CDC28

UniProt ID: P00546
Organism: Saccharomyces cerevisiae
Review Status: COMPLETE
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Gene Description

CDC28 encodes Cdk1, the single essential cyclin-dependent serine/threonine protein kinase of the budding yeast Saccharomyces cerevisiae and the catalytic engine of its cell division cycle. The 298-residue protein is catalytically inert on its own and becomes active only in a holoenzyme with one of nine cyclins (the G1 cyclins Cln1, Cln2 and Cln3, and the B-type cyclins Clb1-6) together with the small phospho-adaptor subunit Cks1; activation also requires phosphorylation of Thr169 in the T-loop by the CDK-activating kinase Cak1. Its activity is restrained by Swe1-mediated phosphorylation of Tyr19 (reversed by the Mih1 phosphatase), which enforces the morphogenesis checkpoint, and by the stoichiometric inhibitors Sic1 (B-type cyclin complexes) and Far1 (G1 cyclin complexes during mating-pheromone arrest). Cyclin identity retargets the kinase through docking motifs (LP motifs for Cln1/2, RxL and NLxxxL for Clb5/6, LxF for Clb2) so that the same catalytic subunit phosphorylates stage-specific sets of S/T-P substrates, with Cks1 enabling processive multisite phosphorylation. Cln3-Cdc28 and then Cln1/2-Cdc28 execute Start, the G1/S commitment, by hyperphosphorylating the SBF inhibitor Whi5 and the CKI Sic1 (freeing Clb5/6-Cdc28) and by silencing pheromone signalling through Ste5 phosphorylation; Clb5/6-Cdc28 triggers replication-origin firing and blocks re-licensing; Clb3/4- and Clb1/2-Cdc28 drive spindle pole body separation, spindle assembly, chromosome segregation and mitotic entry, and keep APC/C-Cdh1 inhibited until cyclin destruction, Cdc14-mediated dephosphorylation and Sic1 re-accumulation reset the cell to G1. Through hundreds of substrates it also couples the cycle to bud emergence and polarized growth, septin and actomyosin-ring assembly, membrane trafficking, histone-gene and basal transcription, telomerase recruitment, cohesion establishment, DNA double-strand-break resection and repair-pathway choice, and carbon-store mobilisation, and it drives premeiotic DNA replication, meiotic recombination initiation and the meiotic divisions. Cdc28 is distributed between the cytoplasm and nucleus in a cell-cycle-regulated manner, with additional pools at spindle pole bodies, cytoplasmic-microtubule plus ends and the bud neck.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0000082 G1/S transition of mitotic cell cycle
IBA
GO_REF:0000033
ACCEPT
Summary: Cdc28 is the budding-yeast Cdk1; with the G1 cyclins Cln1-3 it executes Start, the commitment step of the G1/S transition, by phosphorylating Whi5 and the CKI Sic1 and by triggering the SBF/MBF transcriptional program. The PAINT node is seeded by pombe cdc2, C. albicans CDC28 and Cdc28 itself, which carries direct experimental support.
Reason: Core cell-cycle function of the single essential CDK; the target's own IDA (PMID:2142620) and the classical Start phenotype of cdc28-ts alleles (PMID:7002718) ground the IBD, and inheritance of G1/S control across the Cdk1 clade is phylogenetically sound.
Supporting Evidence:
PMID:2142620
Cln proteins are an essential component of the active protein kinase complex required for the G1 to S transition
PMID:7002718
Each mutation produces stage-specific arrest of cell division at start, the same point where mating pheromone interrupts division.
GO:0000082 G1/S transition of mitotic cell cycle
IDA
PMID:2142620
G1-specific cyclins of S. cerevisiae: cell cycle periodicity...
ACCEPT
Summary: Wittenberg et al. showed that the G1 cyclin Cln2 associates with p34CDC28 to form an active kinase and that the Cln-Cdc28 complex is the kinase required for the G1-to-S transition.
Reason: Direct biochemical demonstration of the G1 cyclin-Cdc28 kinase that drives Start; core function.
Supporting Evidence:
PMID:2142620
we demonstrate that the Cln2 polypeptide interacts with p34CDC28 to form an active protein kinase complex
PMID:2142620
Cln proteins are an essential component of the active protein kinase complex required for the G1 to S transition
GO:0000082 G1/S transition of mitotic cell cycle
IEA
GO_REF:0000117
ACCEPT
Summary: ARBA machine-learning mapping to G1/S transition, consistent with the experimental and phylogenetic evidence.
Reason: Electronic inference that agrees with direct evidence (PMID:2142620) and the IBA; correct and at the right level.
GO:0000086 G2/M transition of mitotic cell cycle
IBA
GO_REF:0000033
ACCEPT
Summary: Clb-Cdc28 (Clb1-4) drives mitotic entry; its activation by Mih1-mediated Tyr19 dephosphorylation and its inhibition by Swe1 constitute the morphogenesis checkpoint. Node seeded by pombe cdc2 and Cdc28 itself.
Reason: Core function; the G2/M requirement of CDC28 is directly demonstrated (PMID:2165600, PMID:1849457) and the IBD placement at the Cdk1 node is sound.
Supporting Evidence:
PMID:2165600
We present evidence that the Cdc28 protein kinase is also required for mitosis and that this function is executed in the G2 interval of the cell cycle.
GO:0000086 G2/M transition of mitotic cell cycle
IDA
PMID:32083180
Phosphoregulation of Rad51/Rad52 by CDK1 functions as a mole...
ACCEPT
Summary: Lim et al. showed that Cdc28 with G2/M cyclins phosphorylates Rad51 and Rad52 in G2/M to switch on homologous recombination. The direct evidence is for G2/M-phase Clb-Cdc28 kinase activity rather than for the transition itself.
Reason: The term is correct for Cdc28 (core G2/M function); this paper documents a G2/M-cyclin-Cdc28-specific phosphorylation event, so it is a somewhat indirect but not wrong support. The HR-specific content is better captured by the DNA-repair terms.
Supporting Evidence:
PMID:32083180
the budding yeast CDK1, Cdc28, phosphorylates the major homologous recombination regulators Rad51 and Rad52. This phosphorylation occurs in the G2/M phase by Cdc28 in combination with G2/M phase cyclins.
GO:0000086 G2/M transition of mitotic cell cycle
IMP
PMID:1427070
Cyclin-B homologs in Saccharomyces cerevisiae function in S ...
ACCEPT
Summary: Richardson et al. showed with clb1-4 quadruple mutants rescued by conditional CLB1 that the Clb cyclins, i.e. Clb-Cdc28 kinase, perform an essential role at the G2/M transition.
Reason: Core mitotic-entry function; the cyclin requirement is a requirement for the Clb-Cdc28 holoenzyme.
Supporting Evidence:
PMID:1427070
we showed that the CLB genes perform an essential role at the G2/M-phase transition, and also a role in S phase.
GO:0000086 G2/M transition of mitotic cell cycle
IMP
PMID:1849457
The role of CDC28 and cyclins during mitosis in the budding ...
ACCEPT
Summary: Surana et al. characterised cdc28-1N, an allele with normal Start function but a mitotic defect, suppressed by the B-type cyclins CLB1-4, establishing a distinct mitotic role for Cdc28 executed with G2 cyclins.
Reason: Classical genetic demonstration that Cdc28 is required for mitosis in addition to Start; core function.
Supporting Evidence:
PMID:1849457
Our results are consistent with CDC28 function being required in both G1 and mitosis. Its mitotic role, we believe, involves interaction with a family of at least four G2-specific cyclins.
GO:0000086 G2/M transition of mitotic cell cycle
NAS
PMID:1427070
Cyclin-B homologs in Saccharomyces cerevisiae function in S ...
ACCEPT
Summary: Author-statement duplicate of the IMP row from the same paper (Clb-Cdc28 essential at G2/M).
Reason: Same evidence as the IMP row; correct core function.
Supporting Evidence:
PMID:1427070
we showed that the CLB genes perform an essential role at the G2/M-phase transition, and also a role in S phase.
GO:0000086 G2/M transition of mitotic cell cycle
NAS
PMID:1849457
The role of CDC28 and cyclins during mitosis in the budding ...
ACCEPT
Summary: Author-statement duplicate of the IMP row from the same paper (mitotic role of CDC28 with Clb cyclins).
Reason: Same evidence as the IMP row; correct core function.
Supporting Evidence:
PMID:1849457
Our results are consistent with CDC28 function being required in both G1 and mitosis. Its mitotic role, we believe, involves interaction with a family of at least four G2-specific cyclins.
GO:0000122 negative regulation of transcription by RNA polymerase II
IMP
PMID:22056777
Cdk1 promotes kinetochore bi-orientation and regulates Cdc20...
KEEP AS NON CORE
Summary: Liang et al. found that Cdk1 sustains CDC20 expression during recovery from spindle-checkpoint arrest, in part because Cdk1 negatively regulates transcription of the repressor YOX1.
Reason: Real but downstream/indirect transcriptional effect of Cdk1 (through the Yox1 repressor); not a core function of the kinase.
Supporting Evidence:
PMID:22056777
we find that YOX1 transcription is negatively regulated by Cdk1
GO:0000278 mitotic cell cycle
IMP
PMID:22056777
Cdk1 promotes kinetochore bi-orientation and regulates Cdc20...
ACCEPT
Summary: Cdk1 is required for efficient recovery from SAC-induced mitotic arrest, promoting kinetochore bi-orientation and sustaining Cdc20 expression.
Reason: Cdc28 is the master regulator of the mitotic cell cycle; the general term is correct, if less informative than the transition-specific terms.
Supporting Evidence:
PMID:22056777
Here, we report two unrecognized functions of yeast Cdk1 required for efficient recovery from SAC-induced arrest.
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IBA
GO_REF:0000033
ACCEPT
Summary: Cdc28 is the catalytic subunit of the cyclin-dependent kinase holoenzyme: it is inactive alone and is activated by binding one of nine cyclins (Cln1-3, Clb1-6) together with the Cks1 subunit. Deep PAINT node covering the CDK family.
Reason: Core cellular component; cyclin-CDK holoenzyme formation is the defining property of the family and is directly shown for Cdc28.
Supporting Evidence:
PMID:2142620
we demonstrate that the Cln2 polypeptide interacts with p34CDC28 to form an active protein kinase complex
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IDA
PMID:1849458
A cyclin B homolog in S. cerevisiae: chronic activation of t...
ACCEPT
Summary: Ghiara et al. identified the B-type cyclin Scb1 (Clb2); a non-degradable allele arrests cells in mitosis and the arrest is reversed by inactivating Cdc28, showing that the cyclin acts through persistent activation of the Cdc28 kinase.
Reason: Direct evidence for a Clb-Cdc28 holoenzyme whose persistent activity blocks mitotic exit.
Supporting Evidence:
PMID:1849458
This arrest can be reversed by inactivation of the Cdc28 protein kinase, suggesting that cyclin-mediated arrest results from persistent protein kinase activation.
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IDA
PMID:2142620
G1-specific cyclins of S. cerevisiae: cell cycle periodicity...
ACCEPT
Summary: Cln2 co-immunoprecipitates with p34CDC28 in an active kinase complex.
Reason: Direct biochemical evidence for the Cln2-Cdc28 holoenzyme.
Supporting Evidence:
PMID:2142620
we demonstrate that the Cln2 polypeptide interacts with p34CDC28 to form an active protein kinase complex
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IEA
GO_REF:0000117
ACCEPT
Summary: ARBA mapping to the CDK holoenzyme complex.
Reason: Electronic inference concordant with direct evidence.
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IPI
PMID:10688190
A comprehensive analysis of protein-protein interactions in ...
ACCEPT
Summary: Genome-wide two-hybrid screen (Uetz et al.) detected Cdc28 interactions with cyclins and with Cks1/Cak1 consistent with holoenzyme membership.
Reason: High-throughput interaction support for a complex that is independently established biochemically; correct component term.
Supporting Evidence:
PMID:10688190
These approaches resulted in the detection of 957 putative interactions involving 1,004 S. cerevisiae proteins.
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IPI
PMID:10913169
Cks1 is required for G(1) cyclin-cyclin-dependent kinase act...
ACCEPT
Summary: Reynard et al. reconstituted Cln2-Cdc28 and Clb-Cdc28 complexes in insect cells and showed that Cks1 is required for G1 cyclin-Cdc28 kinase activity and stabilises Cln2-Cdc28 complexes.
Reason: Direct reconstitution of the cyclin-Cdc28-Cks1 holoenzyme; core component.
Supporting Evidence:
PMID:10913169
Cks1 can both stabilize Cln2-Cdc28 complexes and activate intact complexes in vitro
PMID:10913169
Cdc28 forms stable, active complexes with the B-type cyclins Clb4 and Clb5 regardless of whether Cks1 is present.
GO:0000307 cyclin-dependent protein kinase holoenzyme complex
IPI
PMID:8319908
CLB5 and CLB6, a new pair of B cyclins involved in DNA repli...
ACCEPT
Summary: Schwob and Nasmyth identified the S-phase cyclins Clb5/Clb6 whose association with Cdc28 confers the S-phase-promoting kinase.
Reason: Cyclin-Cdc28 complex identification; correct component term.
Supporting Evidence:
PMID:8319908
The functions of the Cdc28 protein kinase in DNA replication and mitosis in Saccharomyces cerevisiae are thought to be determined by the type of cyclin subunit with which it is associated.
GO:0000706 meiotic DNA double-strand break processing
IGI
PMID:20150422
Processing of meiotic DNA double strand breaks requires cycl...
KEEP AS NON CORE
Summary: Manfrini et al. showed that Cdk1 phosphorylation of Sae2 Ser-267 is required to initiate resection of Spo11-induced meiotic DSBs by allowing Spo11 removal.
Reason: Cdc28 does the work (phosphorylates Sae2) but this is one of many substrate-mediated downstream processes; non-core relative to the cell-cycle transitions it gates.
Supporting Evidence:
PMID:20150422
phosphorylation of Sae2 Ser-267 by cyclin-dependent kinase 1 (Cdk1) is required to initiate meiotic DSB resection by allowing Spo11 removal from DSB ends
GO:0000729 DNA double-strand break processing
IMP
PMID:21841787
Cell cycle regulation of DNA double-strand break end resecti...
KEEP AS NON CORE
Summary: Chen et al. showed Cdk1-dependent phosphorylation of the resection nuclease Dna2 (Thr4, Ser17, Ser237) stimulates its recruitment to DSBs and 5' strand resection.
Reason: Direct substrate-mediated control of DSB end resection; genuine but non-core.
Supporting Evidence:
PMID:21841787
we provide evidence for Cdk1-dependent phosphorylation of the resection nuclease Dna2 at Thr4, Ser17 and Ser237 that stimulates its recruitment to DSBs, resection and subsequent Mec1-dependent phosphorylation
GO:0000993 RNA polymerase II complex binding
IDA
PMID:22689984
Cdc28 kinase activity regulates the basal transcription mach...
KEEP AS NON CORE
Summary: Chymkowitch et al. found by ChIP-seq that Cdc28 localises to >200 genes (e.g. PMA1), that its recruitment is mutually dependent with the CTD kinase Kin28, and that both kinase activities are needed for full CTD-Ser5 phosphorylation. The cached text does not show the physical-binding assay behind the IDA.
Reason: A cell-cycle-independent transcriptional role of Cdc28 at basal-transcription machinery is documented, but it is peripheral to the CDK's core cell-cycle function; the physical RNAPII-binding claim is deferred to the SGD curator who read the full text.
Supporting Evidence:
PMID:22689984
ChIP followed by high-throughput sequencing (ChIP-seq) revealed that Cdc28 localizes to at least 200 genes
PMID:22689984
recruitment of Cdc28 and Kin28 to PMA1 is mutually dependent
GO:0004672 protein kinase activity
HDA
PMID:16319894
Global analysis of protein phosphorylation in yeast.
ACCEPT
Summary: Proteome-chip survey of yeast kinases (Ptacek et al.) recorded in vitro substrates for Cdc28 with several cyclins.
Reason: High-throughput but correct; the parent of the more specific cyclin-dependent Ser/Thr kinase term.
Supporting Evidence:
PMID:16319894
four cyclin-dependent kinases that vary only in their cyclin subunits
GO:0004672 protein kinase activity
IEA
GO_REF:0000002
ACCEPT
Summary: InterPro Ser/Thr kinase active-site signature mapping.
Reason: Correct, general MF from domain signature; more specific terms are present.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:12050115
Phosphorylation of the mitotic regulator Pds1/securin by Cdc...
ACCEPT
Summary: Cdc28 phosphorylates Pds1/securin, promoting Pds1-Esp1 binding and nuclear localisation of separase.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:12050115
Here we show that Pds1 is a substrate of the cyclin-dependent kinase Cdc28. Phosphorylation of Pds1 by Cdc28 is important for efficient binding of Pds1 to Esp1 and for promoting the nuclear localization of Esp1.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:16096060
Cdk1-dependent regulation of the mitotic inhibitor Wee1.
ACCEPT
Summary: Cdk1 directly phosphorylates its own inhibitor Swe1; phosphorylation activates Swe1 and stabilises the Swe1-Cdk1 complex.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:16096060
Phosphorylation of Swe1 by Cdk1 activates Swe1 and is required for formation of a stable Swe1-Cdk1 complex that maintains Cdk1 in the inhibited state.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:17460120
Protein arms in the kinetochore-microtubule interface of the...
ACCEPT
Summary: Miranda et al. studied the DASH/Dam1 kinetochore complex; the cached text (abstract and partial full text) does not mention the Cdc28 kinase assay that SGD used, which is presumably in the full text (in vitro phosphorylation of DASH by Cdk1 is a standard control in this literature).
Reason: The activity is unquestionably correct for Cdc28; the specific assay cannot be verified from the cached text, so the curator's reading of the full text is deferred to.
Supporting Evidence:
PMID:17460120
The yeast DASH complex is a heterodecameric component of the kinetochore necessary for accurate chromosome segregation.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:21498574
Phosphorylation-dependent regulation of the F-BAR protein Ho...
ACCEPT
Summary: Hof1 is phosphorylated by three mitotic kinases including Cdk1, regulating its relocalisation during cytokinesis.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:21498574
three different mitotic kinases phosphorylate Hof1 to regulate its relocalization from septins to the medial AMR to promote cytokinesis: Cdk1
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:22156209
Restriction of histone gene transcription to S phase by phos...
ACCEPT
Summary: Cdk1 and CK2 phosphorylate the chromatin boundary protein Yta7 in S phase, releasing it from HTA1 chromatin.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:22156209
phosphorylation of the conserved chromatin boundary protein Yta7 by both cyclin-dependent kinase 1 (Cdk1) and casein kinase 2 (CK2)
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:22521784
Spatiotemporal regulation of Ipl1/Aurora activity by direct ...
ACCEPT
Summary: Cdk1 phosphorylates Ipl1/Aurora on two N-terminal serines, suppressing Ipl1-Bim1 association until anaphase.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:22521784
We show that Cdk1 phosphorylates Ipl1 on two serine residues in the N-terminal domain, thereby suppressing its association with the microtubule plus-end tracking protein Bim1 until the onset of anaphase.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:22563681
Cyclin-dependent kinase-dependent phosphorylation of Lif1 an...
ACCEPT
Summary: CDK phosphorylates Lif1 Ser261 in S/G2, promoting Lif1 binding to DSBs and imprecise NHEJ.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:22563681
Lif1, a component of the DNA ligase IV complex in Saccharomyces cerevisiae, was phosphorylated by cyclin-dependent kinase (CDK) at Ser261 during the S to G2 phase but not during G1 phase.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:22689984
Cdc28 kinase activity regulates the basal transcription mach...
ACCEPT
Summary: Cdc28 kinase activity contributes with Kin28 to CTD-Ser5 phosphorylation at a subset of genes.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:22689984
the activity of both kinases is required for full phosphorylation of C-terminal domain-Ser5, for efficient transcription, and for mRNA capping
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:23314252
Sequential primed kinases create a damage-responsive phospho...
ACCEPT
Summary: Cdk1 provides the priming phosphorylation on Eco1 that is followed by Cdc7-Dbf4 and Mck1 to create the Cdc4 phosphodegron.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:23314252
Eco1 degradation requires the sequential actions of Cdk1 and two additional kinases, Cdc7-Dbf4 and the GSK-3 homolog Mck1
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:24319056
Global analysis of cdc14 dephosphorylation sites reveals ess...
ACCEPT
Summary: Global Cdc14 dephosphorylation-site study; Smc4 and Bud3 are phosphorylated by Cdk1 and dephosphorylated by Cdc14.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:24319056
The regulation of both Smc4 and Bud3 is mediated by Cdk1 and Cdc14.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:25602519
CDC28 phosphorylates Cac1p and regulates the association of ...
ACCEPT
Summary: CDC28 phosphorylates the CAF-I subunit Cac1 on Ser94/Ser515 in early S phase, regulating its chromatin association.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:25602519
another kinase, CDC28, phosphorylates Cac1p on serines 94 and 515 in early S phase and regulates its association with chromatin, but not its association with PCNA
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:27203178
The Yeast Cyclin-Dependent Kinase Routes Carbon Fluxes to Fu...
ACCEPT
Summary: Cdk1 phosphorylates and activates neutral trehalase Nth1 at G1/S.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:27203178
At the G1/S transition, Cdk1 phosphorylates and activates the enzyme Nth1, which funnels the storage carbohydrate trehalose into central carbon metabolism.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:32083180
Phosphoregulation of Rad51/Rad52 by CDK1 functions as a mole...
ACCEPT
Summary: G2/M Cdc28-cyclin phosphorylates Rad51 and Rad52.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:32083180
the budding yeast CDK1, Cdc28, phosphorylates the major homologous recombination regulators Rad51 and Rad52. This phosphorylation occurs in the G2/M phase by Cdc28 in combination with G2/M phase cyclins.
GO:0004674 protein serine/threonine kinase activity
IDA
PMID:8930895
Interaction between yeast Cdc6 protein and B-type cyclin/Cdc...
ACCEPT
Summary: Cdc6 contains five Cdc28 consensus sites and is a Cdc28 substrate in vitro; it binds preferentially B-type cyclin-Cdc28 complexes.
Reason: Direct demonstration of Cdc28-dependent Ser/Thr phosphorylation of a physiological substrate; core catalytic activity.
Supporting Evidence:
PMID:8930895
Cdc6, which is a phosphoprotein in vivo, contains five Cdc28 consensus sites and is a substrate of the Cdc28 kinase in vitro.
GO:0004674 protein serine/threonine kinase activity
IMP
PMID:21841787
Cell cycle regulation of DNA double-strand break end resecti...
ACCEPT
Summary: In vivo Cdk1-dependent phosphorylation of Dna2 (mutant analysis).
Reason: Mutant evidence corroborating the direct kinase activity documented by the IDA row from the same paper.
Supporting Evidence:
PMID:21841787
we provide evidence for Cdk1-dependent phosphorylation of the resection nuclease Dna2 at Thr4, Ser17 and Ser237
GO:0004674 protein serine/threonine kinase activity
IMP
PMID:22521784
Spatiotemporal regulation of Ipl1/Aurora activity by direct ...
ACCEPT
Summary: Non-phosphorylatable Ipl1 mutants show premature spindle targeting, demonstrating the in vivo relevance of Cdk1 phosphorylation.
Reason: Mutant evidence corroborating the direct kinase activity documented by the IDA row from the same paper.
Supporting Evidence:
PMID:22521784
Failure to phosphorylate Ipl1 leads to its premature targeting to the metaphase spindle
GO:0004674 protein serine/threonine kinase activity
IMP
PMID:8930895
Interaction between yeast Cdc6 protein and B-type cyclin/Cdc...
ACCEPT
Summary: Cdc6 mutant lacking the Cdc28-interaction domain cannot support growth.
Reason: Mutant evidence corroborating the direct kinase activity documented by the IDA row from the same paper.
Supporting Evidence:
PMID:8930895
Deletion of the Cdc28 interaction domain from Cdc6 yields a protein that cannot support growth.
GO:0004693 cyclin-dependent protein serine/threonine kinase activity
IBA
GO_REF:0000033
ACCEPT
Summary: Cdc28 is the archetypal cyclin-dependent Ser/Thr protein kinase: catalytically inert as a monomer, activated by cyclin binding and Cak1 Thr169 phosphorylation, it phosphorylates S/T-P motifs on hundreds of substrates. Family-wide PAINT node with the target among the experimental sources.
Reason: Defining core molecular function; directly demonstrated (PMID:2142620) and inherited across the CDK family.
Supporting Evidence:
PMID:2142620
we demonstrate that the Cln2 polypeptide interacts with p34CDC28 to form an active protein kinase complex
file:yeast/CDC28/CDC28-deep-research-falcon.md
Its best-supported molecular function is transfer of the terminal phosphate of ATP to serine or threonine residues in protein substrates
GO:0004693 cyclin-dependent protein serine/threonine kinase activity
IDA
PMID:2142620
G1-specific cyclins of S. cerevisiae: cell cycle periodicity...
ACCEPT
Summary: Cln2-p34CDC28 forms an active protein kinase complex.
Reason: Direct demonstration of cyclin-dependent kinase activity; core MF.
Supporting Evidence:
PMID:2142620
we demonstrate that the Cln2 polypeptide interacts with p34CDC28 to form an active protein kinase complex
GO:0004693 cyclin-dependent protein serine/threonine kinase activity
IDA
PMID:27068241
Fimbrin phosphorylation by metaphase Cdk1 regulates actin ca...
ACCEPT
Summary: Metaphase cyclin-Cdk1 phosphorylates fimbrin Thr103 in vivo and in vitro, regulating actin cable assembly.
Reason: Direct cyclin-Cdk1 phosphorylation of a physiological substrate; core MF.
Supporting Evidence:
PMID:27068241
Fimbrin is specifically phosphorylated on threonine 103 by the metaphase cyclin-Cdk1 complex, in vivo and in vitro.
GO:0004693 cyclin-dependent protein serine/threonine kinase activity
IEA
GO_REF:0000120
ACCEPT
Summary: EC 2.7.11.22 (cyclin-dependent kinase) mapping.
Reason: Correct EC-based inference of the core MF.
GO:0005515 protein binding
IPI
PMID:10688190
A comprehensive analysis of protein-protein interactions in ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln3 interaction from a genome-wide yeast two-hybrid screen (Uetz et al.). Cln3 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:10688190
These approaches resulted in the detection of 957 putative interactions involving 1,004 S. cerevisiae proteins.
GO:0005515 protein binding
IPI
PMID:10688190
A comprehensive analysis of protein-protein interactions in ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a genome-wide yeast two-hybrid screen (Uetz et al.). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:10688190
These approaches resulted in the detection of 957 putative interactions involving 1,004 S. cerevisiae proteins.
GO:0005515 protein binding
IPI
PMID:10688190
A comprehensive analysis of protein-protein interactions in ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb5 interaction from a genome-wide yeast two-hybrid screen (Uetz et al.). Clb5 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:10688190
These approaches resulted in the detection of 957 putative interactions involving 1,004 S. cerevisiae proteins.
GO:0005515 protein binding
IPI
PMID:10688190
A comprehensive analysis of protein-protein interactions in ...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cak1 interaction from a genome-wide yeast two-hybrid screen (Uetz et al.). Cak1 is the CDK-activating kinase that phosphorylates Cdc28 Thr169; Cdc28 is its substrate.
Reason: Generic protein binding is uninformative; the enzyme-substrate relationship belongs on Cak1 (CDK-activating kinase activity), and no distinct molecular function of Cdc28 is supported by the interaction. Removal does not dispute the interaction.
Supporting Evidence:
PMID:10688190
These approaches resulted in the detection of 957 putative interactions involving 1,004 S. cerevisiae proteins.
GO:0005515 protein binding
IPI
PMID:10913169
Cks1 is required for G(1) cyclin-cyclin-dependent kinase act...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln3 interaction from a reconstitution of cyclin-Cdc28-Cks1 complexes in insect cells (Reynard et al.). Cln3 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:10913169
Cks1 can both stabilize Cln2-Cdc28 complexes and activate intact complexes in vitro
GO:0005515 protein binding
IPI
PMID:10913169
Cks1 is required for G(1) cyclin-cyclin-dependent kinase act...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a reconstitution of cyclin-Cdc28-Cks1 complexes in insect cells (Reynard et al.). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:10913169
Cks1 can both stabilize Cln2-Cdc28 complexes and activate intact complexes in vitro
GO:0005515 protein binding
IPI
PMID:10913169
Cks1 is required for G(1) cyclin-cyclin-dependent kinase act...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a reconstitution of cyclin-Cdc28-Cks1 complexes in insect cells (Reynard et al.). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:10913169
Cks1 can both stabilize Cln2-Cdc28 complexes and activate intact complexes in vitro
GO:0005515 protein binding
IPI
PMID:11283351
A comprehensive two-hybrid analysis to explore the yeast pro...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a comprehensive two-hybrid interactome (Ito et al.). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:11283351
we have completed the comprehensive analysis using this system to identify 4,549 two-hybrid interactions among 3,278 proteins
GO:0005515 protein binding
IPI
PMID:11805826
Functional organization of the yeast proteome by systematic ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a large-scale TAP/mass-spectrometry complex survey (Gavin et al. 2002). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:11805826
We used tandem-affinity purification (TAP) and mass spectrometry in a large-scale approach to characterize multiprotein complexes in Saccharomyces cerevisiae.
GO:0005515 protein binding
IPI
PMID:11805826
Functional organization of the yeast proteome by systematic ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a large-scale TAP/mass-spectrometry complex survey (Gavin et al. 2002). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:11805826
We used tandem-affinity purification (TAP) and mass spectrometry in a large-scale approach to characterize multiprotein complexes in Saccharomyces cerevisiae.
GO:0005515 protein binding
IPI
PMID:11805826
Functional organization of the yeast proteome by systematic ...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a large-scale TAP/mass-spectrometry complex survey (Gavin et al. 2002). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:11805826
We used tandem-affinity purification (TAP) and mass spectrometry in a large-scale approach to characterize multiprotein complexes in Saccharomyces cerevisiae.
GO:0005515 protein binding
IPI
PMID:11805837
Systematic identification of protein complexes in Saccharomy...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a HMS-PCI mass-spectrometric complex identification (Ho et al.). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:11805837
Beginning with 10% of predicted yeast proteins as baits, we detected 3,617 associated proteins covering 25% of the yeast proteome.
GO:0005515 protein binding
IPI
PMID:11805837
Systematic identification of protein complexes in Saccharomy...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a HMS-PCI mass-spectrometric complex identification (Ho et al.). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:11805837
Beginning with 10% of predicted yeast proteins as baits, we detected 3,617 associated proteins covering 25% of the yeast proteome.
GO:0005515 protein binding
IPI
PMID:11805837
Systematic identification of protein complexes in Saccharomy...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a HMS-PCI mass-spectrometric complex identification (Ho et al.). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:11805837
Beginning with 10% of predicted yeast proteins as baits, we detected 3,617 associated proteins covering 25% of the yeast proteome.
GO:0005515 protein binding
IPI
PMID:14690591
Assigning function to yeast proteins by integration of techn...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln3 interaction from a multi-technology functional survey of uncharacterised ORFs (Hazbun et al.). Cln3 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:14690591
These proteins were subjected to affinity purification and mass spectrometry analysis to identify copurifying proteins, two-hybrid analysis to identify interacting proteins
GO:0005515 protein binding
IPI
PMID:14690591
Assigning function to yeast proteins by integration of techn...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a multi-technology functional survey of uncharacterised ORFs (Hazbun et al.). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:14690591
These proteins were subjected to affinity purification and mass spectrometry analysis to identify copurifying proteins, two-hybrid analysis to identify interacting proteins
GO:0005515 protein binding
IPI
PMID:14690591
Assigning function to yeast proteins by integration of techn...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cak1 interaction from a multi-technology functional survey of uncharacterised ORFs (Hazbun et al.). Cak1 is the CDK-activating kinase that phosphorylates Cdc28 Thr169; Cdc28 is its substrate.
Reason: Generic protein binding is uninformative; the enzyme-substrate relationship belongs on Cak1 (CDK-activating kinase activity), and no distinct molecular function of Cdc28 is supported by the interaction. Removal does not dispute the interaction.
Supporting Evidence:
PMID:14690591
These proteins were subjected to affinity purification and mass spectrometry analysis to identify copurifying proteins, two-hybrid analysis to identify interacting proteins
GO:0005515 protein binding
IPI
PMID:16429126
Proteome survey reveals modularity of the yeast cell machine...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a genome-wide affinity-purification complex screen (Gavin et al. 2006). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:16429126
Here we report the first genome-wide screen for complexes in an organism, budding yeast, using affinity purification and mass spectrometry.
GO:0005515 protein binding
IPI
PMID:16429126
Proteome survey reveals modularity of the yeast cell machine...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a genome-wide affinity-purification complex screen (Gavin et al. 2006). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:16429126
Here we report the first genome-wide screen for complexes in an organism, budding yeast, using affinity purification and mass spectrometry.
GO:0005515 protein binding
IPI
PMID:16429126
Proteome survey reveals modularity of the yeast cell machine...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a genome-wide affinity-purification complex screen (Gavin et al. 2006). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:16429126
Here we report the first genome-wide screen for complexes in an organism, budding yeast, using affinity purification and mass spectrometry.
GO:0005515 protein binding
IPI
PMID:17289571
A mechanism for cell-cycle regulation of MAP kinase signalin...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a mechanistic study of Cln/CDK inhibition of the mating MAPK scaffold Ste5 (Strickfaden et al.). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:17289571
Cln/CDK disrupts Ste5 membrane localization by phosphorylating a cluster of sites that flank a small, basic, membrane-binding motif in Ste5.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln3 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Cln3 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln1 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Cln1 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln2 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Cln2 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cks1 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Cks1 is the phospho-adaptor subunit of the CDK holoenzyme.
Reason: Generic protein binding is uninformative; the biologically meaningful relationship (Cks1 as a subunit of the Cdc28 holoenzyme, GO:0000307) is already captured by the complex annotations. Removal does not dispute the interaction, which is well established.
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb5 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Clb5 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb6 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Clb6 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:20489023
A global protein kinase and phosphatase interaction network ...
REMOVE
Summary: Bare protein-binding record of the Cdc28-Cak1 interaction from a global kinase/phosphatase interaction network by AP-MS (Breitkreutz et al.). Cak1 is the CDK-activating kinase that phosphorylates Cdc28 Thr169; Cdc28 is its substrate.
Reason: Generic protein binding is uninformative; the enzyme-substrate relationship belongs on Cak1 (CDK-activating kinase activity), and no distinct molecular function of Cdc28 is supported by the interaction. Removal does not dispute the interaction.
Supporting Evidence:
PMID:20489023
We identified a kinase and phosphatase interaction (KPI) network of 1844 interactions in budding yeast by mass spectrometric analysis of protein complexes.
GO:0005515 protein binding
IPI
PMID:23217712
CDK-dependent Hsp70 Phosphorylation controls G1 cyclin abund...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Cln3 interaction from a study of CDK-dependent Hsp70 (Ssa1) phosphorylation controlling Cln3 abundance (Truman et al.). Cln3 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:23217712
the mitotic CDK Cdk1 phosphorylates T36 to block Cln3 accumulation in G2/M
GO:0005515 protein binding
IPI
PMID:23267104
Proteome-wide protein interaction measurements of bacterial ...
REMOVE
Summary: Bare protein-binding record of a Cdc28-Cdc6 interaction attributed to a microfluidic interaction screen of Streptococcus pneumoniae proteins of unknown function (Meier et al.). The cached title, abstract and available full text of PMID:23267104 concern S. pneumoniae proteins and do not mention yeast, Cdc28 or Cdc6; the Cdc6-Cdc28 interaction itself is well documented elsewhere (PMID:8930895).
Reason: Generic protein binding is uninformative, and the cited reference cannot be verified as supporting a yeast Cdc28-Cdc6 interaction from the cached text (see reference_review). The interaction is real (PMID:8930895) and its functional content (Cdc6 as a Clb-Cdc28 substrate) is captured under the kinase-activity rows.
Supporting Evidence:
PMID:8930895
Cdc6 and Cdc28 can be coimmunoprecipitated from extracts, Cdc6 is retained on the Cdc28-binding matrix p13-agarose, and Cdc28 is retained on an affinity column charged with bacterially produced Cdc6.
GO:0005515 protein binding
IPI
PMID:37968396
The social and structural architecture of the yeast protein ...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb5 interaction from a saturated AP-MS yeast interactome (Michaelis et al.). Clb5 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:37968396
The 4,159 pull-downs generated a highly structured network of 3,927 proteins connected by 31,004 interactions
GO:0005515 protein binding
IPI
PMID:7815540
Simian virus 40 large T antigen affects the Saccharomyces ce...
REMOVE
Summary: Bare protein-binding record of Cdc28 with heterologously expressed SV40 large T antigen (SV40 large T antigen expressed in yeast binds and co-precipitates with p34CDC28 (Nacht et al.)).
Reason: Generic protein binding is uninformative, and the partner is a heterologous viral oncoprotein expressed artificially in yeast, so the interaction has no physiological meaning for CDC28 function. Removal does not dispute the reported interaction.
Supporting Evidence:
PMID:7815540
Both p34CDC28 and p34CDC2Hs were shown to bind to a chimeric T antigen-glutathione S-transferase fusion protein
GO:0005515 protein binding
IPI
PMID:8253070
A new pair of B-type cyclins from Saccharomyces cerevisiae t...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb5 interaction from a identification of the B-type cyclins Clb5/Clb6 (Kuhne and Linder). Clb5 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:8253070
Both cyclins have the potential to interact with the p34CDC28 kinase in vivo.
GO:0005515 protein binding
IPI
PMID:8253070
A new pair of B-type cyclins from Saccharomyces cerevisiae t...
MODIFY
Summary: Bare protein-binding record of the Cdc28-Clb6 interaction from a identification of the B-type cyclins Clb5/Clb6 (Kuhne and Linder). Clb6 is one of the nine cyclins that bind and activate Cdc28, so the informative molecular function is cyclin binding.
Reason: GO:0005515 carries no functional information. The partner is a cyclin, and cyclin binding is the defining regulatory interaction of a CDK, so the row should be recorded as cyclin binding (GO:0030332); the holoenzyme itself is captured by GO:0000307.
Proposed replacements: cyclin binding
Supporting Evidence:
PMID:8253070
Both cyclins have the potential to interact with the p34CDC28 kinase in vivo.
GO:0005524 ATP binding
IEA
GO_REF:0000002
ACCEPT
Summary: InterPro protein-kinase domain / ATP-binding-site signature; Cdc28 binds ATP at Lys40 in the canonical kinase fold (UniProt BINDING 14-22, 40).
Reason: Correct ligand-binding MF implied by the kinase activity and the domain signatures.
GO:0005634 nucleus
IBA
GO_REF:0000033
ACCEPT
Summary: Cdc28 acts in the nucleus on replication, transcription and chromosome-segregation substrates; Cln3-Cdc28 becomes nuclear in late G1 (PMID:14685274). Family-wide PAINT node.
Reason: Core location; directly shown for Cdc28 and inherited across the CDK family.
Supporting Evidence:
PMID:14685274
Both Cln3 and Cdc28 are mainly cytoplasmic during early G1, and become nuclear in late G1.
GO:0005634 nucleus
IDA
PMID:14685274
Recruitment of Cdc28 by Whi3 restricts nuclear accumulation ...
ACCEPT
Summary: Wang et al. showed that Cln3 and Cdc28 are mainly cytoplasmic in early G1 (retained by Whi3) and accumulate in the nucleus in late G1 via the Cln3 NLS.
Reason: Direct localisation evidence; nuclear Cdc28 is where Start and S-phase substrates are phosphorylated.
Supporting Evidence:
PMID:14685274
Both Cln3 and Cdc28 are mainly cytoplasmic during early G1, and become nuclear in late G1.
GO:0005737 cytoplasm
IBA
GO_REF:0000033
ACCEPT
Summary: A substantial pool of Cdc28 is cytoplasmic (early-G1 Cln3-Cdc28 retained by Whi3; SPB/cytoplasmic-microtubule and bud-cortex pools).
Reason: Correct location supported by immunofluorescence (PMID:3312233) and fractionation; Cdc28 acts on cytoplasmic substrates (polarity, septin, actin regulators).
Supporting Evidence:
PMID:3312233
By using immunofluorescence methods, the CDC28 product was shown to be primarily cytoplasmic in distribution.
GO:0005737 cytoplasm
IDA
PMID:3312233
Subcellular localization of a protein kinase required for ce...
ACCEPT
Summary: Wittenberg et al. localised the CDC28 product by immunofluorescence as primarily cytoplasmic, with a fraction tightly associated with the detergent-insoluble cytoplasmic matrix.
Reason: Direct localisation evidence (1987 immunofluorescence); later work shows the distribution is cell-cycle regulated, but a cytoplasmic pool is real.
Supporting Evidence:
PMID:3312233
By using immunofluorescence methods, the CDC28 product was shown to be primarily cytoplasmic in distribution.
GO:0005783 endoplasmic reticulum
IDA
PMID:17560371
Cyclin Cln3 is retained at the ER and released by the J chap...
KEEP AS NON CORE
Summary: Verges et al. showed that Cln3 is retained at the ER in early G1 by binding Cdc28, a fraction of which also associates with the ER, until Ydj1 releases the complex in late G1.
Reason: A minor, cell-cycle-stage-specific ER-associated pool tied to Cln3 retention; genuine but not a principal site of action.
Supporting Evidence:
PMID:17560371
ER retention requires binding of Cln3 to the cyclin-dependent kinase Cdc28, a fraction of which also associates to the ER.
GO:0005935 cellular bud neck
IDA
PMID:12554645
Yeast Cdk1 translocates to the plus end of cytoplasmic micro...
KEEP AS NON CORE
Summary: Maekawa et al. showed that in G1/S Cdc28 associates with SPBs, phosphorylates Kar9, and then moves with Kar9 along cytoplasmic microtubules to their plus ends at the bud neck/bud cortex to align the spindle.
Reason: Transient, cell-cycle-stage-specific localisation of a Cdc28 pool at the bud neck/cortex during spindle orientation; real but non-core. The same paper also documents an SPB-associated pool.
Supporting Evidence:
PMID:12554645
in G(1)/S the yeast Cdk1, Cdc28, associates with SPBs and phosphorylates Kar9. Thirdly, Kar9 and Cdc28 then move from the SPB to the plus end of CMs directed towards the bud.
PMID:12554645
the similar cell cycle-dependent movement of Cdc28 and Kar9 from the SPB to the bud neck
GO:0006303 double-strand break repair via nonhomologous end joining
IMP
PMID:22563681
Cyclin-dependent kinase-dependent phosphorylation of Lif1 an...
KEEP AS NON CORE
Summary: CDK phosphorylation of Lif1 Ser261 in S/G2 promotes stable Lif1 binding to DSBs and imprecise, resection-associated NHEJ in G2/M cells.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:22563681
This phosphorylation was required for efficient NHEJ in G2/M cells, rather than in G1 cells. It also promotes the stable binding of Lif1 protein to DSBs
GO:0006355 regulation of DNA-templated transcription
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: ARBA mapping to regulation of DNA-templated transcription; Cdc28 regulates transcription through SBF/MBF activation (Whi5, Swi6), Swi5 import, Yta7 release and CTD-Ser5 phosphorylation.
Reason: Correct but very general; transcriptional regulation is a downstream output of the CDK, better captured by the specific rows.
GO:0006355 regulation of DNA-templated transcription
IMP
PMID:8319908
CLB5 and CLB6, a new pair of B cyclins involved in DNA repli...
KEEP AS NON CORE
Summary: Schwob and Nasmyth showed that CLB5/CLB6 transcription in late G1 depends on Cln (Cln-Cdc28) activity, i.e. Cdc28 drives the G1/S transcriptional program.
Reason: Cln-Cdc28 activation of the G1/S transcriptional program is real, but the general transcription term is a downstream output subsumed by the G1/S transition core function.
Supporting Evidence:
PMID:8319908
Transcription of CLB5 and CLB6 is normally dependent on Cln activity, but ectopic CLB5 expression allows cells to proliferate in the absence of Cln cyclins.
GO:0006370 7-methylguanosine mRNA capping
IMP
PMID:22689984
Cdc28 kinase activity regulates the basal transcription mach...
MARK AS OVER ANNOTATED
Summary: Chymkowitch et al. found that Cdc28 kinase activity, together with Kin28, is required for full CTD-Ser5 phosphorylation, efficient transcription and mRNA capping at a subset of genes such as PMA1.
Reason: Cdc28 does not perform any step of cap addition (Cet1/Ceg1/Abd1 do); the capping defect is a consequence of reduced CTD-Ser5 phosphorylation and capping-enzyme recruitment. Requirement is not participation, so 'involved_in 7-methylguanosine mRNA capping' over-reads the evidence; the transcriptional regulation rows capture the finding.
Supporting Evidence:
PMID:22689984
the activity of both kinases is required for full phosphorylation of C-terminal domain-Ser5, for efficient transcription, and for mRNA capping
GO:0006892 post-Golgi vesicle-mediated transport
IMP
PMID:22767578
Cdk1-dependent control of membrane-trafficking dynamics.
MODIFY
Summary: McCusker et al. showed that Cdk1 inhibition causes rapid mistargeting of post-Golgi exocytic vesicles away from the bud and disorganises exocytic/endocytic zones, i.e. Cdk1 modulates membrane-trafficking dynamics.
Reason: Cdc28 is not part of the vesicle-transport machinery; it regulates it (through Cdc42/polarity regulators). The regulatory term is the accurate one.
Supporting Evidence:
PMID:22767578
Analysis of post-Golgi vesicle dynamics after Cdk1 inhibition demonstrates that exocytic vesicles are rapidly mistargeted away from the growing bud
PMID:22767578
Cdk1 thus modulates membrane-trafficking dynamics
GO:0006974 DNA damage response
IDA
PMID:26801641
Enrichment of Cdk1-cyclins at DNA double-strand breaks stimu...
KEEP AS NON CORE
Summary: Cdk1-cyclin complexes are enriched at DSBs and phosphorylate the Fun30 remodeler on Ser28 to stimulate resection and checkpoint activation.
Reason: Cdc28 is a genuine, direct participant in the DNA damage response (phosphorylating Sae2, Dna2, Fun30, Rad51/52, Lif1), but this is a non-core output relative to cell-cycle control.
Supporting Evidence:
PMID:26801641
Fun30 is phosphorylated by Cdk1 on Serine 28 to stimulate its functions in DNA damage response including resection of DSB ends.
GO:0007130 synaptonemal complex assembly
IMP
PMID:20825495
Cyclin-dependent kinase promotes formation of the synaptonem...
KEEP AS NON CORE
Summary: Clb5/6-Cdc28 localises to meiotic chromosomes and compromised CDK activity in prophase causes defective synaptonemal complex formation without affecting DSB formation.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:20825495
Compromised CDK activity in meiotic prophase leads to defective SC formation without affecting DSB formation.
GO:0007165 signal transduction
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Generic 'signal transduction' PAINT annotation at a CDK subfamily node (Arabidopsis CDKs, mouse Cdk2/Cdk1, pombe cdc2).
Reason: Cdc28 is an intracellular signalling kinase, so the term is not wrong, but it is far too general to be informative; the specific cell-cycle transition and regulation terms carry the content.
GO:0010494 cytoplasmic stress granule
HDA
PMID:26777405
ATPase-Modulated Stress Granules Contain a Diverse Proteome ...
MARK AS OVER ANNOTATED
Summary: Cdc28 was detected among proteins in purified stress-granule cores in a proteomic survey (Jain et al.).
Reason: Single high-throughput proteomic detection with no functional follow-up for Cdc28; not supported as a site of Cdc28 function.
Supporting Evidence:
PMID:26777405
Proteomic analysis of stress granule cores reveals a dense network of protein-protein interactions
GO:0010568 regulation of budding cell apical bud growth
IGI
PMID:17417630
Cdk1 coordinates cell-surface growth with the cell cycle.
KEEP AS NON CORE
Summary: McCusker et al. showed that G1 cyclin-Cdk1 phosphorylates Cdc24-associated polarity proteins and that a non-phosphorylatable mutant causes bud-growth defects, linking Cdk1 to control of polarized cell-surface growth.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:17417630
we show that G1 cyclin-Cdk1 complexes specifically phosphorylate multiple proteins associated with Cdc24, the guanine nucleotide-exchange factor (GEF) that activates the Cdc42 GTPase. A mutant form of a Cdc24-associated protein that fails to undergo Cdk1-dependent phosphorylation causes defects in bud growth.
GO:0010568 regulation of budding cell apical bud growth
IMP
PMID:17417630
Cdk1 coordinates cell-surface growth with the cell cycle.
KEEP AS NON CORE
Summary: Cdk1 has an unexpected role in controlling bud growth after bud emergence.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:17417630
we show that G1 cyclin-Cdk1 complexes specifically phosphorylate multiple proteins associated with Cdc24, the guanine nucleotide-exchange factor (GEF) that activates the Cdc42 GTPase. A mutant form of a Cdc24-associated protein that fails to undergo Cdk1-dependent phosphorylation causes defects in bud growth.
GO:0010570 regulation of filamentous growth
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: ARBA mapping to regulation of filamentous growth, consistent with the IMP evidence.
Reason: Correct but non-core morphogenetic output of Cdc28 activity (Swe1/Tyr19 axis).
GO:0010570 regulation of filamentous growth
IMP
PMID:9891070
Control of Saccharomyces cerevisiae filamentous growth by cy...
KEEP AS NON CORE
Summary: Edgington et al. found that specific cdc28 alleles cause constitutive filamentous growth and that the Elm1-Hsl1-Swe1 pathway acts through Cdc28 Tyr19 phosphorylation to modulate filamentous growth.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:9891070
A specific amino acid substitution in the cyclin-dependent protein kinase Cdc28 was found to cause constitutive expression of most filamentous growth characteristics.
GO:0010571 positive regulation of nuclear cell cycle DNA replication
IDA
PMID:14747467
Cell cycle-dependent phosphorylation of the DNA polymerase e...
ACCEPT
Summary: Kesti et al. showed that the Pol epsilon subunit Dpb2 is phosphorylated in late G1 by Cdc28 (Ser-144 directly in vitro), facilitating Pol epsilon function at origins.
Reason: Core function: Clb5/6-Cdc28 (S-CDK) is the kinase that triggers origin firing (via Sld2/Sld3 phosphorylation) and prevents re-replication; cdc28 was among the original 'initiation of DNA synthesis' genes.
Supporting Evidence:
PMID:14747467
The appearance of that species in vivo is dependent upon the Cdc28 cyclin-dependent protein kinase (CDK), which can directly phosphorylate Dpb2 in vitro.
GO:0010571 positive regulation of nuclear cell cycle DNA replication
IMP
PMID:12783856
Control of landmark events in meiosis by the CDK Cdc28 and t...
ACCEPT
Summary: Benjamin et al. showed with cdc28-as1 that Cdc28 is required for premeiotic DNA replication.
Reason: Core function: Clb5/6-Cdc28 (S-CDK) is the kinase that triggers origin firing (via Sld2/Sld3 phosphorylation) and prevents re-replication; cdc28 was among the original 'initiation of DNA synthesis' genes. This row documents the same S-CDK requirement in the meiotic program.
Supporting Evidence:
PMID:12783856
Early inhibition of analog-sensitive cdc28-as1 blocked DNA replication, revealing a previously undetected role for Cdc28.
GO:0010571 positive regulation of nuclear cell cycle DNA replication
IMP
PMID:14747467
Cell cycle-dependent phosphorylation of the DNA polymerase e...
ACCEPT
Summary: Non-phosphorylatable Dpb2 CDK-site mutants show synthetic defects with pol2-11.
Reason: Core function: Clb5/6-Cdc28 (S-CDK) is the kinase that triggers origin firing (via Sld2/Sld3 phosphorylation) and prevents re-replication; cdc28 was among the original 'initiation of DNA synthesis' genes.
Supporting Evidence:
PMID:14747467
inactivation of all three CDK consensus sites in Dpb2 results in a synthetic phenotype with the pol2-11 mutation, leading to decreased spore viability, slow growth, and increased thermosensitivity
GO:0010571 positive regulation of nuclear cell cycle DNA replication
IMP
PMID:4580573
Three additional genes required for deoxyribonucleic acid sy...
ACCEPT
Summary: Hartwell (1973) classified cdc28 with cdc4 and cdc7 as genes required for the initiation of DNA synthesis.
Reason: Core function: Clb5/6-Cdc28 (S-CDK) is the kinase that triggers origin firing (via Sld2/Sld3 phosphorylation) and prevents re-replication; cdc28 was among the original 'initiation of DNA synthesis' genes.
Supporting Evidence:
PMID:4580573
Mutations in three genes (cdc 4, 7, and 28) appear to block a precondition for DNA synthesis since cells carrying these lesions cannot start new rounds of DNA replication after a shift from permissive to restrictive temperature, but can finish rounds that were in progress.
GO:0010696 positive regulation of mitotic spindle pole body separation
IGI
PMID:16688214
Cdk1 regulates centrosome separation by restraining proteoly...
ACCEPT
Summary: Crasta et al.: cells unable to activate Cdk1 (Tyr19) lack spindles because APC-Cdh1 degrades Cin8, Kip1 and Ase1; proteolysis-resistant versions restore SPB separation without Cdc28-Clb activity.
Reason: Core mitotic function: Clb-Cdc28 activated by Tyr19 dephosphorylation directly promotes separation of duplicated SPBs, the first step of spindle assembly, by stabilising Cin8/Kip1/Ase1 against APC-Cdh1.
Supporting Evidence:
PMID:16688214
Tyrosine 19 dephosphorylation of Cdk1 is necessary to specifically prevent proteolysis of these proteins.
GO:0010696 positive regulation of mitotic spindle pole body separation
IGI
PMID:16688214
Cdk1 regulates centrosome separation by restraining proteoly...
ACCEPT
Summary: Crasta et al.: cells unable to activate Cdk1 (Tyr19) lack spindles because APC-Cdh1 degrades Cin8, Kip1 and Ase1; proteolysis-resistant versions restore SPB separation without Cdc28-Clb activity.
Reason: Core mitotic function: Clb-Cdc28 activated by Tyr19 dephosphorylation directly promotes separation of duplicated SPBs, the first step of spindle assembly, by stabilising Cin8/Kip1/Ase1 against APC-Cdh1.
Supporting Evidence:
PMID:16688214
Tyrosine 19 dephosphorylation of Cdk1 is necessary to specifically prevent proteolysis of these proteins.
GO:0010696 positive regulation of mitotic spindle pole body separation
IGI
PMID:16688214
Cdk1 regulates centrosome separation by restraining proteoly...
ACCEPT
Summary: Crasta et al.: cells unable to activate Cdk1 (Tyr19) lack spindles because APC-Cdh1 degrades Cin8, Kip1 and Ase1; proteolysis-resistant versions restore SPB separation without Cdc28-Clb activity.
Reason: Core mitotic function: Clb-Cdc28 activated by Tyr19 dephosphorylation directly promotes separation of duplicated SPBs, the first step of spindle assembly, by stabilising Cin8/Kip1/Ase1 against APC-Cdh1.
Supporting Evidence:
PMID:16688214
Tyrosine 19 dephosphorylation of Cdk1 is necessary to specifically prevent proteolysis of these proteins.
GO:0010696 positive regulation of mitotic spindle pole body separation
IMP
PMID:16688214
Cdk1 regulates centrosome separation by restraining proteoly...
ACCEPT
Summary: Cdk1 Tyr19 dephosphorylation is required for SPB separation via stabilisation of microtubule-associated force-generating proteins.
Reason: Core mitotic function: Clb-Cdc28 activated by Tyr19 dephosphorylation directly promotes separation of duplicated SPBs, the first step of spindle assembly, by stabilising Cin8/Kip1/Ase1 against APC-Cdh1.
Supporting Evidence:
PMID:16688214
we suggest that stabilization of these mechanical force-generating proteins is the predominant role of Cdc28-Clb in centrosome separation
GO:0010696 positive regulation of mitotic spindle pole body separation
IMP
PMID:8887667
Spindle pole body separation in Saccharomyces cerevisiae req...
ACCEPT
Summary: Lim et al.: cdc28-E19 (phosphomimetic Tyr19) cells complete DNA synthesis but cannot separate duplicated SPBs; the requirement holds even when spindle formation is uncoupled from mitosis, indicating a direct role.
Reason: Core mitotic function: Clb-Cdc28 activated by Tyr19 dephosphorylation directly promotes separation of duplicated SPBs, the first step of spindle assembly, by stabilising Cin8/Kip1/Ase1 against APC-Cdh1.
Supporting Evidence:
PMID:8887667
These results imply that dephosphorylation of Tyr-19 is required for the segregation of SPBs.
GO:0010898 positive regulation of triglyceride catabolic process
IGI
PMID:19150427
Cdk1/Cdc28-dependent activation of the major triacylglycerol...
KEEP AS NON CORE
Summary: Kurat et al.: Cdk1/Cdc28 phosphorylates and activates the triacylglycerol lipase Tgl4, coupling lipolysis to bud emergence.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:19150427
the major yeast TG lipase Tgl4, the functional ortholog of murine adipose TG lipase ATGL, is phosphorylated and activated by cyclin-dependent kinase 1 (Cdk1/Cdc28)
GO:0010898 positive regulation of triglyceride catabolic process
IMP
PMID:19150427
Cdk1/Cdc28-dependent activation of the major triacylglycerol...
KEEP AS NON CORE
Summary: Cdk1-dependent Tgl4 activation links TG mobilisation to cell-cycle progression.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:19150427
the major yeast TG lipase Tgl4, the functional ortholog of murine adipose TG lipase ATGL, is phosphorylated and activated by cyclin-dependent kinase 1 (Cdk1/Cdc28)
GO:0030163 protein catabolic process
IGI
PMID:21993622
Cascades of multisite phosphorylation control Sic1 destructi...
MODIFY
Summary: Koivomagi et al.: Cln2-Cdk1 and Clb5-Cdk1 act in processive multisite phosphorylation cascades on Sic1, generating the phosphodegrons that direct it to SCF-mediated destruction at the G1/S transition.
Reason: Cdc28 does the work here (it creates the phosphodegron that SCF-Cdc4 recognises), but the general 'protein catabolic process' term misplaces the kinase inside the degradation pathway; the accurate statement is that CDK phosphorylation positively regulates proteasomal, ubiquitin-dependent degradation of specific substrates.
Supporting Evidence:
PMID:21993622
both Cln2-Cdk1 and Clb5-Cdk1 act in processive multiphosphorylation cascades leading to the phosphorylation of a small number of specific phosphodegrons
GO:0030163 protein catabolic process
IMP
PMID:21098119
SCFCdc4 enables mating type switching in yeast by cyclin-dep...
MODIFY
Summary: Liu et al.: Ash1 associates with active Cdc28 and is targeted to SCF-Cdc4 for degradation in a Cdc28-dependent manner via CDK-phosphorylated diphosphodegrons.
Reason: Cdc28 does the work here (it creates the phosphodegron that SCF-Cdc4 recognises), but the general 'protein catabolic process' term misplaces the kinase inside the degradation pathway; the accurate statement is that CDK phosphorylation positively regulates proteasomal, ubiquitin-dependent degradation of specific substrates.
Supporting Evidence:
PMID:21098119
Ash1 associates with active Cdc28 kinase in vivo and is targeted to SCF(Cdc4) in a Cdc28-dependent fashion in vivo and in vitro.
GO:0032210 regulation of telomere maintenance via telomerase
IGI
PMID:19135888
Cdk1-dependent phosphorylation of Cdc13 coordinates telomere...
KEEP AS NON CORE
Summary: Li et al.: Cdk1 phosphorylation of Cdc13 is essential for efficient telomerase recruitment, favouring Cdc13-Est1 over Stn1-Ten1 binding.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:19135888
Cdk1-dependent phosphorylation of Cdc13 is essential for efficient recruitment of the yeast telomerase complex to telomeres by favoring the interaction of Cdc13 with Est1 rather than the competing Stn1-Ten1 complex
GO:0032880 regulation of protein localization
IMP
PMID:29263158
Clb6-Cdc28 Promotes Ribonucleotide Reductase Subcellular Red...
KEEP AS NON CORE
Summary: Wu et al.: Clb6-Cdc28 phosphorylation of Rnr2 disrupts the Rnr2-Wtm1 interaction and releases Rnr2-Rnr4 from the nucleus in S phase.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:29263158
we propose that Rnr2-Rnr4 redistribution in S phase is triggered by Clb6-Cdc28-mediated phosphorylation of Rnr2, which disrupts the Rnr2-Wtm1 interaction and promotes the release of Rnr2-Rnr4 from the nucleus
GO:0034504 protein localization to nucleus
IMP
PMID:19188495
Antagonistic regulation of Fus2p nuclear localization by phe...
MODIFY
Summary: Ydenberg and Rose: Fus2 accumulates in the nucleus in G2/M and is exported only in G1; Cdc28 activity blocks Fus2 export, in late G1/S by inhibiting Fus3 and again in G2/M.
Reason: Cdc28 does not carry out nuclear import of Fus2; it prevents Fus3-dependent export, i.e. it regulates Fus2 nuclear localisation, largely indirectly through inhibition of pheromone signalling. The regulation term is accurate; the process term over-reads the evidence.
Supporting Evidence:
PMID:19188495
cyclin/Cdc28p-dependent inhibition of Fus3p during late G1 through S phase was sufficient to block exit. However, during G2/M, when Fus3p was activated by pheromone signaling, Cdc28p activity again blocked Fus2p export.
GO:0042393 histone binding
IDA
PMID:20855529
Cks1, Cdk1, and the 19S proteasome collaborate to regulate g...
KEEP AS NON CORE
Summary: Chaves et al.: Cks1, Cdk1 and the 19S proteasome are recruited to chromatin by binding directly to the histone H4 N-terminal tail during gene induction.
Reason: Specific binding activity supporting a non-catalytic, gene-induction role of the Cks1-Cdk1 complex; genuine but peripheral to the core kinase function.
Supporting Evidence:
PMID:20855529
Cks1, Cdk1, and the 19S subunit of the proteasome are recruited to chromatin by binding directly to the histone H4 amino-terminal tail.
GO:0045819 positive regulation of glycogen catabolic process
IMP
PMID:27203179
Cyclin-Dependent Kinase Co-Ordinates Carbohydrate Metabolism...
KEEP AS NON CORE
Summary: Zhao et al.: CDK activity directly controls glycogen phosphorylase Gph1, co-ordinating glycogen mobilisation with division.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:27203179
neutral trehalase (Nth1) and glycogen phosphorylase (Gph1), and show that their activities are likely directly controlled by CDK activity
GO:0045875 negative regulation of sister chromatid cohesion
IMP
PMID:21549314
Cdk1-dependent destruction of Eco1 prevents cohesion establi...
KEEP AS NON CORE
Summary: Lyons and Morgan: Cdk1 phosphorylation of Eco1 after S phase targets it to SCF-Cdc4, restricting cohesion establishment to S phase.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:21549314
Phosphorylation of Eco1 after S phase targets it to SCF(Cdc4) for ubiquitination and subsequent degradation. A nonphosphorylatable mutant of Eco1 establishes cohesion after DNA replication
GO:0045892 negative regulation of DNA-templated transcription
IDA
PMID:14993267
Clb6/Cdc28 and Cdc14 regulate phosphorylation status and cel...
KEEP AS NON CORE
Summary: Geymonat et al.: Clb6-Cdc28 phosphorylates Swi6 Ser160, driving nuclear export of the SBF/MBF subunit and terminating G1/S transcription.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:14993267
Cdc28 combined with the S-phase cyclin Clb6 specifically phosphorylates serine 160 of Swi6 in vitro.
GO:0045892 negative regulation of DNA-templated transcription
IMP
PMID:12081645
The CLN3/SWI6/CLN2 pathway and SNF1 act sequentially to regu...
KEEP AS NON CORE
Summary: Purnapatre et al.: the Cln3-Cdc28/Swi6/Cln2 pathway represses IME1 during growth; cdc28-4 mutants express IME1 prematurely.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:12081645
Moderate IME1 expression occurred in cln3Delta, cln1Delta cln2Delta, cdc28-4 and swi6Delta mutants, even during growth.
GO:0045892 negative regulation of DNA-templated transcription
IMP
PMID:14993267
Clb6/Cdc28 and Cdc14 regulate phosphorylation status and cel...
KEEP AS NON CORE
Summary: Clb6/Cdc28 kinase is required for Swi6 nuclear export.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:14993267
We show that Clb6/Cdc28 kinase is required for this nuclear export.
GO:0045893 positive regulation of DNA-templated transcription
IDA
PMID:1652372
The role of phosphorylation and the CDC28 protein kinase in ...
MODIFY
Summary: Moll et al. showed that CDC28-dependent phosphorylation of three serines in/near the Swi5 NLS keeps Swi5 cytoplasmic in S/G2/M; mutation of these serines gives constitutive nuclear entry. Cdc28 thus prevents, rather than promotes, Swi5-dependent transcription (e.g. of HO) until it is inactivated at mitotic exit.
Reason: The direct finding is negative control of Swi5 nuclear import by Cdc28 phosphorylation; a 'positive regulation of transcription' reading is not supported by the abstract's mechanism (Cdc28 inactivation, not activity, permits Swi5 nuclear entry). The most accurate term for what Cdc28 does here is negative regulation of protein import into nucleus. The curator may have had full-text data in view; if so the term direction should be re-checked against it.
Supporting Evidence:
PMID:1652372
As all three serines are phosphorylated by purified CDC28-dependent H1 kinase activity in vitro, we propose a model in which the CDC28 kinase acts directly to control nuclear entry of SWI5.
GO:0045893 positive regulation of DNA-templated transcription
IGI
PMID:1652372
The role of phosphorylation and the CDC28 protein kinase in ...
MODIFY
Summary: Moll et al. showed that CDC28-dependent phosphorylation of three serines in/near the Swi5 NLS keeps Swi5 cytoplasmic in S/G2/M; mutation of these serines gives constitutive nuclear entry. Cdc28 thus prevents, rather than promotes, Swi5-dependent transcription (e.g. of HO) until it is inactivated at mitotic exit.
Reason: The direct finding is negative control of Swi5 nuclear import by Cdc28 phosphorylation; a 'positive regulation of transcription' reading is not supported by the abstract's mechanism (Cdc28 inactivation, not activity, permits Swi5 nuclear entry). The most accurate term for what Cdc28 does here is negative regulation of protein import into nucleus. The curator may have had full-text data in view; if so the term direction should be re-checked against it.
Supporting Evidence:
PMID:1652372
Located within or close to the NLS are three serine residues, mutation of which results in constitutive nuclear entry.
GO:0045930 negative regulation of mitotic cell cycle
IDA
PMID:10074450
Inhibitory phosphorylation of the APC regulator Hct1 is cont...
MODIFY
Summary: Jaspersen et al.: Cdc28-cyclin phosphorylates the APC activator Hct1/Cdh1 at CDK sites, abolishing its ability to activate the APC; Cdc14 reverses this. Cdc28 thereby blocks its own inactivation and prevents premature mitotic exit.
Reason: The evidence describes negative control of the exit from mitosis (inhibition of APC-Cdh1), not negative regulation of the mitotic cell cycle as a whole (Cdc28 is its principal positive driver). The child term 'negative regulation of exit from mitosis' states exactly what the paper shows.
Supporting Evidence:
PMID:10074450
Purified Hct1 was phosphorylated in vitro at these sites by purified Cdc28-cyclin complexes, and phosphorylation abolished the ability of Hct1 to activate the APC in vitro.
PMID:10074450
Phosphorylation of Hct1 provides a mechanism by which Cdc28 blocks its own inactivation during S phase and early mitosis.
GO:0045931 positive regulation of mitotic cell cycle
IMP
PMID:2165600
Mitotic role for the Cdc28 protein kinase of Saccharomyces c...
ACCEPT
Summary: Reed and Wittenberg: Cdc28 is required for the G1/S transition and additionally for mitosis, with the mitotic function executed in G2 when the kinase is highly active.
Reason: Cdc28 is the positive driver of the mitotic cell cycle at both transitions; correct (if general) core term.
Supporting Evidence:
PMID:2165600
We present evidence that the Cdc28 protein kinase is also required for mitosis and that this function is executed in the G2 interval of the cell cycle.
GO:0045931 positive regulation of mitotic cell cycle
IMP
PMID:7002718
The selection of S. cerevisiae mutants defective in the star...
ACCEPT
Summary: Reed (1980) isolated start mutants; cdc28 alleles arrest at Start, the point where mating pheromone interrupts division.
Reason: Classical Start phenotype establishing Cdc28 as a positive driver of cell-cycle progression; core.
Supporting Evidence:
PMID:7002718
Each mutation produces stage-specific arrest of cell division at start, the same point where mating pheromone interrupts division.
GO:0045944 positive regulation of transcription by RNA polymerase II
IMP
PMID:22689984
Cdc28 kinase activity regulates the basal transcription mach...
KEEP AS NON CORE
Summary: Cdc28 kinase activity is required with Kin28 for CTD-Ser5 phosphorylation and efficient transcription of a subset of basal genes (PMA1).
Reason: A cell-cycle-independent transcriptional role of Cdc28 at a subset of genes; genuine but peripheral to the core CDK function.
Supporting Evidence:
PMID:22689984
the activity of both kinases is required for full phosphorylation of C-terminal domain-Ser5, for efficient transcription, and for mRNA capping
GO:0051446 positive regulation of meiotic cell cycle
IDA
PMID:16814718
Cyclin-dependent kinase directly regulates initiation of mei...
ACCEPT
Summary: Henderson et al.: Cdc28 directly phosphorylates Mer2/Rec107 to regulate meiotic DSB formation, coordinating prophase events.
Reason: Core function in the meiotic program: Cdc28 (with Clb5/6 and later Clb1/3/4) drives premeiotic S phase, DSB formation (Mer2 phosphorylation), pachytene exit and the meiotic divisions.
Supporting Evidence:
PMID:16814718
the budding yeast cyclin-dependent kinase Cdc28 directly regulates the formation of the DNA double-strand breaks that initiate recombination by phosphorylating the Mer2/Rec107 protein
GO:0051446 positive regulation of meiotic cell cycle
IMP
PMID:16814718
Cyclin-dependent kinase directly regulates initiation of mei...
ACCEPT
Summary: Non-phosphorylatable Mer2 mutants show that CDK phosphorylation of Mer2 is required for meiotic recombination initiation.
Reason: Core function in the meiotic program: Cdc28 (with Clb5/6 and later Clb1/3/4) drives premeiotic S phase, DSB formation (Mer2 phosphorylation), pachytene exit and the meiotic divisions.
Supporting Evidence:
PMID:16814718
the budding yeast cyclin-dependent kinase Cdc28 directly regulates the formation of the DNA double-strand breaks that initiate recombination by phosphorylating the Mer2/Rec107 protein
GO:0051446 positive regulation of meiotic cell cycle
IMP
PMID:2680756
Pachytene arrest and other meiotic effects of the start muta...
ACCEPT
Summary: Shuster and Byers: start mutants including cdc28 arrest meiotic cells at pachytene after DNA replication and SPB duplication, and CDC28 is additionally required for SPB separation before meiosis II.
Reason: Core function in the meiotic program: Cdc28 (with Clb5/6 and later Clb1/3/4) drives premeiotic S phase, DSB formation (Mer2 phosphorylation), pachytene exit and the meiotic divisions.
Supporting Evidence:
PMID:2680756
CDC28 was additionally found to be required for the SPB separation that precedes spindle formation in preparation for the second meiotic division.
GO:0051447 negative regulation of meiotic cell cycle
IMP
PMID:12081645
The CLN3/SWI6/CLN2 pathway and SNF1 act sequentially to regu...
KEEP AS NON CORE
Summary: Cln3-Cdc28 represses IME1/IME2 during growth, preventing meiotic initiation until growth ceases.
Reason: Genuine regulatory role (Cln3-Cdc28 signalling represses meiotic initiation during growth), but a non-core, developmental-decision output.
Supporting Evidence:
PMID:12081645
during growth IME1 is repressed by the CLN3/SWI6/CLN2 pathway
GO:0051726 regulation of cell cycle
IBA
GO_REF:0000033
ACCEPT
Summary: Family-wide PAINT annotation: CDKs regulate the cell cycle. Cdc28 is the master regulator of the budding-yeast cell cycle.
Reason: Correct and core, although less specific than the transition-level terms also present.
Supporting Evidence:
PMID:22689984
The cyclin-dependent kinase Cdc28 is the master regulator of the cell cycle in Saccharomyces cerevisiae.
file:yeast/CDC28/CDC28-deep-research-falcon.md
Cdc28 is the central catalytic engine of the budding-yeast division cycle. By changing cyclin partners and exploiting cyclin-docking motifs, Cks1-dependent phospho-priming, localization, and kinase/phosphatase feedback, the same catalytic subunit triggers Start, DNA replication, spindle assembly, chromosome segregation, and mitotic exit.
GO:0090169 regulation of spindle assembly
IMP
PMID:21558801
Ase1p phosphorylation by cyclin-dependent kinase promotes co...
ACCEPT
Summary: Juanes et al.: CDK (Clb5/Clb3/Clb4-Cdc28) phosphorylation of the midzone protein Ase1 promotes timely spindle assembly; phenotypes are enhanced in cdc28-4 clb5 mutants and rescued by phosphomimetic Ase1-7D.
Reason: Direct substrate-mediated control of spindle assembly, a core mitotic function of Clb-Cdc28.
Supporting Evidence:
PMID:21558801
Ase1p phosphorylation by CDK promotes the assembly and stability of the mitotic spindle.
GO:0090307 mitotic spindle assembly
IGI
PMID:21558801
Ase1p phosphorylation by cyclin-dependent kinase promotes co...
ACCEPT
Summary: cdc28-4 clb5 double mutants show midzone disruption and transient breaks of the short spindle, rescued by phosphomimetic Ase1.
Reason: Spindle assembly is a core mitotic function of Clb-Cdc28 (SPB separation, Ase1/Cin8/Kip1 stabilisation and phosphorylation).
Supporting Evidence:
PMID:21558801
These phenotypes were enhanced in a cdc28-4 clb5Ξ” mutant to the extent that midzone disruption resulted in transient breaks of the short spindle.
GO:0106310 protein serine kinase activity
IEA
GO_REF:0000116
ACCEPT
Summary: Rhea mapping for protein-serine phosphorylation (RHEA:17989).
Reason: Correct catalytic activity; a component of the cyclin-dependent Ser/Thr kinase activity.
GO:0106310 protein serine kinase activity
ISS
GO_REF:0000024
ACCEPT
Summary: Manual similarity transfer of protein serine kinase activity from human CDK2.
Reason: Correct; directly demonstrated for Cdc28 on numerous serine sites (e.g. Swi6 Ser160, Cac1 Ser94/515).
GO:1901319 positive regulation of trehalose catabolic process
IGI
PMID:27203179
Cyclin-Dependent Kinase Co-Ordinates Carbohydrate Metabolism...
KEEP AS NON CORE
Summary: Zhao et al.: CDK activity directly controls neutral trehalase Nth1, coordinating trehalose mobilisation with division.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:27203179
neutral trehalase (Nth1) and glycogen phosphorylase (Gph1), and show that their activities are likely directly controlled by CDK activity
GO:1901319 positive regulation of trehalose catabolic process
IMP
PMID:27203179
Cyclin-Dependent Kinase Co-Ordinates Carbohydrate Metabolism...
KEEP AS NON CORE
Summary: CDK-dependent activation of Nth1 (also shown by Ewald et al., PMID:27203178).
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:27203179
neutral trehalase (Nth1) and glycogen phosphorylase (Gph1), and show that their activities are likely directly controlled by CDK activity
GO:1902275 regulation of chromatin organization
IPI
PMID:22156209
Restriction of histone gene transcription to S phase by phos...
KEEP AS NON CORE
Summary: Cdk1 (and CK2) phosphorylation of the boundary protein Yta7 releases it from HTA1 chromatin in S phase, permitting histone gene transcription.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:22156209
in S phase, Yta7 is hyperphosphorylated, causing its release from HTA1 chromatin and productive transcription
GO:1902806 regulation of cell cycle G1/S phase transition
NAS
PMID:2569741
A family of cyclin homologs that control the G1 phase in yea...
MODIFY
Summary: Hadwiger et al. isolated CLN1/CLN2 as dosage suppressors of cdc28-ts, defining the G1 cyclins that act with Cdc28 at Start.
Reason: GO:1902806 is defined as a signalling pathway that modulates the activity of a cell-cycle CDK to control G1/S; Cdc28 is that CDK, not its regulator. The finding supports the executing role captured by GO:0000082, which the gene already carries with IBA/IDA evidence.
Supporting Evidence:
PMID:2569741
Two Saccharomyces cerevisiae genes were isolated based upon their dosage-dependent rescue of a temperature-sensitive mutation of the gene CDC28, which encodes a protein kinase involved in control of cell division.
GO:1902889 protein localization to spindle microtubule
IMP
PMID:22521784
Spatiotemporal regulation of Ipl1/Aurora activity by direct ...
KEEP AS NON CORE
Summary: Zimniak et al.: Cdk1 phosphorylation of Ipl1 suppresses its Bim1-dependent targeting to the spindle until anaphase; non-phosphorylatable Ipl1 is prematurely spindle-localised.
Reason: Cdc28 controls the timing of Ipl1 localisation to spindle microtubules by direct phosphorylation (a negative, timing role rather than performing localisation); genuine but non-core.
Supporting Evidence:
PMID:22521784
Failure to phosphorylate Ipl1 leads to its premature targeting to the metaphase spindle and results in constitutive Bim1 phosphorylation, which is normally restricted to anaphase.
GO:1903500 negative regulation of mitotic actomyosin contractile ring assembly
IMP
PMID:24413167
Cdk1-dependent phosphorylation of Iqg1 governs actomyosin ri...
KEEP AS NON CORE
Summary: Naylor and Morgan: Cdk1-dependent phosphorylation of Iqg1 delays bud-neck localisation of Iqg1 and actin, preventing premature actomyosin ring assembly.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:24413167
the Cdk1-dependent phosphorylation state of Iqg1 is a determining factor in the timing of bud neck localization of both Iqg1 and actin, with both proteins accumulating prematurely in cells expressing nonphosphorylatable Iqg1 mutants
GO:1904291 positive regulation of mitotic DNA damage checkpoint
IMP
PMID:15496928
DNA end resection, homologous recombination and DNA damage c...
KEEP AS NON CORE
Summary: Ira et al.: CDK1 (Cdc28) is required for DSB end resection and thereby for Mec1-dependent checkpoint activation and its maintenance.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:15496928
Here we report that DNA damage checkpoint activation by a DSB requires the cyclin-dependent kinase CDK1 (Cdc28) in budding yeast. CDK1 is also required for DSB-induced homologous recombination at any cell cycle stage.
GO:1905168 positive regulation of double-strand break repair via homologous recombination
IMP
PMID:15496928
DNA end resection, homologous recombination and DNA damage c...
KEEP AS NON CORE
Summary: CDK1 is required for DSB-induced homologous recombination (resection, RPA/Rad51 recruitment, and a post-strand-invasion step); its inhibition shifts repair to NHEJ.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:15496928
Here we report that DNA damage checkpoint activation by a DSB requires the cyclin-dependent kinase CDK1 (Cdc28) in budding yeast. CDK1 is also required for DSB-induced homologous recombination at any cell cycle stage.
GO:1905634 regulation of protein localization to chromatin
IDA
PMID:25602519
CDC28 phosphorylates Cac1p and regulates the association of ...
KEEP AS NON CORE
Summary: Jeffery et al.: CDC28 phosphorylation of Cac1 Ser94/Ser515 regulates CAF-I association with chromatin in early S phase.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:25602519
another kinase, CDC28, phosphorylates Cac1p on serines 94 and 515 in early S phase and regulates its association with chromatin, but not its association with PCNA
GO:1990139 protein localization to nuclear periphery
IMP
PMID:20702586
Cdk phosphorylation of a nucleoporin controls localization o...
KEEP AS NON CORE
Summary: Brickner and Brickner: Cdk1 and two CDK sites in the nucleoporin Nup1 are required for peripheral targeting of the active INO1 and GAL1 genes; phosphomimetic Nup1 bypasses Cdk1.
Reason: Genuine Nup1-mediated role, but non-core; note that the phenomenon controlled is localisation of active gene loci (chromatin) to the nuclear periphery rather than protein localisation, so the term is an approximate fit.
Supporting Evidence:
PMID:20702586
the Cdk1 kinase and two Cdk phosphorylation sites in the nucleoporin Nup1 were required for peripheral targeting of INO1 and GAL1
GO:1990758 mitotic sister chromatid biorientation
IGI
PMID:22056777
Cdk1 promotes kinetochore bi-orientation and regulates Cdc20...
KEEP AS NON CORE
Summary: Liang et al.: Cdk1 promotes kinetochore bi-orientation during recovery from SAC arrest by restraining premature spindle elongation.
Reason: Real mitotic function of Cdk1 under checkpoint-recovery conditions; regulatory and context-specific, so non-core.
Supporting Evidence:
PMID:22056777
We show that Cdk1 promotes kinetochore bi-orientation during recovery by restraining premature spindle elongation thereby extinguishing SAC signalling.
GO:2001033 negative regulation of double-strand break repair via nonhomologous end joining
IMP
PMID:19699692
Regulation of repair choice: Cdk1 suppresses recruitment of ...
KEEP AS NON CORE
Summary: Zhang et al.: NHEJ is repressed in G2 through CDK1-dependent end resection and reduced association of Ku/Dnl4-Lif1 with breaks.
Reason: Substrate-mediated downstream process: Cdc28 phosphorylates the named substrate and thereby controls the process, but this is one of dozens of such outputs rather than a core function of the kinase, which is to gate the cell-cycle transitions.
Supporting Evidence:
PMID:19699692
Repression of NHEJ at G(2) is achieved by efficient end resection and by the reduced association of core NHEJ proteins with DNA breaks, both of which depend on the CDK1 activity.

Core Functions

Cyclin-dependent Ser/Thr protein kinase activity of the cyclin-Cdc28-Cks1 holoenzyme: with the G1 cyclins Cln1-3 it executes Start (the G1/S transition) by phosphorylating Whi5, Sic1, Far1 and Ste5, committing the cell to a new division cycle and triggering the SBF/MBF transcriptional program.

Supporting Evidence:
  • PMID:2142620
    Cln proteins are an essential component of the active protein kinase complex required for the G1 to S transition
  • PMID:7002718
    Each mutation produces stage-specific arrest of cell division at start, the same point where mating pheromone interrupts division.

S-phase CDK activity of Clb5/6-Cdc28: phosphorylates replication factors (Sld2/Sld3, Dpb2, Cdc6, ORC, Mcm10) to fire replication origins and prevent re-licensing, and phosphorylates Sic1 in a Cks1-dependent cascade to complete its destruction at the onset of S phase.

Supporting Evidence:
  • PMID:4580573
    Mutations in three genes (cdc 4, 7, and 28) appear to block a precondition for DNA synthesis
  • PMID:14747467
    The appearance of that species in vivo is dependent upon the Cdc28 cyclin-dependent protein kinase (CDK), which can directly phosphorylate Dpb2 in vitro.

Mitotic CDK activity of Clb1-4-Cdc28, activated by Tyr19 dephosphorylation: drives spindle pole body separation and spindle assembly (stabilising Cin8/Kip1/Ase1 and phosphorylating Ase1, Kar9, Ipl1), securin/separase control, and the G2/M transition, while phosphorylating Cdh1 and Swe1 to prevent premature mitotic exit.

Supporting Evidence:
  • PMID:2165600
    We present evidence that the Cdc28 protein kinase is also required for mitosis and that this function is executed in the G2 interval of the cell cycle.
  • PMID:8887667
    These results imply that dephosphorylation of Tyr-19 is required for the segregation of SPBs.

Meiotic CDK activity: Clb5/6-Cdc28 drives premeiotic DNA replication and initiates recombination by phosphorylating Mer2, and Clb-Cdc28 is required for pachytene exit, SPB separation before meiosis II and the meiotic divisions.

Supporting Evidence:
  • PMID:16814718
    the budding yeast cyclin-dependent kinase Cdc28 directly regulates the formation of the DNA double-strand breaks that initiate recombination by phosphorylating the Mer2/Rec107 protein
  • PMID:12783856
    Early inhibition of analog-sensitive cdc28-as1 blocked DNA replication, revealing a previously undetected role for Cdc28.

References

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Suggested Questions for Experts

Q: Of the several hundred in vitro Cdc28 substrates, which individual phosphorylation events are rate-limiting for ordering specific cell-cycle transitions, versus collectively contributing through cyclin-specific activity thresholds?

Q: Is the reported cell-cycle-independent role of Cdc28 at basal-transcription machinery (CTD-Ser5 phosphorylation with Kin28, RNAPII association) a direct catalytic function of a distinct Cdc28 pool, and which cyclin, if any, targets it there?

Q: For substrates where Cdc28 phosphorylation controls localisation (Swi5, Fus2, Ipl1, Rnr2, Cac1), should annotations be recorded as regulation of localisation rather than participation in the localisation process itself?

Suggested Experiments

Experiment: Combine cdc28-as1 chemical-genetic inhibition with cyclin docking-motif mutants (LP, RxL/NLxxxL, LxF) in synchronised cultures and quantitative phosphoproteomics to assign each substrate to a cyclin-Cdc28 complex and a cell-cycle window.

Hypothesis: Cyclin-specific docking, not total CDK activity, determines which substrates are phosphorylated at each transition.

Type: chemical genetics + phosphoproteomics

Experiment: Compare ChIP-seq occupancy and CTD-Ser5 phosphorylation at Cdc28-bound genes in cdc28-as1 cells with and without inhibitor, and in kinase-dead versus wild-type Cdc28 tethered to the PMA1 promoter, to separate catalytic from scaffolding contributions.

Hypothesis: A chromatin-associated Cdc28 pool at basal promoters acts catalytically on the RNAPII CTD or Kin28 rather than structurally.

Type: ChIP-seq / chemical genetics

Deep Research

Falcon

(CDC28-deep-research-falcon.md)

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Notes

(CDC28-notes.md)

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