Affinage

KNL1

Outer kinetochore KNL1 complex subunit KNL1 · UniProt Q8NG31

Length
2342 aa
Mass
265.4 kDa
Annotated
2026-04-28
54 papers in source corpus 24 papers cited in narrative 24 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

KNL1 is a large outer kinetochore scaffold protein that integrates mitotic checkpoint signaling, error correction, and checkpoint silencing by serving as a phosphorylation-regulated platform for the recruitment of checkpoint and phosphatase effectors. MPS1 kinase phosphorylates arrays of MELT (and vertebrate-specific SHT) repeats on KNL1 to create docking sites for BUB3–BUB1 and BUB3–BubR1 complexes, with KI motifs enhancing this recruitment, thereby activating the spindle assembly checkpoint at tensionless kinetochores (PMID:22660415, PMID:25661489, PMID:24344183, PMID:24361068). KNL1 simultaneously recruits PP1 phosphatase via a conserved RVSF motif—in a manner mutually exclusive with microtubule binding—to dephosphorylate Aurora B substrates and silence the checkpoint, while Aurora B phosphorylation of KNL1 disrupts PP1 binding to sustain error correction, and BubR1-associated PP2A-B56 dephosphorylates MELT motifs to further promote silencing (PMID:20231380, PMID:30100357, PMID:25246613). Conditional deletion of KNL1 in the mouse brain causes chromosome missegregation, p53-dependent apoptosis, and microcephaly, and in C. elegans postmitotic neurons KNL1 modulates F-actin dynamics to control dendrite branching and axonal organization (PMID:31197172, PMID:39625434, PMID:38656792).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2003 High

    Establishing KNL-1 as a core outer kinetochore component resolved how centromeric chromatin connects to the microtubule-binding interface: KNL-1 acts downstream of CENP-A/C and is required for recruitment of the Ndc80 complex and other outer kinetochore proteins.

    Evidence RNAi-based genomics, co-immunoprecipitation from C. elegans embryonic extracts, epistasis analysis

    PMID:14522947

    Open questions at the time
    • Human orthologue function not yet tested
    • Structural basis of KNL-1–Ndc80 interaction unknown
  2. 2007 High

    Identification of KNL1 as the direct kinetochore receptor for Bub1 and BubR1 (via their TPR domains) explained how SAC effectors are targeted to kinetochores, linking outer kinetochore architecture to checkpoint signaling.

    Evidence Direct binding assays, co-immunoprecipitation, RNAi in human cells, live-cell imaging

    PMID:17981135 PMID:18045986

    Open questions at the time
    • Which KNL1 motifs mediate each interaction not mapped
    • Post-translational regulation of recruitment unknown
  3. 2010 High

    Discovery that KNL1's RVSF motif recruits PP1 to oppose Aurora B substrates revealed how error correction is switched off at bioriented kinetochores, and that Aurora B phosphorylation of KNL1 creates positive feedback preventing premature PP1 recruitment.

    Evidence Co-immunoprecipitation, in vitro binding with phospho-mutants, live-cell imaging, kinetochore PP1 quantification in human cells; confirmed by yeast gene replacement

    PMID:20231380 PMID:21640906

    Open questions at the time
    • Structural basis of KNL1–PP1 interaction unknown at this stage
    • How PP1 targets specific substrates at the kinetochore unclear
  4. 2012 High

    Demonstrating that MPS1 phosphorylation of MELT repeats creates individual BUB3–BUB1 docking sites resolved the molecular code by which checkpoint activity is generated proportionally on KNL1, and the KI-motif crystal structure provided an atomic-level view of Bub1–KNL1 recognition.

    Evidence In vitro kinase and binding assays, phospho-mutant genetics in fission yeast (two independent labs), X-ray crystallography of Bub1 TPR–KI complex, microtubule-binding reconstitution in C. elegans

    PMID:22331848 PMID:22331849 PMID:22521786 PMID:22660415

    Open questions at the time
    • Vertebrate-specific modifications of the MELT code not yet characterized
    • How KI and MELT motifs cooperate mechanistically unresolved
  5. 2013 High

    Systematic dissection of KNL1's modular MELT/KI repeat array established that a minimum of four active repeats sustains SAC and chromosome congression, with KI motifs acting as enhancers of adjacent MELT function, framing KNL1 as a tunable signaling scaffold.

    Evidence Engineered KNL1 variants with defined repeat numbers, rescue assays in KNL1-depleted human cells, quantitative immunofluorescence

    PMID:24344183 PMID:24361068 PMID:24363448

    Open questions at the time
    • How PP1 tunes effective MELT repeat occupancy in real time unclear
    • Role of N-terminal KNL1 domain in Aurora B activity requires further mechanistic parsing
  6. 2014 High

    Identification of PP2A-B56 (recruited via BubR1) as the phosphatase that dephosphorylates MELT motifs closed the negative feedback loop for checkpoint silencing, showing that the SAC generates its own off-switch through BubR1-associated phosphatase.

    Evidence In vitro phosphatase assay on MELT peptides, cell-based depletion and rescue, quantitative kinetochore protein measurements

    PMID:25246613

    Open questions at the time
    • Relative contributions of PP1 vs PP2A-B56 to MELT dephosphorylation in vivo not quantified
    • Timing of phosphatase engagement during mitotic progression unclear
  7. 2015 High

    Discovery of the vertebrate-specific SHT motif and its sequential phosphorylation by MPS1 (requiring prior MELT phosphorylation) refined the MELT code into a two-step phospho-switch that synergistically enhances BUB3 docking, explaining vertebrate-specific checkpoint amplification.

    Evidence In vitro binding screens, sequential phosphorylation assays, BUB3 surface mutant localization in human cells

    PMID:25661489

    Open questions at the time
    • Structural basis of BUB3 recognition of phospho-SHT not determined
    • Whether SHT phosphorylation is also reversed by PP2A-B56 not tested
  8. 2015 High

    Genetic separation of the KBB (KNL1–Bub3–Bub1) and RZZ pathways for Mad1-Mad2 recruitment revealed that KNL1's checkpoint role is specifically required at tensionless (misaligned) kinetochores, not at fully unattached ones, resolving a long-standing ambiguity about checkpoint pathway redundancy.

    Evidence CRISPR gene editing and RNAi in diploid human cells, Mad2 kinetochore localization assays

    PMID:26581576 PMID:26651294

    Open questions at the time
    • Molecular basis for tension-dependent selectivity of the KBB pathway unknown
    • How RZZ recruitment depends on KNL1/Bub1 not fully resolved
  9. 2018 High

    Structural and biophysical analyses revealed that PP1 and microtubules bind overlapping surfaces on KNL1's N-terminus in a mutually exclusive manner, providing a molecular switch that couples microtubule attachment status to phosphatase activity at the kinetochore.

    Evidence X-ray crystallography, NMR spectroscopy, co-sedimentation and competition assays

    PMID:30100357

    Open questions at the time
    • How the switch operates dynamically in the context of the full KMN network not tested
    • Whether MT binding displaces PP1 or vice versa in vivo not determined
  10. 2019 High

    Conditional KNL1 deletion in the developing mouse brain demonstrated that KNL1 loss causes chromosome missegregation, p53-dependent apoptosis, and microcephaly, directly linking kinetochore dysfunction to neurodevelopmental disease.

    Evidence Conditional Cre-lox knockout in mouse brain, mitotic index analysis, p53 double-KO epistasis

    PMID:31197172

    Open questions at the time
    • Whether human CASC5/KNL1 mutations cause microcephaly through the same apoptotic pathway not established
    • Cell-type specificity of p53 response not explored
  11. 2024 Medium

    Discovery that postmitotic KNL-1 (with KMN partners) modulates F-actin dynamics to control dendrite branching and axonal organization in C. elegans neurons revealed a non-mitotic cytoskeletal function, where PP1 and SAC-signaling motifs are also required for proper neurodevelopment.

    Evidence C. elegans genetics, gene-replacement with motif-specific mutations, live actin/MT imaging, neuronal morphology quantification

    PMID:38656792 PMID:39625434

    Open questions at the time
    • Mechanism by which KNL-1 N-terminus initiates F-actin assembly unknown
    • Whether mammalian KNL1 has analogous postmitotic neuronal functions not tested
    • Signaling pathway linking SAC motifs to actin regulation uncharacterized

Open questions

Synthesis pass · forward-looking unresolved questions
  • How KNL1 coordinates the kinase–phosphatase balance in real time during the metaphase-to-anaphase transition, and whether its postmitotic cytoskeletal functions are conserved in vertebrate neurons, remain open mechanistic questions.
  • No live single-molecule studies tracking KNL1 phospho-state transitions during checkpoint silencing
  • No structural model of full-length KNL1 or its complex with multiple MELT-bound BUB modules
  • Vertebrate postmitotic neuronal role not tested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 5 GO:0008092 cytoskeletal protein binding 3 GO:0098772 molecular function regulator activity 3
Localization
GO:0005694 chromosome 5 GO:0005856 cytoskeleton 2
Pathway
R-HSA-1640170 Cell Cycle 10 R-HSA-1266738 Developmental Biology 3
Partners
BUB1BUB3BUBR1MIS12MPS1NDC80PPP1CA
Complex memberships
KMN networkKNL1-Mis12 complex

Evidence

Reading pass · 24 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2003 KNL-1 (C. elegans ortholog) is required downstream of CeCENP-A and CeCENP-C in a linear kinetochore assembly hierarchy, forms a near-stoichiometric complex with CeNDC-80 and HIM-10, and is required to target multiple outer kinetochore components including the Ndc80 complex to build the microtubule-binding interface. RNAi-based genomics, co-immunoprecipitation from embryonic extracts, epistasis analysis Genes & development High 14522947
2007 Human KNL1 (Blinkin/AF15q14) directly interacts via its N-terminal and middle domains with the TPR domains of Bub1 and BubR1, recruiting them to kinetochores; its C-terminal domain associates with the hMis12 complex. KNL1 knockdown causes checkpoint failure and chromosome misalignment phenocopying Bub1+BubR1 double knockdown. RNAi, direct binding assays, co-immunoprecipitation, live-cell imaging Developmental cell High 17981135
2007 Vertebrate KNL1 and CENP-K coordinately direct localization of the Ndc80 complex to kinetochores; simultaneous depletion of both abolishes all kinetochore assembly downstream of centromeric chromatin, revealing functional redundancy not present in C. elegans. RNAi depletion, immunofluorescence, epistasis analysis in chicken DT40 cells Molecular biology of the cell High 18045986
2010 A conserved RVSF motif in KNL1 directly recruits protein phosphatase 1 (PP1) to the outer kinetochore to dephosphorylate Aurora B substrates and stabilize microtubule attachments; Aurora B phosphorylation of KNL1 disrupts the KNL1-PP1 interaction, creating a positive feedback mechanism that prevents PP1 recruitment where Aurora B is active. Co-immunoprecipitation, in vitro binding assays, phospho-mutant analysis, live-cell imaging, kinetochore PP1 quantification The Journal of cell biology High 20231380
2011 The RVSF motif of yeast Spc105 (KNL1 ortholog) binds PP1/Glc7 and this interaction is essential for viability by silencing the spindle assembly checkpoint; PP1 amount at the kinetochore must be finely tuned as either loss or gain of one extra PP1 copy is detrimental. Exact gene replacement in budding yeast, genetic epistasis, quantitative imaging Current biology : CB High 21640906
2012 Fission yeast Mph1 (MPS1 ortholog) phosphorylates MELT repeat sequences of Spc7 (KNL1/Blinkin ortholog), and this phosphorylation directly promotes binding of the Bub1-Bub3 complex to kinetochores, which is required for SAC activation (Mad1-Mad2-Mad3 localization) and chromosome alignment. Non-phosphorylatable spc7-12A abolishes Bub1-Bub3 kinetochore targeting; phosphomimetic spc7-12E forces constitutive localization. In vitro kinase assay, in vitro binding assay, phosphomutant genetics, live-cell imaging in fission yeast Nature cell biology High 22660415
2012 Mph1 (Mps1) phosphorylation of conserved MELT motifs in S. pombe Spc7 recruits Bub1 and Bub3 to the kinetochore, and this phospho-dependent recruitment is required to maintain the SAC signal. Phospho-mutant genetics, mass spectrometry, in vitro binding, kinetochore localization assays Current biology : CB High 22521786
2012 The N-terminal domain of KNL-1 contains microtubule-binding and -bundling activity; this activity is dispensable for load-bearing attachment and checkpoint activation but contributes independently to checkpoint silencing at the kinetochore, additively with PP1 docking. In vitro microtubule-binding assay, selective point mutagenesis, C. elegans embryo imaging, checkpoint silencing assays The Journal of cell biology High 22331849
2012 Crystal structure of the Bub1 TPR domain in complex with the KI motif of Knl1 was determined; point mutations on the convex TPR surface impaired the Bub1-Knl1 interaction in vitro and in vivo. A 62-residue segment of Bub1 C-terminal to the TPRs (containing the Bub3-binding domain) was found necessary and sufficient for kinetochore recruitment of Bub1. X-ray crystallography, in vitro binding assays, site-directed mutagenesis, cell-based localization The Journal of cell biology High 22331848
2013 KNL1 contains an array of MELT repeats that serve as individual Mps1-phosphorylated docking sites for Bub3 (and thereby Bub1/BubR1); a minimum of four active MELT repeats is sufficient to support chromosome congression and SAC function. PP1 binding to KNL1 during prometaphase reduces Bub protein levels at kinetochores to the equivalent of four active MELT repeats. MELT repeat deletion constructs, rescue assays in KNL1-depleted cells, quantitative immunofluorescence Journal of cell science High 24363448
2013 KNL1 contains an extensive array of short linear BUB recruitment modules (TxxΩ and MELT motifs) that can independently localize BUB1; increasing numbers of modules progressively enhance chromosome biorientation efficiency, while a minimal array suffices for robust checkpoint. A minimal set of artificially designed identical modules maintains normal KNL1 function. Engineered KNL1 variants with defined module numbers, BUB1 quantification at kinetochores, chromosome segregation assays The Journal of cell biology High 24344183
2013 KNL1 N-terminus is essential for Aurora B kinase activity at kinetochores; KNL1 promotes Aurora B activity partly through supporting Bub1 kinase activity, and partly through an additional Bub1-independent pathway. Ectopic targeting of Aurora B does not fully rescue Aurora B activity upon KNL1 depletion. RNAi depletion, phospho-substrate immunofluorescence, ectopic targeting constructs, live-cell imaging The Journal of cell biology High 24344188
2013 KI motifs in the N-terminal region of human Knl1 cooperate with the adjacent MELT motif to assemble comprehensive SAC signaling complexes; Knl1(1-250) containing both KI motifs and one MELT motif can fully restore SAC and chromosome alignment in Knl1-depleted cells. KI motifs function as enhancers of MELT function. Knl1 truncation/rescue constructs, co-immunoprecipitation, kinetochore localization assays, SAC functional assays Current biology : CB High 24361068
2014 BubR1-associated PP2A-B56 dephosphorylates Mps1-phosphorylated MELT motifs on Knl1 in vitro and in vivo, thereby removing the docking sites for Bub1/BubR1 and promoting SAC silencing via a negative feedback loop. In vitro phosphatase assay, cell-based rescue/depletion, quantitative kinetochore protein measurements The Journal of cell biology High 25246613
2014 C. elegans KNL-1 exists as a decameric oligomer mediated by a small hydrophobic N-terminal domain; however, precise disruption of this oligomerization does not alter KNL-1 localization or embryonic viability in gene replacement experiments. Biochemical oligomerization assays, electron microscopy, site-directed mutagenesis, C. elegans gene replacement Molecular biology of the cell Medium 25411336
2015 Human KNL1 MELT repeats contain a vertebrate-specific SHT motif C-terminal to the MELT sequence; MPS1 phosphorylates SHT in a manner requiring prior MELT phosphorylation (sequential multisite phosphorylation), and phospho-SHT synergizes with phospho-MELT to promote BUB3/BUB1 binding in vitro and at kinetochores. BUB3 mutated in its predicted SHpT-binding surface cannot localize to kinetochores. Systematic in vitro binding screens, phosphomutant analysis, cell-based localization assays Molecular cell High 25661489
2015 The KNL1-Bub3-Bub1 (KBB) pathway is required for SAC activation at misaligned (tensionless) kinetochores during normal mitosis but is not required when kinetochores are fully unattached; the RZZ complex provides a separate, KBB-independent pathway for Mad1-Mad2 recruitment to unattached kinetochores. CRISPR/gene editing, RNAi, Mad2 kinetochore localization assays, checkpoint response assays in diploid human cells Developmental cell High 26651294
2015 The RZZ complex localizes to the N-terminus of KNL1 downstream of Bub1 to mediate robust Mad1-Mad2 kinetochore localization in human cells, establishing KNL1 as a platform for RZZ-dependent Mad1-Mad2 recruitment. RNAi, truncation constructs, kinetochore localization assays, co-immunoprecipitation Open biology Medium 26581576
2016 In fission yeast, multisite binding of Bub3 to the Spc7 (KNL1) MELT array is required for Mph1 (Mps1)-dependent interaction of Bub1 with Mad1-Mad2, thereby toggling the spindle checkpoint switch; Bub3-Spc7 binding licenses the Bub1-Mad1-Mad2 interaction. Genetic epistasis in S. pombe, co-immunoprecipitation, phosphomutant analysis, checkpoint functional assays Current biology : CB High 27618268
2018 Crystal structure and NMR analysis show that KNL1 binds PP1 via its RVSF motif and binds microtubules via overlapping binding sites, demonstrating that microtubule and PP1 binding to KNL1 are mutually exclusive; Aurora B phosphorylation of KNL1 causes distinct disruption patterns for each complex, and co-sedimentation assays confirm preferential formation of KNL1:PP1 holoenzyme in the presence of PP1. X-ray crystallography, NMR spectroscopy, in vitro binding/competition assays, co-sedimentation assays Structure High 30100357
2018 RZZ's sole role in SAC activation is to tether Mad1-Mad2 to kinetochores; Bub1 and KNL1 (via MPS1-dependent phosphorylation) activate kinetochore-bound Mad1-Mad2 to produce the 'wait anaphase' signal but are not required for fibrous corona formation. MPS1 kinase triggers fibrous corona formation by phosphorylating N-terminal sites on Rod, not KNL1. Genome editing (KO of Bub1, KNL1, RZZ subunits), kinetochore protein localization assays, checkpoint functional assays Current biology : CB High 30415700
2019 Conditional KNL1 deletion from embryonic mouse brain causes chromosome missegregation in neural progenitor cells, leading to DNA damage on missegregated chromosomes, p53 activation, apoptosis, and microcephaly; without both KNL1 and p53-dependent safeguards, genome-damaged cells persist causing lethality. Conditional Cre-lox knockout in mouse brain, mitotic index analysis, immunofluorescence, p53 double-KO epistasis Nature communications High 31197172
2024 In C. elegans postmitotic neurons, KNL-1 (with KMN network partners) controls dendrite branching by modulating F-actin dynamics; KNL-1 loss causes excess dendritic branching, altered actin and microtubule dynamics, and the N-terminus of KNL-1 can initiate F-actin assembly. C. elegans genetics, live imaging, actin dynamics assays, neuronal morphology quantification, gene replacement constructs The Journal of cell biology Medium 39625434
2024 In C. elegans, postmitotic KNL-1 is required for proper axonal organization; its PP1-recruiting SILK/RVSF motifs and SAC-activating signaling motifs (not just microtubule-binding activity) are required for neurodevelopmental function, while NDC-80 microtubule binding facilitates axon-axon contacts during nerve ring formation. C. elegans gene-replacement, fluorescent imaging, signaling motif mutagenesis, axon organization quantification Molecular biology of the cell Medium 38656792

Source papers

Stage 0 corpus · 54 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2010 Regulated targeting of protein phosphatase 1 to the outer kinetochore by KNL1 opposes Aurora B kinase. The Journal of cell biology 305 20231380
2012 MPS1/Mph1 phosphorylates the kinetochore protein KNL1/Spc7 to recruit SAC components. Nature cell biology 291 22660415
2007 Human Blinkin/AF15q14 is required for chromosome alignment and the mitotic checkpoint through direct interaction with Bub1 and BubR1. Developmental cell 249 17981135
2012 Phosphodependent recruitment of Bub1 and Bub3 to Spc7/KNL1 by Mph1 kinase maintains the spindle checkpoint. Current biology : CB 241 22521786
2003 KNL-1 directs assembly of the microtubule-binding interface of the kinetochore in C. elegans. Genes & development 200 14522947
2011 KNL1/Spc105 recruits PP1 to silence the spindle assembly checkpoint. Current biology : CB 184 21640906
2007 KNL1 and the CENP-H/I/K complex coordinately direct kinetochore assembly in vertebrates. Molecular biology of the cell 162 18045986
2012 Microtubule binding by KNL-1 contributes to spindle checkpoint silencing at the kinetochore. The Journal of cell biology 122 22331849
2012 Structural analysis reveals features of the spindle checkpoint kinase Bub1-kinetochore subunit Knl1 interaction. The Journal of cell biology 114 22331848
2013 Arrayed BUB recruitment modules in the kinetochore scaffold KNL1 promote accurate chromosome segregation. The Journal of cell biology 113 24344183
2014 PP2A-B56 opposes Mps1 phosphorylation of Knl1 and thereby promotes spindle assembly checkpoint silencing. The Journal of cell biology 107 25246613
2013 A minimal number of MELT repeats supports all the functions of KNL1 in chromosome segregation. Journal of cell science 97 24363448
2015 Sequential multisite phospho-regulation of KNL1-BUB3 interfaces at mitotic kinetochores. Molecular cell 94 25661489
2013 KI motifs of human Knl1 enhance assembly of comprehensive spindle checkpoint complexes around MELT repeats. Current biology : CB 91 24361068
2018 Distinct Roles of RZZ and Bub1-KNL1 in Mitotic Checkpoint Signaling and Kinetochore Expansion. Current biology : CB 88 30415700
2015 KNL1-Bubs and RZZ Provide Two Separable Pathways for Checkpoint Activation at Human Kinetochores. Developmental cell 78 26651294
2014 Cotton KNL1, encoding a class II KNOX transcription factor, is involved in regulation of fibre development. Journal of experimental botany 69 24831118
2013 KNL1 facilitates phosphorylation of outer kinetochore proteins by promoting Aurora B kinase activity. The Journal of cell biology 63 24344188
2018 Dysregulation of NCAPG, KNL1, miR-148a-3p, miR-193b-3p, and miR-1179 may contribute to the progression of gastric cancer. Biological research 57 30390708
2013 KNL1: bringing order to the kinetochore. Chromosoma 56 24310619
2015 The RZZ complex requires the N-terminus of KNL1 to mediate optimal Mad1 kinetochore localization in human cells. Open biology 49 26581576
2018 KNL1 Binding to PP1 and Microtubules Is Mutually Exclusive. Structure (London, England : 1993) 42 30100357
2014 The dynamic protein Knl1 - a kinetochore rendezvous. Journal of cell science 41 25052095
2016 Bub3-Bub1 Binding to Spc7/KNL1 Toggles the Spindle Checkpoint Switch by Licensing the Interaction of Bub1 with Mad1-Mad2. Current biology : CB 38 27618268
2019 Robust elimination of genome-damaged cells safeguards against brain somatic aneuploidy following Knl1 deletion. Nature communications 37 31197172
2015 Widespread Recurrent Patterns of Rapid Repeat Evolution in the Kinetochore Scaffold KNL1. Genome biology and evolution 37 26254484
2000 AF15q14, a novel partner gene fused to the MLL gene in an acute myeloid leukaemia with a t(11;15)(q23;q14). Oncogene 36 10980622
2021 Knl1 participates in spindle assembly checkpoint signaling in maize. Proceedings of the National Academy of Sciences of the United States of America 28 33990465
2002 Frequent expression of new cancer/testis gene D40/AF15q14 in lung cancers of smokers. British journal of cancer 28 12087463
2019 Effect of KNL1 on the proliferation and apoptosis of colorectal cancer cells. Technology in cancer research & treatment 17 31315522
2015 Targeted Knockdown of the Kinetochore Protein D40/Knl-1 Inhibits Human Cancer in a p53 Status-Independent Manner. Scientific reports 16 26348410
2024 Kaempferol from Alpinia officinarum hance induces G2/M cell cycle arrest in hepatocellular carcinoma cells by regulating the ATM/CHEK2/KNL1 pathway. Journal of ethnopharmacology 15 38857680
2020 LINC02418 promotes malignant behaviors in lung adenocarcinoma cells by sponging miR-4677-3p to upregulate KNL1 expression. BMC pulmonary medicine 15 32795273
2003 A t(11;15) fuses MLL to two different genes, AF15q14 and a novel gene MPFYVE on chromosome 15. Oncogene 15 12618766
2003 Characterization of the MLL partner gene AF15q14 involved in t(11;15)(q23;q14). Oncogene 13 12618768
2024 A coadapted KNL1 and spindle assembly checkpoint axis orchestrates precise mitosis in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America 10 38170753
2014 The outer kinetochore protein KNL-1 contains a defined oligomerization domain in nematodes. Molecular biology of the cell 9 25411336
2004 The protein encoded by cancer/testis gene D40/AF15q14 is localized in spermatocytes, acrosomes of spermatids and ejaculated spermatozoa. Reproduction (Cambridge, England) 9 15579588
1999 Chromosomal assignment of a novel human gene D40. Nucleic acids symposium series 9 10780384
2024 KNL1 and NDC80 represent new universal markers for the detection of functional centromeres in plants. Chromosome research : an international journal on the molecular, supramolecular and evolutionary aspects of chromosome biology 8 38403686
2008 [The involvement of c-Abl and D40 (AF15q14/CASC5) proteins in the regulation of cell proliferation and cancer]. Tsitologiia 6 18771174
2017 D40/KNL1/CASC5 and autosomal recessive primary microcephaly. Congenital anomalies 4 28901661
2014 Kinetochore signalling: the KIss that MELTs Knl1. Current biology : CB 4 24456977
2025 Arabidopsis KNL1 recruits type one protein phosphatase to kinetochores to silence the spindle assembly checkpoint. Science advances 3 39908360
2024 The kinetochore protein KNL-1 regulates the actin cytoskeleton to control dendrite branching. The Journal of cell biology 3 39625434
2023 Role of Kinetochore Scaffold 1 (KNL1) in Tumorigenesis and Tumor Immune Microenvironment in Pan-Cancer: Bioinformatics Analyses and Validation of Expression. International journal of general medicine 3 37928953
2024 The outer kinetochore components KNL-1 and Ndc80 complex regulate axon and neuronal cell body positioning in the C. elegans nervous system. Molecular biology of the cell 2 38656792
2023 A novel KNL1 intronic splicing variant likely destabilizes the KMN complex, causing primary microcephaly. American journal of medical genetics. Part A 2 37937525
2011 Testis cancer gene D40 expression and its relationship with clinicopathological features in infertile men. International journal of urology : official journal of the Japanese Urological Association 1 21272090
2026 KNL1 Regulates Ferroptosis Resistance and Migration in Lung Adenocarcinoma Cells via AMPK-mTOR Signaling. Oncology research 0 41930176
2026 Proteomics reveals extensive phosphoregulation of outer kinetochore protein KNL1. bioRxiv : the preprint server for biology 0 41959482
2025 The KNL-1/Knl1 outer kinetochore protein caught regulating F-actin. The Journal of cell biology 0 39820668
2002 Isolation of the peri-acrosomal plasma membrane protein D40 enriched fraction from guinea pig spermatozoa using a monoclonal antibody. The Journal of experimental zoology 0 11857474
1999 Isolation of cDNAs that cover the entire coding region of a novel human protein D40. Nucleic acids symposium series 0 10780383