particles are also known as snoRNPs. (483 aa) |
| | APC2 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the catalytic core of the APC/C; has similarity to cullin Cdc53p. (853 aa) |
| | APC9 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition. (265 aa) |
| | SMC4 | Structural maintenance of chromosomes protein 4; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for tRNA gene clustering at the nucleolus; potential Cdc28p substrate; Belongs to the SMC family. SMC4 subfamily. (1418 aa) |
| | SFI1 | Protein SFI1; Centrin (Cdc31p)-binding protein required for SPB duplication; localizes to the half-bridge of the spindle pole body (SPB); required for progression through G(2)-M transition; phosphorylated by Cdc28p-Clb2p and by Cdc5p; dephosphorylated by Cdc14p; has similarity to Xenopus laevis XCAP-C; Belongs to the SFI1 family. (946 aa) |
| | DAD2 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (133 aa) |
| | SDS22 | Regulatory subunit of the type 1 protein phosphatase (PP1) Glc7p; whether it functions as a positive or negative regulator of Glc7p is controversial; involved in the regulation of Glc7p nuclear localization and function. (338 aa) |
| | HSK3 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (69 aa) |
| | MIF2 | Inner kinetochore subunit MIF2; Protein required for structural integrity of elongating spindles; localizes to the kinetochore; interacts with histones H2A, H2B, and H4; phosphorylated by Ipl1p; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-C and fission yeast cnp3; Belongs to the CENP-C/MIF2 family. (549 aa) |
| | ASK1 | Essential subunit of the Dam1 complex (aka DASH complex); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; phosphorylated during the cell cycle by cyclin-dependent kinases; sumoylated in an Mms21p-dependent manner; protein abundance increases in response to DNA replication stress. (292 aa) |
| | CDC16 | Subunit of the anaphase-promoting complex/cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; required for sporulation; relocalizes to the cytosol in response to hypoxia. (840 aa) |
| | MCM22 | Inner kinetochore subunit MCM22; Outer kinetochore protein and component of the Ctf3 subcomplex; binds to centromeric DNA in a Ctf19p-dependent manner; involved in chromosome segregation and minichromosome maintenance; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-K and fission yeast sim4. (239 aa) |
| | NNF1 | Kinetochore-associated protein NNF1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; required for accurate chromosome segregation; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (201 aa) |
| | BFA1 | Mitotic check point protein BFA1; Subunit of a two-component GTPase-activating protein, Bfa1p-Bub2p; contributes to GAP activity, inactivating Tem1 by stimulating GTP hydrolysis following damage or misalignment of the mitotic spindle; functions as a guanine-nucleotide exchange inhibitor (GDI) for Tem1p; involved in multiple cell cycle checkpoint pathways that control mitotic exit; required when telomeres are damaged, but not for all types of chromosomal DNA damage; phosphorylated by the Polo-like kinase Cdc5p; To S.pombe byr4. (574 aa) |
| | TAH11 | Cell division cycle protein CDT1; DNA replication licensing factor; required for pre-replication complex assembly; Belongs to the Cdt1 family. (604 aa) |
| | SWE1 | Mitosis inhibitor protein kinase SWE1; Protein kinase that regulates the G2/M transition; negative regulator of the Cdc28p kinase; morphogenesis checkpoint kinase; positive regulator of sphingolipid biosynthesis via Orm2p; phosphorylates a tyrosine residue in the N-terminus of Hsp90 in a cell-cycle associated manner, thus modulating the ability of Hsp90 to chaperone a selected clientele; localizes to the nucleus and to the daughter side of the mother-bud neck; homolog of S. pombe Wee1p; potential Cdc28p substrate. (819 aa) |
| | NET1 | Nucleolar protein NET1; Core subunit of the RENT complex; involved in nucleolar silencing and telophase exit; stimulates transcription by RNA polymerase I and regulates nucleolar structure; NET1 has a paralog, TOF2, that arose from the whole genome duplication; To yeast YKR010c. (1189 aa) |
| | SMC3 | Structural maintenance of chromosomes protein 3; Subunit of the multiprotein cohesin complex; required for sister chromatid cohesion in mitotic cells; also required, with Rec8p, for cohesion and recombination during meiosis; phylogenetically conserved SMC chromosomal ATPase family member. (1230 aa) |
| | MAD2 | Component of the spindle-assembly checkpoint complex; delays onset of anaphase in cells with defects in mitotic spindle assembly; forms a complex with Mad1p; regulates APC/C activity during prometaphase and metaphase of meiosis I; gene dosage imbalance between MAD1 and MAD2 leads to chromosome instability. (196 aa) |
| | MAD3 | Subunit of spindle-assembly checkpoint complex; involved in delaying anaphase onset in cells with defects in mitotic spindle assembly; pseudosubstrate inhibitor of APC(Cdc20), the anaphase promoting complex involved in securin (Pds1p) turnover; MAD3 has a paralog, BUB1, that arose from the whole genome duplication. (515 aa) |
| | MND2 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); necessary for maintaining sister chromatid cohesion in prophase I of meiosis by inhibiting premature ubiquitination and subsequent degradation of substrates by the APC(Ama1) ubiquitin ligase; Belongs to the APC15 family. (368 aa) |
| | DSN1 | Kinetochore-associated protein DSN1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; Dsn1p phosphorylation promotes interaction between outer and inner kinetochore proteins; kinetochore receptor for monopolin, via interaction with subunit Csm1p; essential for meiotic but not mitotic chromosome segregation; MIND complex consists of Mtw1p, Nnf1p, Nsl1p and Dsn1p; modified by sumoylation; phosphorylated by monopolin subunit Hrr25p. (576 aa) |
| | NDC80 | Component of the kinetochore-associated Ndc80 complex; conserved coiled-coil protein involved in chromosome segregation, spindle checkpoint activity, and kinetochore assembly and clustering; evolutionarily conserved; complex members include Ndc80p, Nuf2p, Scp24p, and Spc25p; modified by sumoylation. (691 aa) |
| | POG1 | Transcriptional activator POG1; Nuclear chromatin-associated protein of unknown function; may have a role in cell cycle regulation; overexpression promotes recovery from pheromone induced arrest and suppresses the stress sensitivity caused by a mutation in the E3 ubiquitin ligase Rsp5p; binds upstream of BAR1 and cell cycle-related genes; phsosphoylated form may be ubiquitinated by Dma2p; potential Cdc28p substrate; SBF regulated. (351 aa) |
| | IRR1 | Subunit of the cohesin complex; which is required for sister chromatid cohesion during mitosis and meiosis and interacts with centromeres and chromosome arms; relocalizes to the cytosol in response to hypoxia; essential for viability; Belongs to the SCC3 family. (1150 aa) |
| | CDC23 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition. (626 aa) |
| | DMA1 | Ubiquitin-protein ligase (E3); controls septin dynamics, spindle position checkpoint (SPOC) with ligase Dma2p by regulating recruitment of Elm1p to bud neck; regulates levels of eIF2 subunit Gcd11p, as well as abundance, localization, and ubiquitination of Cdk inhibitory kinase Swe1p; ubiquitinates cyclin Pcl1p; ortholog of human RNF8, similar to human Chfr; contains FHA, RING fingers; DMA1 has a paralog, DMA2, that arose from the whole genome duplication. (416 aa) |
| | OKP1 | Inner kinetochore subunit OKP1; Outer kinetochore protein required for accurate chromosome segregation; component of COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) a kinetochore sub-complex which functions as a platform for kinetochore assembly; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-Q and fission yeast fta7. (406 aa) |
| | DAM1 | Essential subunit of the Dam1 complex (aka DASH complex); cooperates with Duo1p to connect the DASH complex with the microtubules (MT); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; Ipl1p target for regulating kinetochore-MT attachments. (343 aa) |
| | CLB1 | G2/mitotic-specific cyclin-1; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the transition from G2 to M phase; accumulates during G2 and M, then targeted via a destruction box motif for ubiquitin-mediated degradation by the proteasome; CLB1 has a paralog, CLB2, that arose from the whole genome duplication. (471 aa) |
| | RME1 | Zinc finger protein involved in control of meiosis; prevents meiosis by repressing IME1 expression and promotes mitosis by activating CLN2 expression; directly repressed by a1-alpha2 regulator; mediates cell type control of sporulation; relocalizes from nucleus to cytoplasm upon DNA replication stress. (300 aa) |
| | DOC1 | Anaphase-promoting complex subunit DOC1; Processivity factor; required for the ubiquitination activity of the anaphase promoting complex (APC), mediates the activity of the APC by contributing to substrate recognition; involved in cyclin proteolysis; contains a conserved DOC1 homology domain; Belongs to the APC10 family. (250 aa) |
| | CSE1 | Importin alpha re-exporter; Nuclear envelope protein that acts as a recycling factor; mediates the nuclear export of Srp1p (importin alpha) back to the cytoplasm after its import substrates have been released into the nucleoplasm, thereby allowing the participation of Srp1p in multiple rounds of nuclear import; required for accurate chromosome segregation; homolog of metazoan CAS and human CSE1L, overexpression of which is implicated in cancer progression. (960 aa) |
| | CDC20 | Activator of anaphase-promoting complex/cyclosome (APC/C); APC/C is required for metaphase/anaphase transition; directs ubiquitination of mitotic cyclins, Pds1p, and other anaphase inhibitors; cell-cycle regulated; potential Cdc28p substrate; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the WD repeat CDC20/Fizzy family. (610 aa) |
| | SRM1 | Nucleotide exchange factor for Gsp1p; localizes to the nucleus, required for nucleocytoplasmic trafficking of macromolecules; suppressor of the pheromone response pathway; potentially phosphorylated by Cdc28p; human homolog of the RAN GEF, RCC1, can complement a temperature sensitive point mutant. (482 aa) |
| | MAD1 | Coiled-coil protein involved in spindle-assembly checkpoint; required for inhibition of karyopherin/importin Pse1p (aka Kap121p) upon spindle assembly checkpoint arrest; phosphorylated by Mps1p upon checkpoint activation which leads to inhibition of anaphase promoting complex activity; forms a complex with Mad2p; gene dosage imbalance between MAD1 and MAD2 leads to chromosome instability; Belongs to the MAD1 family. (749 aa) |
| | DUO1 | Essential subunit of the Dam1 complex (aka DASH complex); cooperates with Dam1p to connect the DASH complex with microtubules (MT); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (247 aa) |
| | CDH1 | Activator of anaphase-promoting complex/cyclosome (APC/C); antagonist of the spindle assembly checkpoint; directs ubiquitination of cyclins resulting in mitotic exit; targets the APC/C to specific substrates including: Cdc20p, Ase1p, Cin8p, Fin1p and Clb5p; partially active in metaphase, and fully active in anaphase; cell-cycle regulated; Belongs to the WD repeat CDC20/Fizzy family. (566 aa) |
| | CNN1 | Inner kinetochore subunit CNN1; Kinetochore protein; associated with the essential kinetochore proteins Nnf1p and Spc24p; phosphorylated by Clb5-Cdk1, Mps1p, Ipl1p and to a lesser extent by Clb2-Cdk1; localizes to the lower region of the Ndc80 complex during anaphase and regulates KMN activity by inhibiting the Mtw1 and Spc105 complexes from binding to the Ndc80 complex; similar to metazoan CENP-T; Belongs to the CENP-T/CNN1 family. (361 aa) |
| | CDC26 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; relocalizes to the cytosol in response to hypoxia; Belongs to the CDC26 family. (124 aa) |
| | SMC2 | Structural maintenance of chromosomes protein 2; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; essential SMC chromosomal ATPase family member that forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for clustering of tRNA genes at the nucleolus. (1170 aa) |
| | CDC14 | Tyrosine-protein phosphatase CDC14; Protein phosphatase required for mitotic exit; required for rDNA segregation, cytokinesis, meiosis I spindle disassembly, environmental stress response; held in nucleolus by Cdc55p in early meiosis, liberated by FEAR and Mitotic Exit Network in anaphase, enabling it to effect a decrease in CDK/B-cyclin activity and mitotic exit; sequestered in metaphase II, released upon entry into anaphase II; human homolog CDC14A can complement thermosensitivity of yeast cdc14-1 mutant. (551 aa) |
| | MOB2 | Activator of Cbk1p kinase; component of the RAM signaling network that regulates cellular polarity and morphogenesis; activation of Cbk1p facilitates the Ace2p-dependent daughter cell-specific transcription of genes involved in cell separation; similar to Mob1p. (287 aa) |
| | CDC4 | Cell division control protein 4; F-box protein required for both the G1/S and G2/M phase transitions; modular substrate specificity factor which associates with core SCF (Cdc53p, Skp1p and Hrt1p/Rbx1p) to form the SCFCdc4 complex; SCFCdc4 acts as a ubiquitin-protein ligase directing ubiquitination of cyclin-dependent kinase (CDK) phosphorylated substrates, such as: Sic1p, Far1p, Cdc6p, Clb6p, and Cln3p. (779 aa) |
| | SMC1 | Structural maintenance of chromosomes protein 1; Subunit of the multiprotein cohesin complex; essential protein involved in chromosome segregation and in double-strand DNA break repair; SMC chromosomal ATPase family member, binds DNA with a preference for DNA with secondary structure; Belongs to the SMC family. SMC1 subfamily. (1225 aa) |
| | SCC4 | MAU2 chromatid cohesion factor homolog; Subunit of cohesin loading factor (Scc2p-Scc4p); complex is required for the loading of cohesin complexes onto chromosomes; involved in establishing sister chromatid cohesion during double-strand break repair via phosphorylated histone H2AX. (624 aa) |
| | GLC7 | Serine/threonine-protein phosphatase PP1-2; Type 1 S/T protein phosphatase (PP1) catalytic subunit; involved in glycogen metabolism, sporulation and mitotic progression; interacts with multiple regulatory subunits; regulates actomyosin ring formation; subunit of CPF; recruited to mating projections by Afr1p interaction; regulates nucleocytoplasmic shuttling of Hxk2p; import into the nucleus is inhibited during spindle assembly checkpoint arrest; involved in dephosphorylating Rps6a/b and Bnr1p. (312 aa) |
| | SPC25 | Kinetochore protein SPC25; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (221 aa) |
| | BIM1 | Protein BIM1; Microtubule plus end-tracking protein; together with Kar9p makes up the cortical microtubule capture site and delays the exit from mitosis when the spindle is oriented abnormally; homolog of human end binding protein 1 (EB1); Belongs to the MAPRE family. (344 aa) |
| | CIN8 | Kinesin-like protein CIN8; Kinesin motor protein; involved in mitotic spindle assembly and chromosome segregation. (1000 aa) |
| | NKP1 | Central kinetochore protein and subunit of the Ctf19 complex; mutants have elevated rates of chromosome loss; orthologous to fission yeast kinetochore protein fta4; Belongs to the NKP1 family. (238 aa) |
| | YCG1 | Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation and chromatin binding by the complex; required for tRNA genes clustering at the nucleolus; required for replication slow zone breakage following Mec1p inactivation; transcription is cell cycle regulated, peaking in mitosis and declining in G1; protein is constitutively degraded by the proteasome; rate limiting for condensin recruitment to chromatin. (1035 aa) |
| | DAD4 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (72 aa) |
| | MCM21 | Inner kinetochore subunit MCM21; Component of the kinetochore sub-complex COMA; COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) bridges kinetochore subunits in contact with centromeric DNA with subunits bound to microtubules during kinetochore assembly; involved in minichromosome maintenance; modified by sumoylation; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-O and fission yeast mal2; Belongs to the CENP-O/MCM21 family. (368 aa) |
| | SSD1 | Protein SSD1; Translational repressor with a role in polar growth and wall integrity; regulated by Cbk1p phosphorylation to effect bud-specific translational control and localization of specific mRNAs; interacts with TOR pathway components; contains a functional N-terminal nuclear localization sequence and nucleocytoplasmic shuttling appears to be critical to Ssd1p function; Belongs to the RNR ribonuclease family. (1250 aa) |
| | SWM1 | Subunit of the anaphase-promoting complex (APC); APC is an E3 ubiquitin ligase that regulates the metaphase-anaphase transition and exit from mitosis; required for activation of the daughter-specific gene expression and spore wall maturation; Belongs to the APC13 family. (170 aa) |
| | CHL4 | Inner kinetochore subunit CHL4; Outer kinetochore protein required for chromosome stability; involved in new kinetochore assembly and sister chromatid cohesion; forms a stable complex with Iml3p; peripheral component of the Ctf19 kinetochore complex that interacts with Ctf19p, Ctf3p, and Mif2p; required for the spindle assembly checkpoint; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-N and fission yeast mis15; Belongs to the CENP-N/CHL4 family. (458 aa) |
| | SPC19 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; also localized to nuclear side of spindle pole body. (165 aa) |
| | RGP1 | Subunit of a Golgi membrane exchange factor (Ric1p-Rgp1p); this complex catalyzes nucleotide exchange on Ypt6p; Belongs to the RGP1 family. (663 aa) |
| | APC4 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the platform domain of the APC/C, based on structural analysis; relative distribution to the nucleus increases upon DNA replication stress. (652 aa) |
| | PDS1 | Securin; inhibits anaphase by binding separin Esp1p; blocks cyclin destruction and mitotic exit, essential for meiotic progression and mitotic cell cycle arrest; localization is cell-cycle dependent and regulated by Cdc28p phosphorylation. (373 aa) |
| | DBF4 | Regulatory subunit of Cdc7p-Dbf4p kinase complex; required for Cdc7p kinase activity and initiation of DNA replication; phosphorylates the Mcm2-7 family of proteins; cell cycle regulated; relative distribution to the nucleus increases upon DNA replication stress; co-expression of human CDC7 and DBF4 complements single cdc7 or dbf4 null mutations or the cdc7 dbf4 double null mutation. (704 aa) |
| | DAD1 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (94 aa) |
| | CLB3 | G2/mitotic-specific cyclin-3; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the G2/M transition; may be involved in DNA replication and spindle assembly; accumulates during S phase and G2, then targeted for ubiquitin-mediated degradation; relative distribution to the nucleus increases upon DNA replication stress; CLB3 has a paralog, CLB4, that arose from the whole genome duplication. (427 aa) |
| | ASM4 | FG-nucleoporin component of central core of nuclear pore complex (NPC); contributes directly to nucleocytoplasmic transport; induces membrane tubulation, which may contribute to nuclear pore assembly; ASM4 has a paralog, NUP53, that arose from the whole genome duplication. (528 aa) |
| | UBC9 | SUMO-conjugating enzyme involved in the Smt3p conjugation pathway; nuclear protein required for S- and M-phase cyclin degradation and mitotic control; involved in proteolysis mediated by the anaphase-promoting complex cyclosome (APCC). (157 aa) |
| | SIT4 | Serine/threonine-protein phosphatase PP1-1; Ceramide-activated, type 2A-related serine-threonine phosphatase; functions in G1/S transition of mitotic cycle; controls lifespan, mitochondrial function, cell cycle progression by regulating HXK2 phosphorylation; regulator of COPII coat dephosphorylation; required for ER to Golgi traffic; interacts with Hrr25p kinase; cytoplasmic and nuclear protein that modulates functions mediated by Pkc1p including cell wall and actin cytoskeleton organization; similar to human PP6. (311 aa) |
| | CDC7 | Cell division control protein 7; DDK (Dbf4-dependent kinase) catalytic subunit; required for origin firing and replication fork progression in mitotic S phase through phosphorylation of Mcm2-7p complexes and Cdc45p; kinase activity correlates with cyclical DBF4 expression; required for pre-meiotic DNA replication, meiotic DSB formation, recruitment of monopolin complex to kinetochores during meiosis I, regulation of meiosis-specific Ndt80p; mutation complemented by human CDC7 and DBF4 co-expression. (507 aa) |
| | APC11 | Catalytic core subunit, Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; contains a RING-H2 domain that is required for activity. (165 aa) |
| | MCD1 | Sister chromatid cohesion protein 1; Essential alpha-kleisin subunit of the cohesin complex; required for sister chromatid cohesion in mitosis and meiosis; apoptosis induces cleavage and translocation of a C-terminal fragment to mitochondria; expression peaks in S phase. (566 aa) |
| | BIK1 | Nuclear fusion protein BIK1; Microtubule-associated protein; component of the interface between microtubules and kinetochore, involved in sister chromatid separation; essential in polyploid cells but not in haploid or diploid cells; ortholog of mammalian CLIP-170. (440 aa) |
| | DAD3 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (94 aa) |
| | AME1 | Inner kinetochore subunit AME1; Essential kinetochore protein associated with microtubules and SPBs; component of the kinetochore sub-complex COMA (Ctf19p, Okp1p, Mcm21p, Ame1p); involved in spindle checkpoint maintenance; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-U and fission yeast Mis17; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the CENP-U/AME1 family. (324 aa) |
| | CDC28 | Cyclin-dependent kinase (CDK) catalytic subunit; master regulator of mitotic and meiotic cell cycles; alternately associates with G1, S, G2/M phase cyclins, which provide substrate specificity; regulates metabolism, basal transcription, chromosome dynamics, growth and morphogenesis; transcript induction in osmostress involves antisense RNA; human homologs CDK1, CDK2, CDK3 can complement yeast conditional cdc28 mutants; human CDK1, CDK2 can complement yeast cdc28 null mutant. (298 aa) |
| | AMN1 | Antagonist of mitotic exit network protein 1; Protein required for daughter cell separation; multiple mitotic checkpoints, and chromosome stability; contains 12 degenerate leucine-rich repeat motifs; expression is induced by the Mitotic Exit Network (MEN); Belongs to the AMN1 family. (549 aa) |
| | HSL7 | Protein arginine N-methyltransferase; exhibits septin and Hsl1p-dependent localization to the bud neck in budded cells and periodic Hsl1p-dependent phosphorylation; required with Hsl1p, and Elm1p for the mother-bud neck recruitment, phosphorylation, and degradation of Swe1p; interacts directly with Swe1p; relocalizes away from bud neck upon DNA replication stress; human homolog PRMT5 can complement yeast hsl7 mutant. (827 aa) |
| | IML3 | Inner kinetochore subunit IML3; Outer kinetochore protein and component of the Ctf19 complex; involved in the establishment of pericentromeric cohesion during mitosis; prevents non-disjunction of sister chromatids during meiosis II; forms a stable complex with Chl4p; required for localization of Sgo1p to pericentric sites during meiosis I; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-L and fission yeast fta1; Belongs to the CENP-L/IML3 family. (245 aa) |
| | BRN1 | Subunit of the condensin complex; required for chromosome condensation and for clustering of tRNA genes at the nucleolus; may influence multiple aspects of chromosome transmission. (754 aa) |
| | CDC27 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition. (758 aa) |
| | KIP1 | Kinesin-like protein KIP1; Kinesin-related motor protein; required for mitotic spindle assembly, chromosome segregation, and 2 micron plasmid partitioning; functionally redundant with Cin8p for chromosomal but not plasmid functions; Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family. BimC subfamily. (1111 aa) |
| | STU1 | Protein STU1; Component of the mitotic spindle; binds to interpolar microtubules via its association with beta-tubulin (Tub2p); required for interpolar microtubules to provide an outward force on the spindle poles; Belongs to the CLASP family. (1513 aa) |
| | SHE1 | Mitotic spindle protein; interacts with components of the Dam1 (DASH) complex, its effector Sli15p, and microtubule-associated protein Bim1p; also localizes to nuclear microtubules and to the bud neck in a ring-shaped structure; inhibits dynein function. (338 aa) |
| | FUS3 | Mitogen-activated serine/threonine protein kinase involved in mating; phosphoactivated by Ste7p; substrates include Ste12p, Far1p, Bni1p, Sst2p; inhibits invasive growth during mating by phosphorylating Tec1p, promoting its; inhibits recruitment of Ste5p, Cdc42p-mediated asymmetry and mating morphogenesis. (353 aa) |
| | CDC15 | Cell division control protein 15; Protein kinase of the Mitotic Exit Network; localized to the spindle pole bodies at late anaphase; promotes mitotic exit by directly switching on the kinase activity of Dbf2p; required for spindle disassembly after meiosis II; relocalizes to the cytoplasm upon DNA replication stress. (974 aa) |
| | MTW1 | Kinetochore-associated protein MTW1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; critical to kinetochore assembly; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (289 aa) |
| | LTE1 | Guanine nucleotide exchange factor LTE1; Protein similar to GDP/GTP exchange factors; without detectable GEF activity; required for asymmetric localization of Bfa1p at daughter-directed spindle pole bodies and for mitotic exit at low temperatures. (1435 aa) |
