Affinage

EVC

EvC complex member EVC · UniProt P57679

Length
992 aa
Mass
112.0 kDa
Annotated
2026-06-09
40 papers in source corpus 14 papers cited in narrative 14 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 9/9 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

EVC is a transmembrane ciliary protein that functions as an intracellular transducer of Hedgehog signaling, acting downstream of Smoothened but upstream of Gli3 processing to drive transcriptional activation of Hh target genes such as Ptch1 and Gli1 (PMID:17660199). It forms an obligate heterodimeric complex with EVC2, the two proteins binding directly and depending on each other for both ciliary localization and protein stability (PMID:21356043, PMID:23026747). The complex localizes to a signaling microdomain at the base of the primary cilium, the EvC zone, where it is anchored by the EFCAB7-IQCE complex through binding of EFCAB7 to a C-terminal disordered region of EVC2; loss of this tethering mislocalizes EVC-EVC2 and impairs GLI2 activation (PMID:24582806). Upon Hh activation, phosphorylated Smoothened recruits EVC-EVC2 through its C-terminal tail in a cilium-dependent manner (PMID:22986504), and the complex promotes Sufu/Gli3 dissociation and Gli3 trafficking to cilia tips (PMID:23026747); EVC-EVC2 also potentiates Hh output independently of Sufu (PMID:22986504, PMID:23026747). Ciliary entry of EVC, EVC2, and Smoothened requires the IFT-A protein WDR35, distinct from the retrograde IFT motor (PMID:25908617). The complex is regulated post-translationally, with monoubiquitination of the cytosolic tails reducing protein levels and SUMOylation enhancing EvC zone accumulation via increased EFCAB7-IQCE binding (PMID:37576597). Through this Hh-transducing role, EVC governs chondrocyte proliferation and hypertrophy and osteoblast differentiation during endochondral ossification (PMID:21911092), tooth buccal-lingual axis patterning (PMID:23315474), and nucleus pulposus extracellular matrix homeostasis (PMID:41550739). Hypomorphic versus null EVC alleles differ in their ability to complex with EVC2, and the degree of complex formation correlates with the clinical severity of EVC-related skeletal dysplasia (PMID:32906221); dominant-negative EVC2 variants underlie Weyers acrodental dysostosis (PMID:19810119). EVC and EVC2 are also aberrantly overexpressed in subsets of AML, where they support leukemic proliferation through MYC pathway activation independent of Hedgehog signaling (PMID:41249566).

Mechanistic history

Synthesis pass · year-by-year structured walk · 13 steps
  1. 2007 High

    Established EVC as a bona fide intracellular Hedgehog pathway component by placing its action genetically between Smoothened and Gli3 processing, resolving where in the pathway it operates.

    Evidence Evc knockout mouse and Evc-/- cells with basal body immunofluorescence, Gli3 western blot, and qRT-PCR of Ihh targets

    PMID:17660199

    Open questions at the time
    • Molecular partners mediating transduction not yet identified
    • Mechanism of basal body targeting unknown
  2. 2011 High

    Identified EVC2 as the direct, mutually co-dependent partner of EVC, defining the functional unit as a heterodimer rather than a solo protein.

    Evidence Yeast two-hybrid screen with EVC bait, co-immunoprecipitation, immunofluorescence, and subcellular fractionation

    PMID:21356043

    Open questions at the time
    • Stoichiometry and structure of the complex not determined
    • Functional significance of nuclear full-length EVC2 unexplored
  3. 2011 High

    Extended EVC's role beyond chondrocytes to osteoblasts and to both Pthrp-dependent and -independent Ihh functions, demonstrating cell-type-specific deployment within skeletal development.

    Evidence Evc-/- mice with Ptch1-LacZ reporter, osteoblast cilia immunofluorescence, and marker analysis

    PMID:21911092

    Open questions at the time
    • Basis for differential requirement across perichondrial layers unresolved
  4. 2012 High

    Showed that Hh activation drives phosphorylation-dependent binding of EVC-EVC2 to the Smoothened C-terminal tail and that the complex acts upstream of Sufu, mechanistically coupling Smo activation to downstream Gli activation.

    Evidence Co-IP of Smo with Evc/Evc2, constitutively active Smo, and Sufu-/- / Kif3a-/- cell lines with reporter assays

    PMID:22986504

    Open questions at the time
    • Identity of the kinase generating the phospho-dependent binding site not defined
  5. 2012 High

    Defined the downstream output of EVC-EVC2 as promoting Sufu/Gli3 dissociation and Gli3 ciliary tip recruitment, and showed a dominant Weyers Evc2 mutation mislocalizes the complex and reproduces these defects.

    Evidence Evc and Evc2 knockout mice, Co-IP, cilia tip immunofluorescence, Sufu/Gli3 dissociation western blot, and siRNA in Sufu-/- cells

    PMID:23026747

    Open questions at the time
    • Direct biochemical action on the Sufu-Gli3 complex not reconstituted
  6. 2014 High

    Revealed how EVC-EVC2 is spatially confined to the EvC zone, identifying the EFCAB7-IQCE complex as the anchor that binds the Weyers-deleted region of EVC2 and is required for GLI2 activation.

    Evidence Co-IP with domain mapping, siRNA depletion of EFCAB7, and GLI2 activation assays

    PMID:24582806

    Open questions at the time
    • How EvC zone confinement mechanistically enables signaling not fully resolved
  7. 2015 High

    Identified the trafficking route into cilia, showing IFT-A protein WDR35 — but not the retrograde motor — is required for ciliary entry of EVC, EVC2, and SMO.

    Evidence Wdr35-/- vs Dync2h1-/- fibroblasts, cilia localization imaging, pathway assays, and disease cDNA rescue

    PMID:25908617

    Open questions at the time
    • Direct physical interaction between IFT-A and EVC cargo not demonstrated
  8. 2023 Medium

    Established post-translational control of the complex, showing monoubiquitination lowers EVC-EVC2 levels while SUMOylation boosts EvC zone accumulation, and mapped a second EFCAB7-binding motif plus a USP7 interaction.

    Evidence Endogenous EVC interactome mass spectrometry in control vs Evc-null cells, ubiquitination/SUMO assays, and localization imaging

    PMID:37576597

    Open questions at the time
    • Ubiquitin ligase responsible not identified
    • Functional role of the USP7 interaction unresolved
    • Single lab, some findings await replication
  9. 2020 Medium

    Connected genotype to phenotype by showing that EVC allele severity tracks with residual ability to complex with EVC2, providing a molecular basis for clinical variability.

    Evidence Patient fibroblasts, immunoblot/immunofluorescence, and Evc-/- MEF complementation assays

    PMID:32906221

    Open questions at the time
    • Quantitative threshold of complex formation needed for function not established
    • Single lab
  10. 2009 Medium

    Distinguished the disease mechanisms of two EVC2-linked dysplasias, showing Weyers missense variants act dominant-negatively on Hh signaling while a truncating Ellis-van Creveld-type allele does not in the assay.

    Evidence Ectopic expression of murine EVC2 variants in NIH 3T3 cells with Hh reporter assays

    PMID:19810119

    Open questions at the time
    • Overexpression assay without rescue controls
    • Endogenous-level confirmation lacking
  11. 2013 Medium

    Showed EVC is required for spatiotemporal Shh response and buccal-lingual axis patterning in tooth development, broadening its developmental remit.

    Evidence Evc knockout mice with in situ hybridization, Wnt LacZ reporter, and histology of molar development

    PMID:23315474

    Open questions at the time
    • Mechanism linking EVC loss to displaced Wnt activity not defined
    • Single lab
  12. 2025 Medium

    Extended EVC function to nucleus pulposus ECM homeostasis and uncovered TGF-beta suppression of EVC, identifying pathway crosstalk affecting Shh output and fibrosis.

    Evidence Evc knockout mice and CRISPR human NP cells with Gli3 western blot, proteome profiling, and TGF-beta treatment

    PMID:41550739

    Open questions at the time
    • Mechanism of TGF-beta-mediated EVC repression unknown
    • Single lab
  13. 2025 Medium

    Revealed a non-canonical oncogenic role in AML where EVC/EVC2 overexpression supports leukemic proliferation via MYC, independent of Hedgehog signaling.

    Evidence shRNA/CRISPR loss-of-function in AML lines, in vivo transplantation, AML1-ETO ChIP-seq, and MYC pathway expression analysis

    PMID:41249566

    Open questions at the time
    • Direct molecular link between EVC and MYC unresolved
    • Hh-independence requires further confirmation
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the EVC-EVC2 complex biochemically catalyzes Sufu/Gli3 dissociation, the structure of the assembled complex, and the identity of the enzymes controlling its ubiquitination remain unresolved.
  • No reconstituted biochemical mechanism for Sufu/Gli3 dissociation
  • No structural model of the EVC-EVC2 complex
  • Ubiquitin ligase identity unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060089 molecular transducer activity 3 GO:0060090 molecular adaptor activity 3
Localization
GO:0005929 cilium 4 GO:0005815 microtubule organizing center 2 GO:0005886 plasma membrane 1
Pathway
R-HSA-1266738 Developmental Biology 3 R-HSA-162582 Signal Transduction 3
Partners
EFCAB7EVC2IQCESMOUSP7
Complex memberships
EVC-EVC2 complexEvC zone (EVC-EVC2-EFCAB7-IQCE)

Evidence

Reading pass · 14 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2007 EVC protein localizes to the base of the primary cilium (basal body) in chondrocytes, and loss of Evc in mice impairs transcriptional activation of Ihh target genes (Ptch1, Gli1) downstream of Smo but upstream of Gli3 processing, establishing EVC as an intracellular component of the hedgehog signal transduction pathway required for normal transcriptional activation. Immunofluorescence with anti-Evc antibodies; Evc knockout mouse model; in vitro studies with Evc-/- cells; western blot analysis of Gli3 processing; qRT-PCR of Ihh target genes Development (Cambridge, England) High 17660199
2011 EVC2 (Evc2) interacts directly with EVC (Evc) as demonstrated by yeast two-hybrid and co-immunoprecipitation; the two proteins co-localize at the basal body and cilia membrane, and their localization to basal body and cilia is mutually co-dependent. EVC2 is a transmembrane protein with an extracellular portion and intracellular C-terminus, while EVC lacks an extracellular portion. Additionally, full-length EVC2 (but not EVC) is also found in the nucleus. Yeast two-hybrid screen using EVC as bait; co-immunoprecipitation; immunofluorescence with anti-Evc2 antibodies; subcellular fractionation western blot; co-transfection localization studies BMC biology High 21356043
2012 The EVC-EVC2 complex acts downstream of Smo to transduce Hedgehog signal; Hh stimulation induces phosphorylation-dependent binding of EVC/EVC2 to the Smo C-terminal intracellular tail; this binding is abolished in Kif3a-/- cilium-deficient cells. EVC/EVC2 are dispensable for constitutive Gli activity in Sufu-/- cells, indicating they act upstream of Sufu to promote Gli activation. Co-immunoprecipitation (Smo with Evc/Evc2); constitutively active Smo constructs; Sufu-/- and Kif3a-/- cell lines; Smo phosphorylation assays; Hh pathway reporter assays Cell research High 22986504
2012 EVC and EVC2 are mutually required for localizing to primary cilia and for maintaining their normal protein levels. Smo translocation to the cilium is normal in Evc2-/- chondrocytes after Hh activation, but Gli3 recruitment to cilia tips is reduced and Sufu/Gli3 dissociation is impaired. Smo co-precipitates with EVC/EVC2 complex. A dominantly acting Evc2 mutation (Evc2Δ43, Weyers) causes mislocalization of EVC/EVC2Δ43 within the cilium and reproduces Gli3-related molecular defects. Evc silencing in Sufu-/- cells attenuates Hh pathway output, indicating EVC/EVC2 also promote Hh signaling in the absence of Sufu. Evc and Evc2 knockout mouse models; Co-immunoprecipitation; immunofluorescence for cilia tip localization; western blot for Sufu/Gli3 dissociation; siRNA silencing in Sufu-/- cells; SAG (Smo agonist) stimulation assays Human molecular genetics High 23026747
2014 EFCAB7 and IQCE form a complex that anchors EVC-EVC2 in a signaling microdomain at the base of primary cilia (EvC zone). EFCAB7 binds directly to a C-terminal disordered region of EVC2 that is deleted in Weyers syndrome patients. Depletion of EFCAB7 causes mislocalization of EVC-EVC2 within cilia and impairs activation of GLI2, phenocopying the Weyers cellular defect. Co-immunoprecipitation; immunofluorescence; siRNA depletion of EFCAB7; GLI2 activation assays; domain mapping of EFCAB7-EVC2 interaction Developmental cell High 24582806
2011 Evc mediates Hedgehog signaling in osteoblasts (localizing to osteoblast primary cilia) in addition to chondrocytes; Evc promotes chondrocyte proliferation, chondrocyte hypertrophy, and osteoblast differentiation in the perichondrium, implicating it in both Pthrp-dependent and Pthrp-independent Ihh functions. Evc mediates Hh target gene expression in inner perichondrial cells but is dispensable in the external perichondrial layers. Evc-/- mouse model in defined genetic background; Ptch1-LacZ reporter mice; immunofluorescence for osteoblast cilia localization; qRT-PCR and in situ hybridization of osteoblast and Wnt/β-catenin markers Bone High 21911092
2009 Ectopic expression of EVC2 Weyers acrodental dysostosis missense variants (in the final exon of EVC2) in NIH 3T3 cells impairs Hedgehog signal transduction, consistent with a dominant-negative mechanism, whereas expression of a truncated EVC2 protein mimicking an Ellis-van Creveld loss-of-function mutation does not impair Hh signaling in this assay. Ectopic expression of murine EVC2 Weyers variants in NIH 3T3 cells; Hedgehog pathway reporter assays Human mutation Medium 19810119
2013 Evc is required for a spatially and temporally regulated Shh signaling response during molar development; absence of Evc causes progressive loss of Shh pathway response in a buccal-to-lingual direction and is associated with displaced Wnt pathway activity, establishing Evc as necessary for normal buccal-lingual axis patterning during tooth morphogenesis. Evc knockout mouse model; in situ hybridization for Shh target genes; LacZ reporter analysis for Wnt pathway; histological analysis of molar development Journal of dental research Medium 23315474
2015 WDR35 (IFT121), a retrograde intraflagellar transport protein, is specifically required for entry of EVC, EVC2, and Smoothened into the ciliary compartment. In Wdr35-/- cells all three proteins fail to localize to cilia, but this defect is not seen in Dync2h1-/- (retrograde motor mutant) cilia, establishing that IFT-A but not the retrograde IFT motor is required for EVC/EVC2/SMO ciliary targeting. Wdr35-/- and Dync2h1-/- mouse fibroblast cell lines; immunofluorescence for EVC, EVC2, and SMO cilia localization; Hedgehog pathway activity assays; expression of Wdr35 disease cDNAs for rescue Human molecular genetics High 25908617
2023 The EVC-EVC2 complex is subject to ubiquitination and SUMOylation of its cytosolic tails. Monoubiquitination of EVC-EVC2 cytosolic tails reduces protein levels. SUMOylation with SUMO3 enhances EVC-EVC2 accumulation at the EvC zone at the base of cilia, likely via increased binding to the EFCAB7-IQCE complex. A second EFCAB7-binding motif in EVC2's Weyers-deleted peptide was mapped, in addition to the previously characterized one. Known interactors EVC2, IQCE, EFCAB7 were confirmed, and USP7 (a deubiquitinase involved in Hh signaling) was identified as a new interactor, though ubiquitination of EVC-EVC2 was found to be independent of USP7 and USP48. Endogenous EVC protein interactome by mass spectrometry in control and Evc-null cells; ubiquitination assays; SUMO modification assays; immunofluorescence for EvC zone localization; domain mapping of EFCAB7-binding motifs Frontiers in cell and developmental biology Medium 37576597
2020 Hypomorphic EVC missense mutations (p.Arg663Pro and splice variant c.1316-7A>G) produce proteins that retain partial ability to complex with EVC2, whereas the p.Arg622Ter nonsense mutation is a complete loss-of-function allele that cannot complex with EVC2, establishing that the degree of EVC-EVC2 complex formation correlates with clinical severity. Patient-derived fibroblasts; complementary DNA sequencing; immunoblot; immunofluorescence; Evc-/- mouse embryonic fibroblast complementation assays Human mutation Medium 32906221
2025 EVC protein facilitates Shh signaling in notochord-derived nucleus pulposus cells; loss of EVC reduces Gli3 processing, impairs Shh pathway activity, and alters extracellular matrix organization toward fibrosis. TGF-β signaling suppresses EVC expression, indicating crosstalk between TGF-β and Hedgehog pathways at the level of EVC. Evc knockout mouse model; CRISPR-engineered human NP cells; proteome profiling; Gli3 processing western blot; ECM composition analysis; TGF-β treatment assays iScience Medium 41550739
2025 EVC and EVC2 are aberrantly overexpressed in a subset of AMLs (particularly those with ASXL1 mutations or t(8;21) translocation); functional loss of EVC/EVC2 impairs leukemia cell proliferation, promotes differentiation, and blocks AML progression in vivo. The leukemogenic role of EVC/EVC2 is mediated through MYC pathway activation, independent of their canonical role in Hedgehog signaling. shRNA/CRISPR loss-of-function in AML cell lines; in vivo AML transplantation models; ChIP-seq for AML1-ETO occupancy at EVC/EVC2 promoters; chromatin interaction analysis; MYC pathway gene expression analysis Leukemia Medium 41249566
2004 Evc protein is expressed in prehypertrophic and hypertrophic chondrocytes of the epiphyseal growth plate in rat, as determined by immunohistochemistry and in situ hybridization, consistent with a function in chondrocyte differentiation during endochondral ossification. Immunohistochemistry with anti-Evc antibodies; in situ hybridization; RT-PCR on growth plate tissue The anatomical record. Part A, Discoveries in molecular, cellular, and evolutionary biology Low 15278943

Source papers

Stage 0 corpus · 40 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 Evc is a positive mediator of Ihh-regulated bone growth that localises at the base of chondrocyte cilia. Development (Cambridge, England) 156 17660199
2012 The ciliary Evc/Evc2 complex interacts with Smo and controls Hedgehog pathway activity in chondrocytes by regulating Sufu/Gli3 dissociation and Gli3 trafficking in primary cilia. Human molecular genetics 94 23026747
2012 Smoothened transduces Hedgehog signal by forming a complex with Evc/Evc2. Cell research 86 22986504
2006 Sequencing EVC and EVC2 identifies mutations in two-thirds of Ellis-van Creveld syndrome patients. Human genetics 80 17024374
2014 EFCAB7 and IQCE regulate hedgehog signaling by tethering the EVC-EVC2 complex to the base of primary cilia. Developmental cell 77 24582806
2011 Evc2 is a positive modulator of Hedgehog signalling that interacts with Evc at the cilia membrane and is also found in the nucleus. BMC biology 77 21356043
2012 Novel and recurrent EVC and EVC2 mutations in Ellis-van Creveld syndrome and Weyers acrofacial dyostosis. European journal of medical genetics 59 23220543
2009 Widening the mutation spectrum of EVC and EVC2: ectopic expression of Weyer variants in NIH 3T3 fibroblasts disrupts Hedgehog signaling. Human mutation 52 19810119
2013 Evc regulates a symmetrical response to Shh signaling in molar development. Journal of dental research 50 23315474
2015 Specific variants in WDR35 cause a distinctive form of Ellis-van Creveld syndrome by disrupting the recruitment of the EvC complex and SMO into the cilium. Human molecular genetics 49 25908617
2011 Evc works in chondrocytes and osteoblasts to regulate multiple aspects of growth plate development in the appendicular skeleton and cranial base. Bone 36 21911092
2008 Extending the spectrum of Ellis van Creveld syndrome: a large family with a mild mutation in the EVC gene. BMC medical genetics 22 18947413
2017 Ellis-van Creveld syndrome and profound deafness resulted by sequence variants in the EVC/EVC2 and TMC1 genes. Journal of genetics 19 29321360
2016 Truncation and microdeletion of EVC/EVC2 with missense mutation of EFCAB7 in Ellis-van Creveld syndrome. Congenital anomalies 12 26748586
2020 Common atrium/atrioventricular canal defect and postaxial polydactyly: A mild clinical subtype of Ellis-van Creveld syndrome caused by hypomorphic mutations in the EVC gene. Human mutation 11 32906221
2014 Complete coding regions of the prototypes enterovirus B93 and C95: phylogenetic analyses of the P1 and P3 regions of EV-B and EV-C strains. Journal of medical virology 9 25163640
2023 Establishing an objective clinical spectrum, genotype-phenotype correlations, and CRMP1 as a modifier in the Ellis-van Creveld syndrome: The first systematic review of EVC- and EVC2-associated conditions. Genetics in medicine open 8 39669252
2022 Blocking chondrocyte hypertrophy in conditional Evc knockout mice does not modify cartilage damage in osteoarthritis. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 8 35334131
2016 Novel homozygous mutations in the EVC and EVC2 genes in two consanguineous families segregating autosomal recessive Ellis-van Creveld syndrome. Clinical dysmorphology 8 26580685
2009 A novel missense mutation in the EVC gene underlies Ellis-van Creveld syndrome in a Pakistani family. Pediatrics international : official journal of the Japan Pediatric Society 8 19744229
2023 EVC-EVC2 complex stability and ciliary targeting are regulated by modification with ubiquitin and SUMO. Frontiers in cell and developmental biology 7 37576597
2017 Phenotypic Variation in Patients with Homozygous c.1678G>T Mutation in EVC Gene: Report of Two Mexican Families with Ellis-van Creveld Syndrome. The American journal of case reports 7 29229899
2019 Identification of a novel EVC variant in a Han-Chinese family with Ellis-van Creveld syndrome. Molecular genetics & genomic medicine 6 31338997
2001 Automatic analysis of image of surface structure of cell wall-deficient EVC. Bio-medical materials and engineering 6 11564900
2023 A homozygous EVC mutation in a prenatal fetus with Ellis-van Creveld syndrome. Molecular genetics & genomic medicine 5 37157924
2015 Novel mutations in EVC cause aberrant splicing in Ellis-van Creveld syndrome. Molecular genetics and genomics : MGG 3 26621368
2004 Expression of the Ellis-van Creveld (Evc) gene in the rat tibial growth plate. The anatomical record. Part A, Discoveries in molecular, cellular, and evolutionary biology 2 15278943
2024 Shugan Tongluo Qiangjing recipe protects against varicocele of EVC rats through modulating sperm DNA damage, telomere expression and oxidative stress. Tissue & cell 1 38865824
2022 Novel large deletion involving EVC and EVC2 in Ellis-van Creveld syndrome. Human genome variation 1 35581188
2006 [Inhibition of rhodiola on the growth of EVC-304 cell line]. Xi bao yu fen zi mian yi xue za zhi = Chinese journal of cellular and molecular immunology 1 16806023
2004 Haplotype and linkage disequilibrium analysis of the CRMP1 and EVC genes. International journal of molecular medicine 1 15492864
2026 Aberrant Proliferation and Cell Fate Underlie Oral Defects in a Mouse Model of EvC Syndrome. Journal of dental research 0 41913599
2026 Identification of EVC variants and the preimplantation genetic testing in a Chinese family. Frontiers in medicine 0 42200062
2025 Oncogenic activation of EVC/EVC2 defines a therapeutically targetable subset of acute myeloid leukemia. Leukemia 0 41249566
2025 Case Report of a Novel EVC Gene Mutation in Ellis-van Creveld Syndrome: Implications for Pediatric Dental Management. Case reports in dentistry 0 41438176
2025 EVC protein regulates Sonic hedgehog signaling during human intervertebral disc development and degeneration. iScience 0 41550739
2025 Mandibular morphogenesis: Genetic insights into malocclusion from EVC perspective. Journal of oral biosciences 0 41714013
2024 Ellis-Van Creveld Syndrome with Severe Mitral Valve Insufficiency Caused by a Homozygous Intragenic Deletion of the EVC Gene. Molecular syndromology 0 40475170
2015 EVC gene polymorphisms and risks of isolated hypospadias - a preliminary study. Central European journal of urology 0 26251756
2005 [From gene to disease; EVC, EVC2, and Ellis-van Creveld syndrome]. Nederlands tijdschrift voor geneeskunde 0 15884406

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