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Showing CENPUMLF1IP is a alias.

CENPU

Centromere protein U · UniProt Q71F23

Length
418 aa
Mass
47.5 kDa
Annotated
2026-06-09
26 papers in source corpus 15 papers cited in narrative 15 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

CENPU (PBIP1/MLF1IP/CENP-50/CENP-U) is a constitutive inner-kinetochore protein of the CENP-O/CCAN class that couples centromere architecture to mitotic kinase signaling and kinetochore–microtubule attachment (PMID:16287847, PMID:34551298). Its centromere localization depends on CENP-H/CENP-I and on the CENP-P/CENP-Q subunits of the CENP-O complex, and CENPU loss causes prolonged mitosis and premature sister chromatid separation under spindle-checkpoint stress (PMID:16287847, PMID:34551298). CENPU forms a mutually stabilizing complex with CENP-Q, binds microtubules directly, and cooperates with Hec1/NDC80 to support kinetochore–microtubule attachment, an interaction tuned by Aurora B phosphorylation that drives error correction (PMID:21056971, PMID:21454580). CENPU is a principal kinetochore receptor for PLK1: CDK1 priming followed by PLK1 self-phosphorylation generates a phospho-Thr78 docking motif that binds the PLK1 polo-box domain, recruiting PLK1 to interphase and mitotic kinetochores, where it acts redundantly with BUB1 (PMID:17081991, PMID:33248027, PMID:34551298). PLK1 then phosphorylates CENP-Q within the ternary complex at multiple sites, delocalizing the CENPU–CENP-Q complex and driving its degradation, with both timely localization and removal being required for faithful chromosome segregation (PMID:21454580, PMID:25670858). Genetic ablation in mice is embryonic-lethal by E6.5 from epiblast degeneration, establishing an essential developmental role (PMID:23724000). Beyond its mitotic function, CENPU has been implicated in cancer through stabilization of partner proteins, including E2F6-dependent control of the E2F1–CENPU feedback loop in hepatocellular carcinoma and COX-2 stabilization driving angiogenesis in triple-negative breast cancer (PMID:31705927, PMID:35844791).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2004 Medium

    Before any kinetochore role was known, CENPU was first isolated as an MLF1-associating nuclear/cytoplasmic protein, providing the initial molecular handle on the gene.

    Evidence Yeast two-hybrid and pulldown with co-localization in human cells

    PMID:15116101

    Open questions at the time
    • Functional consequence of the MLF1 interaction was not characterized
    • No link to kinetochore biology established at this stage
  2. 2005 High

    Established CENPU as a constitutive centromere component whose localization depends on CENP-H/CENP-I and is required for recovery from spindle damage, defining its inner-kinetochore role.

    Evidence DT40 knockout, reciprocal Co-IP, and immunofluorescence

    PMID:16287847

    Open questions at the time
    • Molecular basis of the mitotic recovery defect not defined
    • Direct microtubule or kinase connections not yet known
  3. 2006 High

    Resolved how a mitotic kinase is targeted to kinetochores: PLK1 phosphorylates CENPU at Thr78 to create a self-tethering polo-box-binding motif, identifying CENPU as a PLK1 receptor.

    Evidence In vitro kinase assays, T78 mutagenesis, PBD-binding and live-cell imaging

    PMID:17081991

    Open questions at the time
    • Did not define the priming kinase upstream of PLK1
    • Mechanism of T78-dependent degradation left unresolved
  4. 2010 High

    Showed CENPU directly binds microtubules and cooperates with Hec1/NDC80, and that Aurora B phosphorylation weakens this attachment, embedding CENPU in error correction.

    Evidence Co-IP, in vitro microtubule-binding and Aurora B kinase assays, shRNA phenotyping

    PMID:21056971

    Open questions at the time
    • Aurora B phospho-sites on CENPU not exhaustively mapped
    • Quantitative contribution of CENPU vs NDC80 to attachment unclear
  5. 2011 High

    Defined the CENPU–CENP-Q complex as a mutually stabilizing unit and identified a PLK1–CENPU–CENP-Q ternary complex whose PLK1-dependent phosphorylation drives delocalization.

    Evidence Co-IP, in vitro PLK1 kinase assay, mutagenesis, immunofluorescence

    PMID:21454580

    Open questions at the time
    • Specific CENP-Q phosphosites not yet enumerated
    • Fate of delocalized complex not determined
  6. 2015 High

    Mapped nine PLK1 phosphosites on CENP-Q and demonstrated that both timely localization and PLK1-driven delocalization/degradation of the CENPU–CENP-Q complex are required for segregation.

    Evidence In vitro kinase assay, 9A/9D-E mutants, chromatin fractionation, segregation assays

    PMID:25670858

    Open questions at the time
    • Degradation machinery for cytosolic CENPU/CENP-Q not identified
    • How localization timing is set in interphase unclear
  7. 2020 High

    Defined the recruitment logic of PLK1 to CENPU: CDK1 priming followed by PLK1 itself generates docking motifs, placing CENPU alongside BUB1 as one of two main kinetochore PLK1 receptors.

    Evidence Ectopic localization, in vitro reconstitution, kinetochore localization and mutagenesis

    PMID:33248027

    Open questions at the time
    • Number and role of PLK1-docking sites later contested (see 2025 preprint)
    • Stoichiometry of PLK1 at the kinetochore not resolved
  8. 2021 High

    Established functional redundancy: BUB1 and CENP-U redundantly recruit PLK1 to stabilize attachments, with CENP-U dependent on CENP-P/CENP-Q, and showed human CENP-O does not control Aurora B localization unlike yeast.

    Evidence Double depletion epistasis, chemical-genetic sensitization, immunofluorescence

    PMID:34551298

    Open questions at the time
    • Why two parallel PLK1 receptors exist mechanistically unclear
    • Spatial division of labor between BUB1 and CENP-U not mapped
  9. 2021 High

    In budding yeast, showed the CENPU ortholog Ame1 is controlled by CDK1-activated phospho-degrons recognized by SCF-Cdc4, with Mis12 binding shielding kinetochore-bound Ame1 from degradation.

    Evidence Phosphorylation mapping, ubiquitination assays, cdc4 genetics in yeast

    PMID:34308839

    Open questions at the time
    • Conservation of degron regulation to human CENPU not tested
    • Identity of relevant human E3 not established
  10. 2013 Medium

    Demonstrated CENPU is essential for early mammalian development, with knockout embryos dying at E6.5 from epiblast degeneration.

    Evidence Germline knockout mouse, in situ hybridization, immune phenotyping

    PMID:23724000

    Open questions at the time
    • Molecular cause of epiblast death not defined
    • Link between lethality and kinetochore function not directly shown
  11. 2019 Medium

    Extended CENPU function to oncogenic signaling, showing it stabilizes COX-2 by blocking its ubiquitination to drive angiogenesis in triple-negative breast cancer.

    Evidence shRNA, ubiquitination assay, tube formation, xenograft model

    PMID:31705927

    Open questions at the time
    • Direct CENPU–COX-2 interaction not fully validated
    • Relationship to kinetochore role unknown
  12. 2022 Medium

    Identified an E2F1–CENPU positive feedback loop in which CENPU degrades E2F6 to de-repress E2F1, accelerating G1/S transition in hepatocellular carcinoma.

    Evidence Co-IP, ubiquitination assay, ChIP, knockdown/overexpression

    PMID:35844791

    Open questions at the time
    • Mechanism by which CENPU promotes E2F6 ubiquitination unclear
    • Generality beyond hepatocellular carcinoma untested
  13. 2025 Medium

    Reopened the architecture of PLK1 docking, with biophysical analysis arguing for a single high-affinity master motif engaging multiple PBD pockets rather than PBD dimerization.

    Evidence Biochemical/biophysical binding assays and structural modelling (preprint)

    PMID:bio_10.1101_2025.03.28.645803

    Open questions at the time
    • Preprint not peer-reviewed
    • Conflicts with earlier dimerization model and awaits reconciliation
  14. 2025 Medium

    Provided structural basis for Mis12-mediated connection: a Dsn1 auto-inhibitory segment occludes the Ame1/CENP-U binding site and Aurora B phosphorylation would relieve it to strengthen inner–outer kinetochore links.

    Evidence Cryo-EM with biochemical and genetic validation in yeast (preprint)

    PMID:bio_10.1101_2025.06.03.657598

    Open questions at the time
    • Preprint not peer-reviewed
    • Demonstrated for yeast Ame1; human relevance untested
  15. 2025 Low

    Suggested a further cancer mechanism in which CENPU stabilizes furin to enhance proNGF-to-NGF processing and breast cancer stem cell properties.

    Evidence Co-IP, furin stability western blots, ELISA, proteomics, mammosphere and xenograft assays

    PMID:41041859

    Open questions at the time
    • Single lab, not independently replicated
    • Mechanism of how CENPU blocks lysosomal furin degradation undefined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How CENPU's essential mitotic/kinetochore role mechanistically connects to its reported cancer-associated protein-stabilization activities and to its embryonic-lethal phenotype remains unresolved.
  • No unifying mechanism links kinetochore scaffolding to COX-2/E2F6/furin stabilization
  • Molecular driver of epiblast lethality not identified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 2 GO:0140096 catalytic activity, acting on a protein 2 GO:0008092 cytoskeletal protein binding 1
Localization
GO:0005634 nucleus 2 GO:0005694 chromosome 2
Pathway
R-HSA-1640170 Cell Cycle 3
Complex memberships
CENP-O complex (CENP-O/P/Q/U)CENPU–CENP-Q complexkinetochore (CCAN)

Evidence

Reading pass · 15 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 PLK1 phosphorylates PBIP1 (CENPU) at Thr-78, creating a self-tethering phosphopeptide motif that specifically binds the polo-box domain (PBD) of PLK1 (but not PLK2 or PLK3), recruiting PLK1 to interphase and mitotic kinetochores. Later in mitosis, PLK1 also induces PBIP1 degradation in a T78-dependent manner, enabling PLK1 to interact with other kinetochore components required for proper chromosome segregation. In vitro kinase assays, mutagenesis of T78, PBD-binding assays, live-cell imaging, spindle checkpoint assays Molecular cell High 17081991
2005 CENP-50 (CENPU) is a constitutive centromere component that colocalizes with CENP-A and CENP-H throughout the cell cycle; its centromere localization depends on CENP-H and CENP-I. CENP-50 physically interacts with the CENP-H/CENP-I complex (shown by co-immunoprecipitation). Loss of CENP-50 causes prolonged mitosis and premature sister chromatid separation upon spindle checkpoint activation, indicating a role in recovery from spindle damage. Knockout in chicken DT40 cells, co-immunoprecipitation, immunofluorescence Molecular and cellular biology High 16287847
2010 CENPU directly interacts with Hec1 (NDC80) at kinetochores. CENPU itself binds microtubules directly in vitro and displays cooperative microtubule-binding activity with Hec1. Aurora B phosphorylates CENPU, and this phosphorylation reduces kinetochore–microtubule interaction, contributing to Aurora B's error-correction function. shRNA-mediated knockdown of CENPU impairs kinetochore–microtubule attachment. Co-immunoprecipitation, in vitro microtubule-binding assays, in vitro Aurora B kinase assay, shRNA knockdown with mitotic phenotype analysis The Journal of biological chemistry High 21056971
2011 PBIP1 (CENPU) directly interacts with CENP-Q, and this interaction is mutually required for their stability and centromere localization. PLK1 forms a ternary complex with PBIP1 and CENP-Q through the self-generated p-T78 motif on PBIP1, and PLK1-dependent phosphorylation of CENP-Q within this complex leads to delocalization of the PBIP1–CENP-Q complex from mitotic centromeres. Co-immunoprecipitation, in vitro PLK1 kinase assay, mutagenesis, immunofluorescence localization The Journal of biological chemistry High 21454580
2015 PLK1 phosphorylates the CENP-Q subunit of the PBIP1–CENP-Q complex at multiple sites; phosphorylation of nine sites drives delocalization of the complex from kinetochores. Phospho-mimetic (9D/E) mutations prevent localization to interphase prekinetochores, while phospho-null (9A) mutations prolong kinetochore residence. Both mutants impair proper chromosome segregation, demonstrating that timely localization and delocalization of the PBIP1–CENP-Q complex are both critical for mitotic progression. PLK1-dependent delocalization indirectly leads to cytosolic degradation of PBIP1 and CENP-Q. In vitro PLK1 kinase assay, mutagenesis (9A and 9D/E), chromatin fractionation, immunofluorescence, chromosome segregation assay The Journal of biological chemistry High 25670858
2020 BUB1 (outer kinetochore) and CENP-U (inner kinetochore) are the main PLK1 receptors at kinetochores. Both share a constellation of sequence motifs (a PP2A-docking motif and two PLK1-docking sites). PLK1 recruitment to CENP-U requires priming phosphorylation by CDK1 followed by PLK1 itself. The two PLK1-docking sites on CENP-U promote PLK1 dimerization. This was demonstrated by ectopic localization assays, in vitro reconstitution, and kinetochore localization studies. Ectopic localization assay, in vitro reconstitution of PLK1–CENP-U interaction, kinetochore localization studies, mutagenesis Molecular cell High 33248027
2021 BUB1 and CENP-U redundantly recruit PLK1 to kinetochores to stabilize kinetochore–microtubule attachments. CENP-U is recruited to kinetochores by the CENP-P and CENP-Q subunits of the CENP-O complex. Depletion of both BUB1 and CENP-U causes chromosome mis-segregation; depletion of either alone is insufficient, demonstrating functional redundancy. Cells depleted of BUB1 or CENP-U are sensitized to PLK1 inhibition but not Aurora B inhibition. Unlike its budding yeast homolog, the CENP-O complex does not regulate centromeric localization of Aurora B in human cells. Stable depletion by siRNA/shRNA, epistasis analysis (double depletion), chemical inhibitor sensitization assays, immunofluorescence Cell reports High 34551298
2004 CENPU (MLF1IP) was identified as a protein that specifically associates with MLF1 by yeast two-hybrid analysis and pulldown assays, and colocalizes with MLF1 in both nuclei and cytoplasm. The protein contains two bipartite and two classical nuclear localization signals, two nuclear receptor-binding motifs (LXXLL), two leucine zippers, and multiple potential phosphorylation sites. Yeast two-hybrid, pulldown assay, co-localization by immunofluorescence Oncogene Medium 15116101
2021 In budding yeast, Cdk1 phosphorylation of the CENPU ortholog Ame1 activates phospho-degrons that are recognized by the SCF-Cdc4 E3 ubiquitin ligase complex, targeting Ame1 for proteasomal degradation during M-phase. Binding of the Mtw1 (Mis12) complex shields the proximal phospho-degron, protecting kinetochore-bound Ame1 from degradation. This mechanism ensures efficient centromere-dependent kinetochore assembly. Comprehensive phosphorylation analysis of native CCAN subunits, biochemical ubiquitination assays, genetic assays in budding yeast (cdc4 mutants, overexpression toxicity) eLife High 34308839
2019 CENPU promotes angiogenesis in triple-negative breast cancer by inhibiting the ubiquitination and proteasomal degradation of COX-2, leading to increased COX-2-p-ERK-HIF-1α-VEGFA signaling. CENPU knockdown reduced VEGFA production, tube formation by endothelial cells, and tumor microvessel density in xenograft models. shRNA knockdown, ubiquitination assay, western blot (COX-2 protein stability), endothelial tube formation assay, xenograft mouse model Cancer letters Medium 31705927
2022 CENPU physically interacts with the transcription factor E2F6 and promotes its ubiquitin-mediated degradation, thereby de-repressing E2F1 transcription. E2F1 in turn binds the CENPU promoter to increase CENPU transcription, forming a positive feedback loop that accelerates G1/S transition in hepatocellular carcinoma cells. Co-immunoprecipitation, ubiquitination assay, chromatin immunoprecipitation (E2F1 binding to CENPU promoter), western blot, knockdown and overexpression International journal of biological sciences Medium 35844791
2020 MLF1-IP (CENPU) KO mice die by embryonic day 6.5 due to degeneration of epiblasts, establishing an essential role in early embryonic development. Heterozygous KO mice are viable and fertile with no apparent immune system defects, indicating that ~50% expression is sufficient for normal postnatal development. Germline knockout mouse (homozygous lethal, heterozygous viable), in situ hybridization, immune phenotyping PloS one Medium 23724000
2025 CENPU promotes furin activity by inhibiting lysosomal degradation of furin in triple-negative breast cancer cells. Furin, a precursor-processing enzyme, converts proNGF to NGF, which promotes breast cancer stem cell properties. Co-immunoprecipitation experiments demonstrated association between CENPU, furin, and NGF/proNGF. A furin inhibitor suppressed CENPU-promoted tumor growth in xenograft models. Co-immunoprecipitation, western blot (furin protein stability), ELISA (NGF), 4D-DIA quantitative proteomics, mammosphere formation, xenograft mouse model International journal of molecular medicine Low 41041859
2025 CENP-U contains a single high-affinity master PLK1-docking motif that is generated by initial CDK1 priming and subsequent PLK1 phosphorylation. Biochemical and biophysical analyses showed this motif forms extensive interactions with multiple pockets on the PBD surface. Evidence did not support a requirement for PBD dimerization in CENP-U-mediated PLK1 docking. Biochemical binding assays, biophysical measurements, structural modelling bioRxivpreprint Medium bio_10.1101_2025.03.28.645803
2025 In budding yeast, an N-terminal auto-inhibitory segment of Dsn1 (Mis12 complex subunit) occludes binding sites for both CENP-C/Mif2 and CENP-U/Ame1 on the Mis12 complex head domain. Aurora B/Ipl1 phosphorylation of this auto-inhibitory segment would release auto-inhibition and strengthen inner–outer kinetochore connections involving CENP-U/Ame1. This was established by cryo-EM structure combined with biochemical and genetic experiments. Cryo-EM, biochemical binding assays, genetic assays in budding yeast bioRxivpreprint Medium bio_10.1101_2025.06.03.657598

Source papers

Stage 0 corpus · 26 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Self-regulated Plk1 recruitment to kinetochores by the Plk1-PBIP1 interaction is critical for proper chromosome segregation. Molecular cell 227 17081991
2020 BUB1 and CENP-U, Primed by CDK1, Are the Main PLK1 Kinetochore Receptors in Mitosis. Molecular cell 91 33248027
2005 The constitutive centromere component CENP-50 is required for recovery from spindle damage. Molecular and cellular biology 66 16287847
2004 cDNA cloning and characterization of a novel gene encoding the MLF1-interacting protein MLF1IP. Oncogene 57 15116101
2010 CENP-U cooperates with Hec1 to orchestrate kinetochore-microtubule attachment. The Journal of biological chemistry 56 21056971
2011 Mammalian polo-like kinase 1-dependent regulation of the PBIP1-CENP-Q complex at kinetochores. The Journal of biological chemistry 44 21454580
2021 Bub1 and CENP-U redundantly recruit Plk1 to stabilize kinetochore-microtubule attachments and ensure accurate chromosome segregation. Cell reports 36 34551298
2019 Centromere protein U (CENPU) enhances angiogenesis in triple-negative breast cancer by inhibiting ubiquitin-proteasomal degradation of COX-2. Cancer letters 32 31705927
2008 Self-regulated mechanism of Plk1 localization to kinetochores: lessons from the Plk1-PBIP1 interaction. Cell division 32 18215321
2022 A positive feedback loop of CENPU/E2F6/E2F1 facilitates proliferation and metastasis via ubiquitination of E2F6 in hepatocellular carcinoma. International journal of biological sciences 29 35844791
2005 Regulation of myeloid leukemia factor-1 interacting protein (MLF1IP) expression in glioblastoma. Brain research 27 15893739
2019 Short communication: A splice site mutation in CENPU is associated with recessive embryonic lethality in Holstein cattle. Journal of dairy science 20 31733857
2015 Mammalian Polo-like kinase 1 (Plk1) promotes proper chromosome segregation by phosphorylating and delocalizing the PBIP1·CENP-Q complex from kinetochores. The Journal of biological chemistry 19 25670858
2022 The centromere-associated protein CENPU promotes cell proliferation, migration, and invasiveness in lung adenocarcinoma. Cancer letters 16 35176420
2019 Reduced CENPU expression inhibits lung adenocarcinoma cell proliferation and migration through PI3K/AKT signaling. Bioscience, biotechnology, and biochemistry 16 30849291
2021 Centromere protein U (CENPU) promotes gastric cancer cell proliferation and glycolysis by regulating high mobility group box 2 (HMGB2). Bioengineered 10 34872447
2017 MLF1IP promotes cells proliferation and apoptosis by regulating CyclinD1 in breast cancer. International journal of clinical and experimental pathology 10 31966511
2017 MLF1IP promotes normal erythroid proliferation and is involved in the pathogenesis of polycythemia vera. FEBS letters 9 28173615
2022 Knockdown of CENPU inhibits cervical cancer cell migration and stemness through the FOXM1/Wnt/β-catenin pathway. Tissue & cell 6 36608638
2013 Investigation of tissue-specific expression and functions of MLF1-IP during development and in the immune system. PloS one 6 23724000
2021 Cdc4 phospho-degrons allow differential regulation of Ame1CENP-U protein stability across the cell cycle. eLife 5 34308839
2022 Overexpression of MLF1IP promotes colorectal cancer cell proliferation through BRCA1/AKT/p27 signaling pathway. Cellular signalling 4 35122991
2021 Mitosis-related gene CENP-U as a potential biomarker in malignancy. Annals of translational medicine 4 35071438
2020 CENP-50 is required for papilloma development in the two-stage skin carcinogenesis model. Cancer science 3 32535988
2025 DPHC from Alpinia officinarum Hance specifically modulates the function of CENPU in the cell cycle and apoptosis to ameliorate hepatocellular carcinoma. Journal of ethnopharmacology 1 40058474
2025 CENPU promotes tumorigenesis and stem cell properties in triple‑negative breast cancer by suppressing lysosomal furin degradation. International journal of molecular medicine 1 41041859

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