Affinage

NUMA1

Nuclear mitotic apparatus protein 1 · UniProt Q14980

Length
2115 aa
Mass
238.3 kDa
Annotated
2026-04-29
100 papers in source corpus 51 papers cited in narrative 49 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

NuMA is a large coiled-coil protein that functions as a master organizer of mitotic spindle architecture and a regulator of nuclear processes during interphase. During mitosis, CDK1 phosphorylation triggers NuMA's release from the nucleus; NuMA is then transported poleward by cytoplasmic dynein–dynactin, where it self-assembles into an insoluble cross-linking matrix that focuses microtubule minus-ends at spindle poles—a function essential for centrosome–spindle attachment under tension and for acentrosomal spindle bipolarization (PMID:8898198, PMID:19255246, PMID:31782546). At the cell cortex, NuMA is recruited by LGN/Gαi and directly by phosphoinositides to cluster dynein–dynactin force generators that pull on astral microtubules for spindle orientation, with cortical NuMA levels dynamically tuned by CDK1, Aurora-A, and Plk1 phosphorylation and opposed by PP2A dephosphorylation (PMID:15537540, PMID:24996901, PMID:23921553, PMID:26832443). In interphase, NuMA resides in the nuclear matrix where it contributes to p53 target-gene selectivity by recruiting CDK8/Mediator, facilitates rDNA transcription through RNA Pol I association, and regulates DNA double-strand break repair by controlling SNF2h-mediated chromatin remodeling and 53BP1 dynamics (PMID:23589328, PMID:28981686, PMID:24753406, PMID:30812030).

Mechanistic history

Synthesis pass · year-by-year structured walk · 17 steps
  1. 1992 High

    Cloning NuMA established it as a large nuclear protein with a tripartite domain architecture—globular heads flanking a long coiled-coil rod—that dynamically redistributes between the nucleus and spindle poles during mitosis, framing the central question of how a nuclear protein reorganizes into a mitotic structural element.

    Evidence cDNA cloning, sequence analysis, and double immunofluorescence with anti-lamin antibodies in human cells

    PMID:1541630 PMID:1541636

    Open questions at the time
    • No functional data; structure-function relationship of domains unknown
    • No interacting partners identified
  2. 1995 High

    Immunodepletion and reconstitution experiments demonstrated that NuMA is essential for organizing microtubule asters and that its C-terminal tail mediates microtubule bundling, establishing NuMA as a direct structural organizer of mitotic spindle poles rather than a passive nuclear scaffold.

    Evidence Cell-free mitotic aster assembly, immunodepletion with recombinant protein rescue, EM of purified protein, CDK1 phosphosite mutagenesis

    PMID:7593190 PMID:7769006 PMID:7781599

    Open questions at the time
    • Motor-dependent transport mechanism to poles not yet defined
    • Regulation of NuMA–MT interaction beyond CDK1 unknown
  3. 1999 High

    EM and self-assembly studies revealed that NuMA oligomerizes into multiarm structures through C-terminal interactions and forms an insoluble matrix at spindle poles independent of microtubules, explaining how NuMA provides a structural scaffold that anchors MT minus-ends.

    Evidence In vitro assembly with EM, biochemical fractionation, immunogold EM of spindle poles

    PMID:10075938 PMID:10098933

    Open questions at the time
    • Regulation of assembly/disassembly cycle unclear
    • How the matrix integrates with pericentriolar material not resolved
  4. 2000 High

    Live imaging and biochemistry established that dynein–dynactin actively transports NuMA to spindle poles after nuclear envelope breakdown, forming a mitosis-specific complex whose disruption blocks pole assembly—linking NuMA's motor-dependent transport to its structural role.

    Evidence Live GFP imaging, immunoprecipitation, gel filtration, dynamitin overexpression in mammalian cells

    PMID:10811826

    Open questions at the time
    • Which NuMA domains contact dynein LIC not yet mapped
    • Regulation of complex formation/dissociation unclear
  5. 2004 High

    Discovery that LGN acts as a conformational switch binding both NuMA and Gαi resolved how NuMA reaches the cell cortex for spindle orientation, establishing the LGN/Gαi/NuMA ternary complex as the cortical force-generating pathway, with CDK1-mediated cyclin B degradation releasing NuMA from dynein at anaphase.

    Evidence FRET biosensor, yeast two-hybrid, RNAi, live imaging; Xenopus egg extract with non-degradable cyclin B

    PMID:14710193 PMID:15537540

    Open questions at the time
    • Whether LGN and direct MT binding are mutually exclusive in vivo not tested
    • Cortical force magnitude not measured
  6. 2009 High

    Conditional NuMA knockout in mice proved NuMA is essential in vivo for tethering spindle microtubules to centrosomes under tension, resolving the question of whether NuMA has a unique function beyond what centrosomes and other crosslinkers provide.

    Evidence Conditional loss-of-function allele in mouse primary cells, live imaging

    PMID:19255246

    Open questions at the time
    • Whether NuMA's tethering and cortical functions are independently essential in vivo not separated
    • Tissue-specific requirements not fully explored
  7. 2011 High

    Crystal structures of LGN bound to NuMA versus mInsc revealed mutually exclusive binding to the same TPR surface, establishing that Par3/mInsc/LGN and NuMA/LGN/Gαi act sequentially rather than simultaneously in orienting the spindle.

    Evidence X-ray crystallography with in vitro competition binding assays

    PMID:21816348

    Open questions at the time
    • Temporal handoff mechanism from mInsc to NuMA in vivo not directly visualized
    • Whether additional factors modulate the switch unknown
  8. 2013 High

    Multiple groups established that CDK1 phosphorylation of NuMA T2055 suppresses cortical NuMA during metaphase while PP2A counteracts this, and that a C-terminal lipid-binding domain mediates LGN-independent cortical association in anaphase—revealing a phospho-switch coupling mitotic progression to spindle positioning forces.

    Evidence In vitro kinase assays, phosphomutant rescue, lipid-binding assays, live imaging in human cells and keratinocytes

    PMID:23921553 PMID:24109598 PMID:24371089

    Open questions at the time
    • Precise PIP species specificity not determined
    • How cortical NuMA domains are spatially restricted to the cell equator at anaphase unclear
  9. 2014 High

    Direct binding of NuMA to phosphoinositides (PIP/PIP2) was demonstrated, and PIP depletion abolished cortical NuMA, establishing phosphoinositides as the primary cortical receptor for NuMA during anaphase when LGN/Gαi are dispensable.

    Evidence In vitro lipid-binding assay, chemical/enzymatic PIP depletion, immunofluorescence, siRNA rescue

    PMID:24996901

    Open questions at the time
    • Whether PIP2 clustering or local PIP2 generation provides spatial specificity unknown
    • Structural basis of NuMA–PIP2 interaction not resolved
  10. 2016 High

    Aurora-A phosphorylation was shown to control NuMA's dynamic exchange between spindle poles and the cortex without affecting MT or LGN binding affinity, and in vivo mutagenesis of NuMA's MT-binding domain in mouse skin caused spindle orientation defects and neonatal lethality, demonstrating NuMA's direct MT engagement is essential for tissue morphogenesis.

    Evidence In vitro kinase assay, FRAP, phosphomutants; conditional MT-binding domain knockout in mouse epidermis

    PMID:26765568 PMID:26832443

    Open questions at the time
    • How Aurora-A phosphorylation alters NuMA mobility mechanistically (conformational change?) not defined
    • Relative contributions of dynein-mediated vs. direct MT-binding functions in different tissues not quantified
  11. 2017 High

    Quantitative imaging and structural studies revealed that NuMA localizes to MT minus-ends independently of dynein and recruits dynactin there, while a crystal structure showed Importin-α/β masks NuMA's MT-binding region with RanGTP releasing the inhibition—establishing NuMA as a bona fide minus-end cargo adaptor regulated by the Ran gradient.

    Evidence Laser ablation, domain deletion rescue, X-ray crystallography of Importin-α/NuMA complex, in vitro MT-binding assay

    PMID:28939615 PMID:29185983

    Open questions at the time
    • Whether Ran gradient regulation operates at the cortex as well as at poles unclear
    • Minus-end recognition mechanism (structural feature of minus-end) not identified
  12. 2018 High

    Optogenetic cortical targeting showed that NuMA—not dynein alone—is sufficient to generate spindle-pulling forces by assembling cortical focal clusters of dynein–dynactin; Plk1 was identified as an additional kinase phosphorylating NuMA to tune cortical levels, adding a third kinase input to the cortical regulatory circuit.

    Evidence Light-induced cortical targeting with AID depletion and domain dissection; Plk1 in vitro kinase assay with pharmacological inhibition and FRAP

    PMID:29848445 PMID:30456393

    Open questions at the time
    • How CDK1, Aurora-A, and Plk1 phosphorylation events are integrated on the same NuMA molecule not resolved
    • Stoichiometry of NuMA within cortical clusters unknown
  13. 2020 High

    Structural determination of NuMA–LIC1/LIC2 interfaces identified two distinct dynein light-intermediate-chain-binding sites (hook domain and CC1-box), both essential for spindle positioning, providing the molecular basis for NuMA's function as a dynein cargo adaptor.

    Evidence X-ray crystallography of NuMA–LIC1/2 interfaces, mutagenesis, functional cell division assay

    PMID:32413290

    Open questions at the time
    • Whether the two LIC-binding sites engage one or two dynein complexes simultaneously unknown
    • Regulation of LIC-binding site accessibility not addressed
  14. 2013 Medium

    Discovery of NuMA's interphase nuclear functions revealed it binds p53 and recruits CDK8/Mediator selectively to p21 but not PUMA promoters, establishing NuMA as a transcriptional cofactor that confers target-gene selectivity to the p53 response.

    Evidence Co-immunoprecipitation, ChIP, siRNA knockdown with selective gene expression readout

    PMID:23589328

    Open questions at the time
    • Structural basis for selective promoter targeting unknown
    • Whether NuMA acts on other transcription factors besides p53 not tested
    • Independent replication needed
  15. 2014 Medium

    NuMA was found to accumulate at DNA damage sites in a PARylation-dependent manner, interact with SNF2h to promote chromatin decompaction and homologous recombination, linking NuMA's nuclear matrix role to active DNA repair.

    Evidence Co-IP, FRAP, laser microirradiation, siRNA with chromosomal DSB repair assays

    PMID:24753406

    Open questions at the time
    • Whether NuMA is directly PARylated at breaks or recruited via PAR binding unclear
    • Relationship to NuMA's interaction with tankyrase-1 at spindle poles not addressed
  16. 2019 Medium

    NuMA was shown to sequester 53BP1 in undamaged nuclei and limit its access to breaks, with NuMA depletion altering PARP inhibitor sensitivity in BRCA1-null cells, revealing NuMA as a regulator of DNA repair pathway choice.

    Evidence Co-IP, FRAP, NHEJ assay, class-switch recombination assay, PARP inhibitor sensitivity

    PMID:30812030

    Open questions at the time
    • Mechanism by which NuMA restrains 53BP1 mobility not structurally defined
    • In vivo relevance for tumor suppression not tested
  17. 2021 Medium

    NuMA was shown to undergo liquid–liquid phase separation at mitotic entry, regulated by Aurora-A, concentrating tubulins and the MT depolymerase Kif2A at poles to promote poleward flux—providing a biophysical framework for how NuMA's self-assembly creates a functional spindle pole compartment.

    Evidence In vitro LLPS reconstitution, live cell condensate imaging, phosphomimetic/phosphodead mutants, siRNA

    PMID:34887424

    Open questions at the time
    • Whether LLPS and the previously described insoluble matrix represent the same or distinct states unclear
    • Contribution of LLPS versus oligomeric cross-linking to pole integrity not separated
    • Independent replication of LLPS findings needed

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include how the three mitotic kinase inputs (CDK1, Aurora-A, Plk1) are integrated on a single NuMA molecule to produce graded cortical versus polar localization, the structural basis of NuMA's phosphoinositide binding and minus-end recognition, and the extent to which NuMA's interphase nuclear functions (p53 cofactor, rDNA transcription, DNA repair pathway choice) are mechanistically independent or coupled through a common scaffolding principle.
  • Integrated phospho-regulation model lacking
  • No structural data for NuMA–PIP2 or NuMA–minus-end interfaces
  • Whether nuclear and mitotic NuMA pools are regulated by shared or distinct mechanisms unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 6 GO:0005198 structural molecule activity 3 GO:0060090 molecular adaptor activity 3 GO:0008289 lipid binding 2 GO:0140110 transcription regulator activity 2
Localization
GO:0005815 microtubule organizing center 5 GO:0005886 plasma membrane 5 GO:0005634 nucleus 4 GO:0005856 cytoskeleton 4 GO:0005730 nucleolus 1
Pathway
R-HSA-1640170 Cell Cycle 8 R-HSA-162582 Signal Transduction 3 R-HSA-73894 DNA Repair 2 R-HSA-74160 Gene expression (Transcription) 2
Partners
AURKADCTN1DYNC1H1EPB41GNAI1GPSM2TNKSTP53
Complex memberships
LGN/Gαi/NuMANuMA–dynein–dynactinNuMA–tankyrase-1

Evidence

Reading pass · 49 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 NuMA forms a complex with cytoplasmic dynein and dynactin; immunodepletion of NuMA from frog egg extracts abolishes normal spindle pole assembly, producing chromatin-associated irregular microtubule arrays; a NuMA tail subdomain induces microtubule aster formation by mediating microtubule bundling. Immunodepletion from Xenopus egg extracts, Co-immunoprecipitation, in vitro aster assembly assay Cell High 8898198
2000 NuMA transport to spindle poles upon nuclear envelope breakdown is powered by cytoplasmic dynein and dynactin along microtubule minus ends; large cytoplasmic NuMA aggregates stream poleward in association with dynactin (Arp1) and dynein; immunoprecipitation and gel filtration demonstrate a reversible, mitosis-specific NuMA–dynein–dynactin complex; disruption of dynactin or dynein blocks NuMA translocation and spindle pole assembly. Live GFP imaging, immunoprecipitation, gel filtration, dynamitin overexpression, dynein antibody inhibition The Journal of cell biology High 10811826
1992 NuMA is a 236–238 kDa nuclear protein with a long central coiled-coil domain (~1,485 aa) flanked by globular N- and C-terminal domains; it dissociates from condensing chromosomes early in prophase before lamina disintegration and reassociates with telophase chromosomes before lamin accumulation, indicating a role in nuclear reformation. cDNA cloning, sequence analysis, double immunofluorescence with anti-NuMA and anti-lamin antibodies The Journal of cell biology High 1541630 1541636
1995 NuMA is required for organizing microtubules into aster-like mitotic arrays; immunodepletion of NuMA from mitotic cell extracts prevents aster formation, and addition of purified recombinant NuMA fully rescues aster assembly; NuMA is phosphorylated upon aster assembly and is only required in late stages of aster formation. Cell-free mitotic aster assembly assay, immunodepletion, recombinant protein rescue The Journal of cell biology High 7593190
1995 Mutation of the predicted CDK1 (p34cdc2) phosphorylation site at Thr2040 of NuMA (alone or combined with other site mutations) abolishes NuMA's ability to associate with spindle microtubules; instead the mutant concentrates at the plasma membrane, causing disorganized spindles, cytokinesis failure, and micronucleation. This demonstrates that mitosis-specific phosphorylation controls NuMA's spindle interaction. Site-directed mutagenesis, transient expression, immunofluorescence, flow cytometry Journal of cell science High 7769006
1993 Expression of NuMA lacking its globular head domain results in cytokinesis failure and micronuclei formation; expression of NuMA lacking the globular tail domain prevents nuclear targeting and spindle binding, causing micronucleation in daughter cells. Wild-type NuMA overexpression rescues nuclear assembly defects in RCC1 temperature-sensitive cells. NuMA function is required for terminal phases of chromosome separation and/or nuclear reassembly. Dominant-negative expression, temperature-sensitive mutant cell rescue, immunofluorescence The Journal of cell biology High 8432734
2004 LGN (mammalian Pins homolog) directly binds NuMA through its N-terminal domain and recruits NuMA to the cell cortex during mitosis; LGN behaves as a conformational switch—its closed state has N- and C-termini interacting, but NuMA or Gαi can open the switch, allowing simultaneous binding to both, resulting in their cortical localization. LGN–NuMA interaction is required for spindle oscillation. FRET biosensor, yeast two-hybrid, overexpression, RNAi, live imaging Cell High 15537540
2001 LGN (human Pins-related protein) binds the C-terminal tail of NuMA; LGN blocks NuMA-dependent stabilization and bundling of microtubules. In vitro assays show NuMA binds MTs directly, and the MT-binding domain overlaps by 10 aa with the LGN-binding domain, indicating steric exclusion explains LGN's inhibitory effect on NuMA spindle function. In vitro MT binding/stabilization assays, Xenopus egg extract aster assay, domain mapping Nature cell biology / Current biology High 11781568 12445386
1999 A 135 kDa nonerythroid isoform of protein 4.1R directly interacts with NuMA; minimal binding involves 4.1R exons 20–21 and NuMA residues 1788–1810; 4.1R and NuMA co-localize in interphase nuclei and redistribute to spindle poles, where 4.1R is part of a NuMA–dynein–dynactin complex during mitosis. Yeast two-hybrid, in vitro binding assay, co-immunoprecipitation, co-immunolocalization The Journal of cell biology Medium 10189366
2002 NuMA contains an RXXPDG motif that mediates direct binding to the ankyrin-repeat domain of tankyrase-1 and tankyrase-2 (poly-ADP-ribose polymerases); this interaction contributes to the known co-localization of tankyrase and NuMA at mitotic spindle poles. Yeast two-hybrid, in vitro binding, co-localization The Journal of biological chemistry Medium 12080061
2005 NuMA is a major acceptor of poly(ADP-ribosyl)ation (PARsylation) by tankyrase 1 in mitosis; tankyrase 1 and NuMA association increases at mitotic onset concomitant with NuMA PARsylation; siRNA knockdown of tankyrase 1 eliminates NuMA PARsylation; conversely, NuMA knockdown causes complete loss of tankyrase 1 from spindle poles. Co-immunoprecipitation, immunofluorescence, siRNA knockdown The Biochemical journal Medium 16076287
2009 Poly(ADP-ribose) (pADPr) made by PARP-5a/tankyrase-1 localizes to spindle poles; pADPr-coated beads trigger microtubule aster assembly in mitotic HeLa lysate; the rod domain of NuMA binds directly to pADPr chains, suggesting pADPr provides dynamic cross-linking at spindle poles by binding NuMA. In vitro pADPr bead aster assay, recombinant domain binding, immuno-EM Molecular biology of the cell Medium 19759176
1997 NuMA is phosphorylated by CDK1 (Cdc2 kinase) in a mitosis-specific manner as shown in vivo with 32P-labeling; phosphorylation shifts NuMA to a higher apparent molecular weight form; this modification coincides with NuMA's release from the nucleus and redistribution to the spindle. 32P metabolic labeling, phosphatase treatment, immunoblot in synchronized cells Journal of cell science Medium 9202389
2004 Cyclin B degradation at anaphase onset leads to NuMA dephosphorylation and its release from dynein, dynactin, and spindle poles; a non-degradable cyclin B (Δ90) keeps NuMA phosphorylated and stably associated with dynein/dynactin at persistent spindle poles that fail to disassemble. Xenopus egg extract, non-degradable cyclin B expression, co-immunoprecipitation, immunofluorescence EMBO reports Medium 14710193
2013 CDK1 phosphorylates NuMA at T2055 to negatively regulate its cortical localization during metaphase; PPP2CA (PP2A) phosphatase counteracts this; CDK1 inactivation in anaphase increases dephosphorylated cortical NuMA, enriching cortical dynein and driving spindle elongation. Thus, NuMA phosphorylation status by CDK1 couples mitotic progression with spindle behavior. In vitro kinase assay, phosphorylation-site mutagenesis, immunofluorescence, siRNA knockdown in human cells The EMBO journal High 23921553
2013 A lipid/membrane-binding domain at the C-terminus of NuMA mediates LGN/Gαi-independent cortical association during anaphase; this binding is inhibited by CDK1 phosphorylation during prophase and metaphase and activated upon CDK1 inactivation at anaphase; replacing endogenous NuMA with membrane-binding-deficient NuMA reduces anaphase cortical dynein and impairs chromosome separation. Domain mapping, phosphorylation-site mutagenesis, live cell imaging, siRNA rescue experiments, lipid-binding assays Molecular biology of the cell High 24371089
2014 NuMA directly associates with phosphoinositides PtdInsP (PIP) and PtdInsP2 (PIP2) in vitro; depletion of PIP/PIP2 by chemical or enzymatic means prevents NuMA cortical localization during mitosis; increasing PIP2 augments cortical NuMA. During anaphase, LGN/Gαi are dispensable for NuMA cortical enrichment but PIP/PIP2 are required. In vitro lipid-binding assay, chemical/enzymatic PIP depletion, immunofluorescence, siRNA knockdown The EMBO journal High 24996901
2016 Aurora-A directly phosphorylates the C-terminus of NuMA on three serine residues, including Ser1969; this phosphorylation governs the dynamic exchange of NuMA between cytoplasm/spindle poles and cortex; partial Aurora-A inhibition traps NuMA at spindle poles and prevents cortical enrichment. Aurora-A phosphorylation of NuMA does not affect LGN or MT binding affinity but controls NuMA mobility at poles. In vitro kinase assay, phospho-site mutagenesis, FRAP, immunofluorescence, kinase inhibition in human cells Current biology High 26832443
2018 Plk1 directly interacts with and phosphorylates NuMA; acute Plk1 inactivation enriches cortical NuMA/LGN/dynein during metaphase; Plk1-mediated NuMA phosphorylation controls NuMA cortical localization dynamics and spindle orientation. In vitro kinase assay, co-immunoprecipitation, pharmacological Plk1 inhibition, FRAP, immunofluorescence Life science alliance Medium 30456393
2009 NuMA is an essential tether linking bulk spindle microtubules to centrosomes; conditional loss of NuMA function in mice shows that centrosomes provide initial spindle focusing but centrosome-spindle attachment under tension fails without NuMA, and kinetochore fiber maintenance at poles is lost; without both centrosomes and NuMA, all spindle focusing fails. Conditional loss-of-function allele (mouse genetics), primary cell culture, immunofluorescence, live imaging The Journal of cell biology High 19255246
1999 NuMA self-assembles in vitro into multiarm oligomers (up to 10–12 arms) through C-terminal globular domain interactions; each arm corresponds to a NuMA dimer; electron microscopy and computer modeling show these 12-arm oligomers are the structural units of a quasi-hexagonal nuclear scaffold observed upon NuMA overexpression in HeLa cells. In vitro assembly, electron microscopy, computer modeling, overexpression in HeLa cells The EMBO journal High 10075938
1995 The NuMA rod domain forms a 207 nm long parallel, in-register double-stranded coiled-coil (the longest known); electron microscopy of purified recombinant fragments and full-length protein confirms tripartite structure with flanking globular domains; the SPN-3 antibody epitope maps to residues 255–267 in the first helical subdomain. Electron microscopy of recombinant protein, chemical cross-linking, circular dichroism, epitope mapping The EMBO journal High 7781599
2017 NuMA recruits dynactin to microtubule minus-ends; NuMA localizes to new minus-ends independently of dynein and faster than dynactin; a C-terminal region outside NuMA's canonical MT-binding domain mediates minus-end localization independently of γ-TuRC, CAMSAP1, or KANSL1/3; both NuMA's minus-end-binding and dynein-dynactin-binding modules are required to rescue bipolar spindle organization. Quantitative live imaging, laser ablation, domain deletion mutants, siRNA knockdown with rescue eLife High 29185983
2018 Cortical targeting of NuMA (but not dynein alone) is sufficient to generate spindle-pulling forces; NuMA assembles specialized focal structures at the cortex that cluster multiple dynein-dynactin force-generating modules; NuMA's N-terminal long arm (dynein-dynactin recruitment), dynein-based MT gliding, and direct MT-binding activity are all required; NuMA clustering is essential for spindle positioning but not spindle-pole focusing. Light-induced cortical targeting (optogenetics/reconstitution), AID-based depletion, live imaging, domain dissection eLife High 29848445
2011 Crystal structures of LGN/NuMA and LGN/mInsc complexes reveal that NuMA and Inscuteable (mInsc) interact with the same LGN TPR domain in a mutually exclusive manner; mInsc binds with higher affinity; biochemical competition assays confirm this exclusivity, suggesting Par3/mInsc/LGN and NuMA/LGN/Gαi complexes play sequential rather than simultaneous roles. X-ray crystallography, in vitro competition binding assays, cell biology studies Molecular cell High 21816348
2005 Ric-8A, a Gαi guanine nucleotide exchange factor, catalytically dissociates Gαi-GDP/LGN/NuMA complexes in vitro by releasing activated Gαi-GTP, which concomitantly liberates NuMA from LGN; this mechanism may regulate microtubule pulling forces on centrosomes during cell division. In vitro GEF assay with purified components, biochemical complex dissociation assay PNAS Medium 16275912
2010 Ric-8A and Gαi recruit LGN, NuMA, and dynein to the cell cortex during mitosis to orient the mitotic spindle; pertussis toxin (blocking Ric-8A GEF activity for Gαi), Ric-8A siRNA, or Gαi siRNA each impair cortical LGN, NuMA, and dynein localization and disturb integrin-dependent spindle orientation. siRNA knockdown, pertussis toxin treatment, live GFP-tubulin imaging, immunofluorescence Molecular and cellular biology Medium 20479129
2014 CYLD deubiquitinase deubiquitinates the cortical polarity protein dishevelled, enhancing dishevelled–NuMA interaction and promoting cortical localization of NuMA and the dynein/dynactin complex; CYLD also stabilizes astral microtubules to regulate spindle orientation. Co-immunoprecipitation, ubiquitination assay, siRNA knockdown, immunofluorescence PNAS Medium 24469800
2013 NuMA's 4.1-binding domain is required for stabilizing NuMA at the cell cortex of keratinocytes (shown by FRAP); loss of 4.1/NuMA interaction causes spindle orientation defects; NuMA is required to recruit dynactin to the cortex; at anaphase, a CDK1-independent, LGN/4.1-independent mechanism also drives cortical NuMA accumulation. FRAP, domain mutant expression, siRNA knockdown, immunofluorescence in primary keratinocytes Molecular biology of the cell Medium 24109598
2016 NuMA's direct MT-binding domain (targeting MT tips) is essential for spindle orientation establishment in keratinocytes; dynein/dynactin alone is insufficient; loss of NuMA–MT interactions in mouse skin causes spindle orientation defects, epidermal differentiation failure, and neonatal lethality; in adult mice, loss of NuMA-MT interactions disrupts hair follicle morphogenesis. MT-binding domain mutagenesis, conditional knockout in mouse skin, live imaging, immunofluorescence eLife High 26765568
2017 Crystal structure of Importin-α bound to the NuMA C-terminus reveals a novel NLS binding pattern enabling selective NLS recognition; Importin-β, in the presence of Importin-α, sterically masks a high-affinity MT-binding region of NuMA C-terminal to the NLS, thereby inhibiting NuMA's MT-bundling activity; RanGTP releases this inhibition. X-ray crystallography, in vitro MT-binding assay, Importin-β binding assay, mutagenesis The Journal of cell biology High 28939615
2020 NuMA contains two dynein LIC-binding sites: a hook domain contacting LIC1/LIC2 through a conserved hydrophobic patch (shared with Hook adaptors) and a CC1-box-like motif within its coiled-coil; both LIC-binding sites are essential for correct spindle positioning and cell division. X-ray crystallography of NuMA–LIC1/2 interfaces, mutagenesis, functional cell division assay Structure High 32413290
1999 Once NuMA is incorporated into the insoluble spindle pole matrix in vivo or in vitro, its insolubility becomes independent of microtubules; immunogold EM shows NuMA at an electron-dense material distinct from pericentriolar material that appears to anchor microtubule ends; NuMA is essential for formation of this insoluble matrix. Immunogold electron microscopy, biochemical fractionation, immunodepletion of cell-free extracts Cell motility and the cytoskeleton Medium 10098933
2006 Rae1 (mRNA export factor) has a mitosis-specific interaction with NuMA; Rae1 binds a specific site on NuMA that would convert a NuMA dimer to a tetravalent MT crosslinker; reducing Rae1 or increasing NuMA disrupts spindle architecture; co-depletion or co-overexpression of Rae1 with NuMA prevents aberrant spindle formation. Co-immunoprecipitation, domain mapping, overexpression, siRNA knockdown in HeLa cells PNAS Medium 17172455
2004 NuMA dynamics at spindle poles involve continuous exchange between soluble and spindle-associated pools (t½ ~3 min by FRAP); this exchange requires cellular energy; LGN binding and protein kinase activity both regulate NuMA's dynamic exchange rate on spindle pole asters in cell-free extracts. FRAP in live cells, cell-free aster exchange assay, LGN addition, kinase inhibitors Journal of cell science Medium 15561764
2021 NuMA undergoes liquid-liquid phase separation (LLPS) at mitotic entry; this is regulated by Aurora-A phosphorylation; KifC1 facilitates NuMA condensate concentration at spindle poles; phase separation is mediated by NuMA's C-terminus and dynein-dynactin binding motif; phase-separated NuMA concentrates tubulins, binds MTs, and enriches Kif2A (a spindle MT depolymerizer) at poles to promote poleward MT flux. In vitro phase separation assay, live cell imaging of condensates, siRNA knockdown, phosphomimetic/phosphodead mutants Nature communications Medium 34887424
2014 NuMA accumulates at DNA damage sites in a poly-ADP-ribosylation (PARylation)-dependent manner; NuMA co-immunoprecipitates with the chromatin remodeler SNF2h/SMARCA5 and regulates SNF2h diffusion and its accumulation at DNA breaks; NuMA knockdown impairs chromatin decompaction after DNA cleavage, reduces homologous recombination repair factor recruitment, and impairs DSB repair in chromosomal (not episomal) contexts. Co-immunoprecipitation, FRAP, siRNA knockdown, laser microirradiation, DNA repair assays Nucleic acids research Medium 24753406
2013 NuMA binds p53 and is required for CDK8 (Mediator complex component) recruitment to p53 target gene promoters; acute NuMA knockdown selectively attenuates p21 induction after DNA damage (impairing cell cycle arrest) but does not affect PUMA induction, demonstrating NuMA confers selectivity to p53-mediated transcription. Co-immunoprecipitation, siRNA knockdown, chromatin immunoprecipitation, transcriptional reporter assays Molecular and cellular biology Medium 23589328
2019 NuMA interacts with 53BP1 and controls 53BP1 diffusion throughout the nucleoplasm; this interaction is reduced after DNA damage; NuMA prevents 53BP1 accumulation at DNA breaks in the undamaged state; manipulating NuMA expression alters PARP inhibitor sensitivity in BRCA1-null cells and affects end-joining activity and immunoglobulin class switching. Co-immunoprecipitation, FRAP, siRNA knockdown, NHEJ assay, class-switch recombination assay Nucleic acids research Medium 30812030
2020 In neurons, NuMA1 is transiently located at the axon initial segment (AIS) where it interacts with scaffolding protein 4.1B and the dynein regulator Lis1; NuMA1 inhibits endocytosis of AIS protein NF186 by impeding Lis1's interaction with doublecortin; silencing NuMA1 or 4.1B disrupts AIS assembly but not maintenance. Differential proteomics, co-immunoprecipitation, shRNA knockdown, endocytosis assay, immunofluorescence in mouse neurons The Journal of cell biology Medium 31727776
2021 NUMA1 is downregulated in Huntington's disease neural progenitors by miR-124; suppression of NUMA1 in wild-type cells recapitulates HD microtubule bundling defects within axonal growth cones and impairs axon growth; restoring NUMA1 levels (via antagomiR-124) or stabilizing microtubules with epothilone B rescues microtubule organization and axonal growth. Proteomics of growth cones, miRNA manipulation, shRNA knockdown, rescue by antagomiR or drug, live imaging of growth cones Neuron Medium 34793694
2000 GAS41 (a protein amplified in gliomas) directly binds the C-terminal rod region of NuMA (Kd ~2×10⁻⁷ M) as shown by dot overlay and surface plasmon resonance; GAS41 is nucleolar in interphase and becomes diffuse in mitosis. Yeast two-hybrid, dot overlay, surface plasmon resonance The Journal of biological chemistry Low 10913114
2017 NuMA controls rDNA transcription and mediates nucleolar stress response in a p53-independent manner; NuMA co-immunoprecipitates with RNA Pol I, ribosomal proteins RPL26/RPL24, and B-WICH complex components; NuMA binds 18S and 28S rRNAs and localizes to rDNA promoter regions; NuMA knockdown decreases nascent pre-rRNA synthesis and triggers p27kip1 upregulation. Co-immunoprecipitation, RNA-binding assay, ChIP, siRNA knockdown, nascent RNA labeling Nucleic acids research Medium 28981686
2016 NuMA directly binds Astrin (C-terminal tail of NuMA to C-terminus of Astrin); NuMA is required for Astrin recruitment to the mitotic spindle; LGN N-terminus (which blocks NuMA MT binding) also displaces Astrin from NuMA; dynein-mediated transport is required for spindle pole accumulation of Astrin; conversely, reduced Astrin impairs NuMA concentration at spindle poles. Co-immunoprecipitation, GST pulldown, siRNA knockdown, immunofluorescence The Journal of biological chemistry Medium 27462074
1997 NuMA is cleaved during apoptosis; the cleavage site lies between residues 1701 and 1725 generating a stable ~180–200 kDa fragment; cleavage is retarded by TPCK but not by ICE inhibitors or other protease inhibitors tested. Immunoblot, protease inhibitor treatment, apoptosis induction in cell lines Experimental cell research Medium 9184071
2003 NuMA is preferentially cleaved by caspase-3 during Fas-mediated apoptosis; NuMA cleavage coincides with lamin B and PARP-1 cleavage; NuMA redistribution during apoptosis involves condensation, central nuclear concentration, and encirclement of apoptotic body fragments; NuMA was not cleaved in caspase-3-null MCF-7 cells treated with staurosporine. Immunofluorescence, immunoblot, caspase inhibitors, caspase-3-null cell line, Fas stimulation Journal of cell science Medium 12508117
1994 NuMA residues 1972–2007 constitute a nuclear localization signal (NLS); Lys1988 is essential for nuclear targeting; residues 1538–2115 are necessary and sufficient for spindle association; NuMA isoforms NuMA-m and NuMA-s (arising from alternative splicing) localize to centrosomes in interphase and spindle poles in mitosis rather than the nucleus. Linker scanning mutagenesis, chimeric protein expression, transfection in CHO cells, immunofluorescence Journal of cell science Medium 7962183
2017 Galectin-3 associates with NuMA at spindle poles in an O-GlcNAcylation-dependent manner during metaphase; loss of Galectin-3 impairs spindle pole cohesion and stable NuMA localization at the spindle pole in epithelial cells. Co-immunoprecipitation, siRNA knockdown, immunofluorescence, experimental mouse model Scientific reports Medium 28469279
2019 In acentrosomal human cells, NuMA forms small microtubule asters at nuclear envelope breakdown that are assembled by dynein and NuMA's clustering activity; NuMA organizes radial MT arrays incorporating Eg5 to promote spindle bipolarization; in cells with centrosomes, NuMA also promotes the initial step of spindle bipolarization. siRNA depletion of centrosome components, live imaging, NuMA siRNA in acentrosomal cells, immunofluorescence The EMBO journal Medium 31782546

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1996 A complex of NuMA and cytoplasmic dynein is essential for mitotic spindle assembly. Cell 488 8898198
2004 Mammalian Pins is a conformational switch that links NuMA to heterotrimeric G proteins. Cell 320 15537540
2000 Formation of spindle poles by dynein/dynactin-dependent transport of NuMA. The Journal of cell biology 278 10811826
2006 The NuMA-related Mud protein binds Pins and regulates spindle orientation in Drosophila neuroblasts. Nature cell biology 267 16648843
2006 The Drosophila NuMA Homolog Mud regulates spindle orientation in asymmetric cell division. Developmental cell 251 16740476
2001 A mammalian Partner of inscuteable binds NuMA and regulates mitotic spindle organization. Nature cell biology 223 11781568
1995 NuMA is required for the organization of microtubules into aster-like mitotic arrays. The Journal of cell biology 220 7593190
1992 NuMA: an unusually long coiled-coil related protein in the mammalian nucleus. The Journal of cell biology 220 1541630
1997 Fusion of retinoic acid receptor alpha to NuMA, the nuclear mitotic apparatus protein, by a variant translocation in acute promyelocytic leukaemia. Nature genetics 211 9288109
1992 Primary structure of NuMA, an intranuclear protein that defines a novel pathway for segregation of proteins at mitosis. The Journal of cell biology 207 1541636
1993 NuMA is required for the proper completion of mitosis. The Journal of cell biology 186 8432734
2010 Ric-8A and Gi alpha recruit LGN, NuMA, and dynein to the cell cortex to help orient the mitotic spindle. Molecular and cellular biology 139 20479129
2010 NuMA after 30 years: the matrix revisited. Trends in cell biology 136 20137953
2005 NuMA is a major acceptor of poly(ADP-ribosyl)ation by tankyrase 1 in mitosis. The Biochemical journal 131 16076287
2002 Identification of a tankyrase-binding motif shared by IRAP, TAB182, and human TRF1 but not mouse TRF1. NuMA contains this RXXPDG motif and is a novel tankyrase partner. The Journal of biological chemistry 131 12080061
2002 LGN blocks the ability of NuMA to bind and stabilize microtubules. A mechanism for mitotic spindle assembly regulation. Current biology : CB 126 12445386
2011 A lateral belt of cortical LGN and NuMA guides mitotic spindle movements and planar division in neuroepithelial cells. The Journal of cell biology 123 21444683
2009 Requirements for NuMA in maintenance and establishment of mammalian spindle poles. The Journal of cell biology 119 19255246
2010 The Fz-Dsh planar cell polarity pathway induces oriented cell division via Mud/NuMA in Drosophila and zebrafish. Developmental cell 111 21074723
1999 A nonerythroid isoform of protein 4.1R interacts with the nuclear mitotic apparatus (NuMA) protein. The Journal of cell biology 111 10189366
2018 Dynein-Dynactin-NuMA clusters generate cortical spindle-pulling forces as a multi-arm ensemble. eLife 110 29848445
2011 LGN/mInsc and LGN/NuMA complex structures suggest distinct functions in asymmetric cell division for the Par3/mInsc/LGN and Gαi/LGN/NuMA pathways. Molecular cell 108 21816348
2009 NuMA-related LIN-5, ASPM-1, calmodulin and dynein promote meiotic spindle rotation independently of cortical LIN-5/GPR/Galpha. Nature cell biology 107 19219036
1995 Mutation of the predicted p34cdc2 phosphorylation sites in NuMA impair the assembly of the mitotic spindle and block mitosis. Journal of cell science 100 7769006
1999 Self assembly of NuMA: multiarm oligomers as structural units of a nuclear lattice. The EMBO journal 99 10075938
2006 Rae1 interaction with NuMA is required for bipolar spindle formation. Proceedings of the National Academy of Sciences of the United States of America 96 17172455
2014 CYLD regulates spindle orientation by stabilizing astral microtubules and promoting dishevelled-NuMA-dynein/dynactin complex formation. Proceedings of the National Academy of Sciences of the United States of America 95 24469800
1994 Localization of NuMA protein isoforms in the nuclear matrix of mammalian cells. Cell motility and the cytoskeleton 95 7820866
2013 NuMA phosphorylation by CDK1 couples mitotic progression with cortical dynein function. The EMBO journal 88 23921553
2017 NuMA recruits dynein activity to microtubule minus-ends at mitosis. eLife 86 29185983
1994 NuMA, a nuclear protein involved in mitosis and nuclear reformation. Current opinion in cell biology 84 7917323
2016 NuMA Phosphorylation by Aurora-A Orchestrates Spindle Orientation. Current biology : CB 78 26832443
2014 SLK-dependent activation of ERMs controls LGN-NuMA localization and spindle orientation. The Journal of cell biology 78 24958772
2009 Interaction between Poly(ADP-ribose) and NuMA contributes to mitotic spindle pole assembly. Molecular biology of the cell 78 19759176
1998 The role of NuMA in the interphase nucleus. Journal of cell science 78 9394013
2016 NuMA-microtubule interactions are critical for spindle orientation and the morphogenesis of diverse epidermal structures. eLife 77 26765568
2005 Resistance to inhibitors of cholinesterase 8A catalyzes release of Galphai-GTP and nuclear mitotic apparatus protein (NuMA) from NuMA/LGN/Galphai-GDP complexes. Proceedings of the National Academy of Sciences of the United States of America 77 16275912
2013 NuMA localization, stability, and function in spindle orientation involve 4.1 and Cdk1 interactions. Molecular biology of the cell 75 24109598
1986 Redistribution of the nuclear mitotic apparatus protein (NuMA) during mitosis and nuclear assembly. Properties of purified NuMA protein. Experimental cell research 75 3527729
1999 NuMA is a component of an insoluble matrix at mitotic spindle poles. Cell motility and the cytoskeleton 71 10098933
2011 Inscuteable and NuMA proteins bind competitively to Leu-Gly-Asn repeat-enriched protein (LGN) during asymmetric cell divisions. Proceedings of the National Academy of Sciences of the United States of America 67 22171003
2014 NuMA interacts with phosphoinositides and links the mitotic spindle with the plasma membrane. The EMBO journal 65 24996901
1996 Dynamic changes of NuMA during the cell cycle and possible appearance of a truncated form of NuMA during apoptosis. Journal of cell science 65 8838651
2002 Unraveling the organization of the internal nuclear matrix: RNA-dependent anchoring of NuMA to a lamin scaffold. Experimental cell research 64 12243746
1998 Induction of a regular nuclear lattice by overexpression of NuMA. Experimental cell research 63 9743603
1996 NuMA assembles into an extensive filamentous structure when expressed in the cell cytoplasm. Journal of cell science 63 8907707
1993 Microinjection of a monoclonal antibody against SPN antigen, now identified by peptide sequences as the NuMA protein, induces micronuclei in PtK2 cells. Journal of cell science 63 8449992
2006 Role of NuMA in vertebrate cells: review of an intriguing multifunctional protein. Frontiers in bioscience : a journal and virtual library 60 16146802
1995 NuMA: a protein involved in nuclear structure, spindle assembly, and nuclear re-formation. Trends in cell biology 60 14731413
2006 NuMA influences higher order chromatin organization in human mammary epithelium. Molecular biology of the cell 59 17108325
1997 Phosphorylation regulates the assembly of NuMA in a mammalian mitotic extract. Journal of cell science 59 9202389
2021 NuMA regulates mitotic spindle assembly, structural dynamics and function via phase separation. Nature communications 58 34887424
2000 NuMA: a nuclear protein involved in mitotic centrosome function. Microscopy research and technique 58 10842374
1995 Epitope mapping and direct visualization of the parallel, in-register arrangement of the double-stranded coiled-coil in the NuMA protein. The EMBO journal 56 7781599
1994 Nuclear mitotic apparatus protein (NuMA): spindle association, nuclear targeting and differential subcellular localization of various NuMA isoforms. Journal of cell science 55 7962183
1984 NuMA protein is a human autoantigen. Arthritis and rheumatism 55 6378210
2011 aPKC phosphorylates NuMA-related LIN-5 to position the mitotic spindle during asymmetric division. Nature cell biology 54 21857670
1993 Nuclear proteins of the bovine esophageal epithelium. II. The NuMA gene gives rise to multiple mRNAs and gene products reactive with monoclonal antibody W1. Journal of cell science 54 8505359
2014 NuMA promotes homologous recombination repair by regulating the accumulation of the ISWI ATPase SNF2h at DNA breaks. Nucleic acids research 52 24753406
2004 Multiple mechanisms regulate NuMA dynamics at spindle poles. Journal of cell science 52 15561764
2013 Cell cycle-regulated membrane binding of NuMA contributes to efficient anaphase chromosome separation. Molecular biology of the cell 51 24371089
2021 Developmental defects in Huntington's disease show that axonal growth and microtubule reorganization require NUMA1. Neuron 50 34793694
2019 NuMA assemblies organize microtubule asters to establish spindle bipolarity in acentrosomal human cells. The EMBO journal 49 31782546
1997 Cleavage of the nuclear matrix protein NuMA during apoptosis. Experimental cell research 49 9184071
2021 The Nuclear Mitotic Apparatus (NuMA) Protein: A Key Player for Nuclear Formation, Spindle Assembly, and Spindle Positioning. Frontiers in cell and developmental biology 48 33869212
2004 Cell and molecular biology of spindle poles and NuMA. International review of cytology 47 15364196
2013 Evidence for dynein and astral microtubule-mediated cortical release and transport of Gαi/LGN/NuMA complex in mitotic cells. Molecular biology of the cell 45 23389635
2003 A functional relationship between NuMA and kid is involved in both spindle organization and chromosome alignment in vertebrate cells. Molecular biology of the cell 44 12972545
2011 Rab5 GTPase controls chromosome alignment through Lamin disassembly and relocation of the NuMA-like protein Mud to the poles during mitosis. Proceedings of the National Academy of Sciences of the United States of America 42 21987826
2005 Association of the NuMA region on chromosome 11q13 with breast cancer susceptibility. Proceedings of the National Academy of Sciences of the United States of America 41 15684076
2004 Cyclin B degradation leads to NuMA release from dynein/dynactin and from spindle poles. EMBO reports 41 14710193
2000 GAS41, a highly conserved protein in eukaryotic nuclei, binds to NuMA. The Journal of biological chemistry 38 10913114
2012 Interconnected contribution of tissue morphogenesis and the nuclear protein NuMA to the DNA damage response. Journal of cell science 37 22331358
2009 NuMA is required for proper spindle assembly and chromosome alignment in prometaphase. BMC research notes 37 19400937
1993 Primary structure and microtubule-interacting domain of the SP-H antigen: a mitotic MAP located at the spindle pole and characterized as a homologous protein to NuMA. Journal of cell science 37 8408288
2020 The Aurora-A/TPX2 Axis Directs Spindle Orientation in Adherent Human Cells by Regulating NuMA and Microtubule Stability. Current biology : CB 36 33275894
2020 NuMA1 promotes axon initial segment assembly through inhibition of endocytosis. The Journal of cell biology 35 31727776
2017 Katanin p80, NuMA and cytoplasmic dynein cooperate to control microtubule dynamics. Scientific reports 35 28079116
2000 Preferential expression of NuMA in the nuclei of proliferating cells. Experimental cell research 34 10739661
1995 Phosphorylation of NUMA occurs during nuclear breakdown and not mitotic spindle assembly. Journal of cell science 34 8586651
2017 The nuclear mitotic apparatus protein NuMA controls rDNA transcription and mediates the nucleolar stress response in a p53-independent manner. Nucleic acids research 33 28981686
2013 NuMA is required for the selective induction of p53 target genes. Molecular and cellular biology 33 23589328
2004 Myeloid leukemia with promyelocytic features in transgenic mice expressing hCG-NuMA-RARalpha. Oncogene 33 14737102
2017 Regulation of mitotic spindle assembly factor NuMA by Importin-β. The Journal of cell biology 32 28939615
2016 SAPCD2 Controls Spindle Orientation and Asymmetric Divisions by Negatively Regulating the Gαi-LGN-NuMA Ternary Complex. Developmental cell 32 26766442
2013 Par1b links lumen polarity with LGN-NuMA positioning for distinct epithelial cell division phenotypes. The Journal of cell biology 32 24165937
1996 NuMA: a bipartite nuclear location signal and other functional properties of the tail domain. Experimental cell research 32 8635513
2019 The nuclear structural protein NuMA is a negative regulator of 53BP1 in DNA double-strand break repair. Nucleic acids research 30 30812030
2018 Plk1 regulates spindle orientation by phosphorylating NuMA in human cells. Life science alliance 30 30456393
1999 The plant nucleoskeleton: ultrastructural organization and identification of NuMA homologues in the nuclear matrix and mitotic spindle of plant cells. Experimental cell research 30 9925768
2003 NuMA and nuclear lamins behave differently in Fas-mediated apoptosis. Journal of cell science 28 12508117
2014 Isoform-specific functions of Mud/NuMA mediate binucleation of Drosophila male accessory gland cells. BMC developmental biology 27 25527079
2016 Nuclear Mitotic Apparatus (NuMA) Interacts with and Regulates Astrin at the Mitotic Spindle. The Journal of biological chemistry 26 27462074
2012 On the inscrutable role of Inscuteable: structural basis and functional implications for the competitive binding of NuMA and Inscuteable to LGN. Open biology 26 22977735
2011 The nuclear mitotic apparatus (NuMA) protein: localization and dynamics in human oocytes, fertilization and early embryos. Molecular human reproduction 25 21297155
2020 Organizational Principles of the NuMA-Dynein Interaction Interface and Implications for Mitotic Spindle Functions. Structure (London, England : 1993) 24 32413290
2017 Spindle pole cohesion requires glycosylation-mediated localization of NuMA. Scientific reports 24 28469279
2002 Association of the nuclear matrix component NuMA with the Cajal body and nuclear speckle compartments during transitions in transcriptional activity in lens cell differentiation. European journal of cell biology 24 12437190
2003 Essential role for the dimerization domain of NuMA-RARalpha in its oncogenic activities and localization to NuMA sites within the nucleus. Oncogene 23 12584566
2018 Activated ezrin controls MISP levels to ensure correct NuMA polarization and spindle orientation. Journal of cell science 22 29669740