Affinage

FOXC1

Forkhead box protein C1 · UniProt Q12948

Length
553 aa
Mass
56.8 kDa
Annotated
2026-06-09
100 papers in source corpus 45 papers cited in narrative 45 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

FOXC1 is a forkhead-domain transcription factor that directs cell-type-specific developmental programs and is co-opted in cancer, acting through direct promoter/enhancer binding and combinatorial protein-protein interactions (PMID:11782474, PMID:34551306, PMID:33501917). Its activity is governed by a modular architecture: N- and C-terminal activation domains flank a central phosphorylated inhibitory domain (residues 215–366), and forkhead-domain residues control DNA binding and nuclear localization (PMID:11782474, PMID:14578375). Stability and output are set by post-translational control — p38 MAPK phosphorylates Ser241/Ser272 to block ubiquitin-mediated degradation, with PP2A reversing this modification (PMID:31650548). FOXC1 partners with developmental cofactors to tune transcription, interacting with PITX2A and p32, both of which negatively regulate its transactivation, and with Gli2 to drive Smoothened-independent Hedgehog target gene expression in endochondral ossification and breast cancer (PMID:16449236, PMID:18676636, PMID:26565916, PMID:25808752). As a pioneer-type regulator it establishes chromatin accessibility at lineage-specific enhancers co-bound with SOX9 in cartilage and at H3K4me2-marked corneal epithelial enhancers, with loss causing fate reprogramming (PMID:33414365, PMID:33501917). In adult tissue homeostasis FOXC1 maintains the haematopoietic stem cell niche by sustaining CXCL12/SCF in CAR cells, enforces hair follicle stem cell quiescence via Nfatc1/BMP, and controls vascular development through suppression of ERK and ROCK signaling and activation of CD98/mTOR and CXCL12/RSPO3 programs (PMID:24590069, PMID:26912704, PMID:27214551, PMID:32510325, PMID:37154714, PMID:38755144). In cancer FOXC1 drives invasion, stemness, and metabolic reprogramming through targets including WNT5A, β-catenin, FBP1, and serine-synthesis enzymes, and cooperates with RUNX1/TLE3/HDAC1 to repress differentiation enhancers in AML (PMID:29249801, PMID:30189871, PMID:30171256, PMID:34551306, PMID:39773485). Mutations in FOXC1 cause dominant glaucoma and Axenfeld-Rieger spectrum disease through both gain- and loss-of-function mechanisms affecting stability, DNA binding, and partner interaction (PMID:18676636, PMID:25808752, PMID:25786029).

Mechanistic history

Synthesis pass · year-by-year structured walk · 42 steps
  1. 2002 High

    Established the domain logic of FOXC1 — how a single factor can be switched between active and inhibited states — by defining its activation domains, a phosphorylation-controlled inhibitory domain, and nuclear-localization determinants.

    Evidence Deletion mutagenesis, reporter transactivation, nuclear localization and phosphorylation analysis in cells

    PMID:11782474

    Open questions at the time
    • Did not identify the kinase or specific residues responsible for inhibitory-domain phosphorylation
    • No structural model of the domains
  2. 2002 Medium

    Placed FOXC1 downstream of TGF-β1 growth-inhibitory signaling by showing it is TGF-β1-inducible and restores G0/G1 arrest in FOXC1-null cells.

    Evidence Cloning from TGF-β1-treated cells plus cDNA rescue and cell-cycle analysis in HeLa

    PMID:12408963

    Open questions at the time
    • Direct transcriptional targets mediating arrest not identified
    • Single cell-line system
  3. 2003 Medium

    Demonstrated that forkhead helix 1 residues are critical for FOXC1 DNA binding and transactivation independent of nuclear import, linking disease missense mutations to a defined molecular defect.

    Evidence Site-directed mutagenesis with DNA-binding, localization and reporter assays

    PMID:14578375

    Open questions at the time
    • In vitro only
    • Did not test endogenous target genes
  4. 2006 High

    Identified PITX2A as a direct FOXC1 partner that negatively regulates its transactivity, placing the two factors in a common anterior segment morphogenesis pathway.

    Evidence Co-IP, colocalization, domain-deletion and reporter assays

    PMID:16449236

    Open questions at the time
    • Mechanism of repression (sequestration vs. complex formation) not resolved
    • No genome-wide target consequence
  5. 2006 High

    Showed FOXC1 directly activates FGF19 to drive FGFR4-MAPK signaling in ocular mesenchyme, providing a first direct developmental target.

    Evidence Chromatin enrichment, overexpression/knockdown in cell culture and zebrafish, MAPK assays

    PMID:17000708

    Open questions at the time
    • Binding-site sequence within FGF19 not precisely mapped
  6. 2007 High

    Defined a homeostatic role by showing FOXC1 directly regulates FOXO1A to protect cells from oxidative stress, linking FOXC1 dysfunction to cellular death.

    Evidence Inducible expression with protein-synthesis blockade, ChIP, knockdown in human cells and zebrafish, oxidative-stress viability assays

    PMID:17993506

    Open questions at the time
    • Did not connect FOXO1A regulation to in vivo glaucoma pathology
  7. 2008 High

    Identified p32 as a forkhead-domain partner that inhibits FOXC1 transactivation without blocking DNA binding, and showed a disease mutation disrupts this interaction.

    Evidence Y2H, pull-down, Co-IP, EMSA and reporter assays in trabecular meshwork cells

    PMID:18676636

    Open questions at the time
    • Mechanism by which p32 dampens activity downstream of DNA binding unknown
  8. 2004 High

    Established FOXC1/FOXC2 redundant control of paraxial versus intermediate mesoderm fate via loss- and gain-of-function across two species.

    Evidence Mouse double-knockout and chick electroporation with marker in situ hybridization

    PMID:15196959

    Open questions at the time
    • Direct transcriptional targets in mesoderm specification not identified
  9. 2005 Medium

    Positioned Foxc1 as an integrator of FGF and BMP signaling in calvarial osteogenesis required for BMP regulation of Msx2.

    Evidence Foxc1-null mouse analysis with FGF2 treatment and osteogenic differentiation assays

    PMID:15906377

    Open questions at the time
    • Whether Foxc1 acts directly on Msx2 not shown
    • Single lab
  10. 2013 Medium

    Showed Foxc1 patterns neural-crest-derived jaw osteogenesis through genetic interaction with Fgf8, preventing ectopic bone fusion.

    Evidence Foxc1-null phenotyping with Fgf8 compound genetics and osteogenic marker in situ

    PMID:24385915

    Open questions at the time
    • Direct Foxc1 targets in jaw patterning not defined
  11. 2014 High

    Defined Foxc1 as a transcriptional regulator of the haematopoietic stem cell niche by maintaining CXCL12/SCF in CAR mesenchymal cells and preventing marrow adipogenesis.

    Evidence Conditional/inducible Foxc1 knockout, flow cytometry, qPCR/ISH and histology in mice

    PMID:24590069

    Open questions at the time
    • Direct binding of Foxc1 to Cxcl12/Scf loci not shown in this study
  12. 2014 High

    Showed Foxc1 directly activates SDF1α in head mesenchyme to drive cerebellar radial glial proliferation and Purkinje migration, validated by Cxcr4-receptor epistasis and SDF1α rescue.

    Evidence Foxc1-null cerebellar phenotyping, conditional Cxcr4 KO and SDF1α rescue

    PMID:25513817

    Open questions at the time
    • Other Foxc1 targets in cerebellar development not explored
  13. 2014 High

    Demonstrated foxc1 is required upstream of acta2 for vascular smooth muscle differentiation from head mesenchyme, distinguishing vSMC from pericyte programs.

    Evidence Live imaging, CRISPR/morpholino loss-of-function and RNA-seq in zebrafish

    PMID:31199900

    Open questions at the time
    • Whether foxc1 binds acta2 regulatory elements directly not shown
  14. 2015 High

    Showed FOXC1 directly interacts with Gli2 to activate Hedgehog target genes independently of Smoothened, a mechanism shared between endochondral ossification and basal-like breast cancer stemness.

    Evidence Domain-mapping pulldowns, Co-IP, ChIP, rescue and xenograft/mouse models across two studies

    PMID:25808752 PMID:26565916

    Open questions at the time
    • Structural basis of the FOXC1-Gli2 interface not resolved
  15. 2011 High

    Identified upstream repression of FOXC1 by a BRCA1-GATA3 co-repressor complex at the distal promoter, connecting FOXC1 deregulation to breast cancer.

    Evidence Co-IP, ChIP, reporter assays and mutant BRCA1 analysis

    PMID:22120723

    Open questions at the time
    • Repression is not FOXC1-specific; selectivity determinants unknown
  16. 2016 High

    Established Foxc1 as an enforcer of hair follicle stem cell quiescence by activating Nfatc1 and BMP signaling specifically in activated stem cells.

    Evidence Stage-specific conditional deletion, lineage tracing and hair-cycle phenotyping

    PMID:26912704

    Open questions at the time
    • Direct vs. indirect activation of Nfatc1/BMP components not fully separated
  17. 2016 Medium

    Showed Foxc1 mediates BMP4-SMAD signaling in osteogenic progenitors, with SMAD binding the Foxc1 promoter and Foxc1 restraining premature osteoblast differentiation.

    Evidence BMP4 treatment, SMAD ChIP, reporter and siRNA differentiation assays

    PMID:26666591

    Open questions at the time
    • Downstream Foxc1 osteogenic targets not identified
    • Single lab
  18. 2016 Medium

    Implicated Foxc1 in early cardiomyogenic specification, controlling Isl1/Mef2c/Nkx2.5 and directly targeting Myh7 within a defined differentiation window.

    Evidence ESC differentiation KD/OE, RNA-seq and electrophysiology

    PMID:26824887

    Open questions at the time
    • Direct binding confirmed only for Myh7
    • Single in vitro model
  19. 2016 High

    Defined FOXC1/FOXC2 control of lymphangiogenesis through suppression of ERK signaling, with ERK inhibition rescuing the lymphatic phenotype in vivo.

    Evidence LEC-specific conditional knockout, ERK phosphorylation profiling and in utero ERK inhibitor rescue

    PMID:27214551

    Open questions at the time
    • Direct targets controlling Ras-regulator expression not pinned down
  20. 2017 High

    Mapped a FOXC1-driven WNT5A-NF-κB-MMP7 invasion axis in triple-negative breast cancer and identified hnRNPs as nuclear FOXC1 partners.

    Evidence ChIP, enChIP-MS, genetic rescue and xenograft metastasis models

    PMID:29249801

    Open questions at the time
    • Functional role of hnRNP interaction in transcription not fully dissected
  21. 2017 Medium

    Showed NF-κB-p65 directly binds and activates the FOXC1 promoter downstream of EGF/EGFR, linking inflammatory/growth signaling to FOXC1 expression in basal-like breast cancer.

    Evidence Reporter, EMSA, oligonucleotide pulldown, ChIP and NF-κB knockdown

    PMID:28629477

    Open questions at the time
    • Single lab
    • In vivo relevance not tested
  22. 2017 Medium

    Showed FOXC1 directly activates an FGFR1-IIIc isoform-specific promoter element following TGFβ1-induced EMT to promote invasive migration.

    Evidence RNA-seq isoform analysis, ChIP, reporter and FGFR1 rescue with 3D migration assays

    PMID:28684636

    Open questions at the time
    • Mechanism of isoform-specific selection not defined
    • Single lab
  23. 2017 Medium

    Established that FOXC1 loss reduces retinal ganglion cells through impaired atoh7-dependent differentiation rather than survival defects.

    Evidence CRISPR/morpholino foxc1 loss-of-function in zebrafish with marker analysis

    PMID:30684501

    Open questions at the time
    • Whether foxc1 binds atoh7 regulatory elements directly not shown
  24. 2018 Medium

    Showed FOXC1 directly activates β-catenin transcription to confer cancer stem-cell properties in NSCLC, defined by epistasis rescue.

    Evidence Reporter, ChIP and β-catenin rescue with sphere/tumorigenicity assays

    PMID:30189871

    Open questions at the time
    • Single lab
    • In vivo β-catenin dependence not fully tested
  25. 2018 Medium

    Identified FOXC1 as a direct repressor of FBP1 that reprograms glycolysis (Warburg effect) to fuel colorectal cancer proliferation.

    Evidence ChIP, reporter, metabolic assays and xenograft

    PMID:30171256

    Open questions at the time
    • Single lab
    • Cofactors mediating repression not identified
  26. 2019 High

    Defined the stability switch for FOXC1: p38 phosphorylation of Ser241/Ser272 blocks ubiquitin-mediated degradation, with PP2A reversing it, stabilizing FOXC1 to drive metastatic target genes.

    Evidence Phospho-site mutagenesis, Co-IP, ubiquitination/dephosphorylation assays and ChIP

    PMID:31650548

    Open questions at the time
    • E3 ubiquitin ligase for FOXC1 not identified
  27. 2019 Medium

    Showed FOXC1 directly activates TLR3/TLR4 promoters and exacerbates inflammation and infarct injury, linking FOXC1 to innate immune signaling.

    Evidence Reporter, ChIP, OE/KD and mouse myocardial ischemia model

    PMID:31517441

    Open questions at the time
    • Single lab
    • Mechanism of TLR-driven cardiac injury indirect
  28. 2020 High

    Defined FOXC1/FOXC2 as laminar-shear-stress effectors restraining ROCK activity to maintain endothelial cytoskeletal and junctional integrity, with ROCK inhibition rescuing in vivo valve defects.

    Evidence Shear-stress culture, endothelial conditional KO, siRNA and ROCK-inhibitor rescue

    PMID:32510325

    Open questions at the time
    • Direct transcriptional link between FOXC1 and ROCK regulators not mapped
  29. 2020 Medium

    Showed EZH2-mediated H3K27me3 represses FOXC1, with OGT modulating EZH2 occupancy and histone acetylation at the FOXC1 locus in breast cancer.

    Evidence ChIP, EZH2 inhibitors and OGT siRNA with chromatin-mark analysis

    PMID:33057161

    Open questions at the time
    • Single lab
    • Context-dependence of OGT effect not mechanistically resolved
  30. 2020 Medium

    Placed FOXC1 upstream of a LINC00301-EZH2-EAF2-pVHL-HIF1α regulatory cascade in NSCLC via direct binding to the LINC00301 promoter.

    Evidence ChIP, RNA pulldown, domain mapping, RIP, EMSA and reporter

    PMID:32878637

    Open questions at the time
    • Single lab
    • In vivo validation of the full cascade limited
  31. 2021 High

    Revealed FOXC1 as a chromatin organizer that interacts with RUNX1 (Forkhead-Runt domains) to stabilize RUNX1/HDAC1/TLE3 repressor complexes at differentiation enhancers, blocking AML differentiation.

    Evidence MS proteomics, ChIP-seq, ATAC-seq, domain-mapped Co-IP and KD transcriptomics

    PMID:34551306

    Open questions at the time
    • How FOXC1 directs repressor redistribution between enhancers and promoters mechanistically unresolved
  32. 2021 High

    Demonstrated FOXC1 functions as a lineage pioneer at H3K4me2-marked corneal enhancers, with loss reprogramming corneal epithelium toward skin and abolishing IRF1/PAX6.

    Evidence ATAC-seq, H3K4me2 ChIP-seq, KD and patient-sample transcriptomics

    PMID:33414365

    Open questions at the time
    • Whether FOXC1 binding precedes or follows enhancer marking not temporally resolved
  33. 2021 High

    Established that Foxc1, downstream of SOX9, establishes chromatin accessibility at Sox9/Fox co-bound cartilage enhancers to drive chondrogenesis, validated by conditional KO and transgenic enhancer assays.

    Evidence SOX9-promoter regulation, conditional chondrocyte KO with RNA-seq, ATAC-seq and transgenic enhancer reporters across mouse and zebrafish

    PMID:33501917 PMID:34331943

    Open questions at the time
    • Whether Foxc1 acts as a true pioneer factor opening closed chromatin vs. maintaining accessibility not fully separated
  34. 2021 Medium

    Showed FOXC1 promotes Wnt signaling both by repressing DKK1 transcription and by forming a cytoplasmic complex with unphosphorylated β-catenin to drive its nuclear translocation and c-MYC expression in gastric cancer.

    Evidence ChIP, Co-IP, reporter and KD/OE Wnt readouts

    PMID:33987183

    Open questions at the time
    • Single lab
    • Structural basis of cytoplasmic FOXC1-β-catenin complex unknown
  35. 2021 High

    Defined an HCC feed-forward loop in which FOXC1 activates DNMT3B to silence CTH, raising ROS that activates ERK1/2-ELK1 to further induce FOXC1.

    Evidence ChIP, bisulfite sequencing, ROS assays, CTH rescue and xenograft metastasis

    PMID:33522955

    Open questions at the time
    • Direct DNMT3B binding-site within FOXC1-target promoter not mapped
  36. 2023 High

    Showed FOXC1/FOXC2 directly bind CXCL12 and RSPO3 regulatory elements in blood vs. lymphatic endothelium to drive Wnt/ISC-mediated intestinal regeneration after ischemia-reperfusion.

    Evidence EC/LEC-specific conditional KO, intestinal I/R model, ChIP and cytokine rescue

    PMID:37154714

    Open questions at the time
    • Cell-type specificity of CXCL12 vs RSPO3 target choice not mechanistically explained
  37. 2023 Medium

    Identified a Sirt1/Foxc1/Ubqln1 proteostasis axis in cerebral ischemia, where Sirt1 deacetylates EZH2 to derepress Foxc1, which then activates Ubqln1.

    Evidence Co-IP, ChIP, reporter, siRNA and OGD/R and MCAO/R models

    PMID:38452414

    Open questions at the time
    • Single lab
    • Direct Foxc1 binding to Ubqln1 mapped but axis tested in limited models
  38. 2023 Medium

    Showed FOXC1 overexpression suppresses microglial neuroinflammation in sepsis-associated encephalopathy by elevating IκBα and inhibiting NF-κB-p65.

    Evidence BV-2 LPS model, CLP mouse model, cognitive testing and IκBα-inhibitor rescue

    PMID:35103290

    Open questions at the time
    • Whether FOXC1 directly activates IκBα transcription not established
    • Single lab
  39. 2023 Low

    Showed O-GlcNAcylation stabilizes FOXC1 and that icaritin reduces this modification to suppress endometrial cancer growth.

    Evidence Western blot for O-GlcNAcylated FOXC1, icaritin treatment, KD/OE and xenograft

    PMID:37683586

    Open questions at the time
    • O-GlcNAc site inferred from western blot without mass-spectrometry residue identification
    • Single lab
  40. 2024 High

    Defined FOXC1 control of endothelial CD98/LAT1 amino-acid transport and mTOR activation required for retinal angiogenesis and blood-retina barrier formation, validated by pharmacological mTOR rescue.

    Evidence EC-specific conditional KO, retinal flat-mount analysis, Slc3a2/Slc7a5 expression, mTOR agonist rescue and oxygen-induced retinopathy model

    PMID:38755144

    Open questions at the time
    • Direct FOXC1 binding to Slc3a2/Slc7a5 loci not detailed
  41. 2024 Medium

    Linked telomere shortening to FOXC1 derepression: telomeric chromatin decompaction increases FOXC1 accessibility, and elevated FOXC1 induces cardiomyocyte aging and contractile dysfunction.

    Evidence Hi-C, RNA-seq, isogenic hiPSC-CM telomere model and OE/KD with patient correlation

    PMID:38634789

    Open questions at the time
    • FOXC1 target genes driving cardiomyocyte aging not defined
    • Single lab
  42. 2025 Medium

    Showed serine deprivation activates ERK1/2-ELK1 to induce FOXC1, which transactivates serine-synthesis enzymes PHGDH/PSAT1/PSPH to support 5-FU resistance in colorectal cancer.

    Evidence ChIP, reporter, serine-deprivation model, pathway inhibitors and 5-FU resistance assays

    PMID:39773485

    Open questions at the time
    • Single lab
    • Clinical relevance of the resistance mechanism not validated

Open questions

Synthesis pass · forward-looking unresolved questions
  • How FOXC1 selects distinct target gene sets and cofactors in different cell types, and what determines its pioneer activity at closed versus marked chromatin, remains unresolved.
  • No structural model of FOXC1-cofactor or FOXC1-DNA complexes
  • E3 ligase and full PTM code controlling FOXC1 stability incompletely defined
  • Mechanism distinguishing pioneer establishment vs maintenance of accessibility unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 9 GO:0003677 DNA binding 4 GO:0140097 catalytic activity, acting on DNA 1
Localization
GO:0005634 nucleus 2 GO:0005829 cytosol 1
Pathway
R-HSA-1266738 Developmental Biology 6 R-HSA-74160 Gene expression (Transcription) 6 R-HSA-162582 Signal Transduction 5 R-HSA-1643685 Disease 5 R-HSA-4839726 Chromatin organization 4 R-HSA-1430728 Metabolism 3 R-HSA-168256 Immune System 2

Evidence

Reading pass · 45 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 FOXC1 transcriptional regulation is mediated by two activation domains (extreme N- and C-terminal regions) and a central phosphorylated transcriptional inhibitory domain (residues 215-366). Two regions in the forkhead domain are necessary for nuclear localization, with only the basic amino-acid-rich region being sufficient. FOXC1 is a phosphoprotein and phosphorylation of residues in the inhibitory domain reduces transactivation; removal of the inhibitory domain produces a hyperactive, hypophosphorylated protein. Deletion mutagenesis, reporter-gene transactivation assays, nuclear-localization assays, phosphorylation analysis in cell-based experiments The Journal of biological chemistry High 11782474
2006 FOXC1 and PITX2A physically interact; this interaction requires the C-terminal activation domain of FOXC1 and the homeodomain of PITX2A. PITX2A and FOXC1 colocalize within a common nuclear subcompartment. PITX2A negatively regulates FOXC1 transactivity, placing both proteins in a common developmental pathway for anterior segment morphogenesis. Co-immunoprecipitation, immunofluorescence colocalization, reporter-gene transactivation assays, domain-deletion analysis Human molecular genetics High 16449236
2007 FOXC1 directly regulates FOXO1A transcription by binding to a conserved element in the FOXO1A promoter in vivo. Reduced FOXC1 expression decreases FOXO1A expression in human trabecular meshwork cells and zebrafish eyes, and increases cell death in response to oxidative stress, establishing FOXC1 as a mediator of cellular homeostasis through FOXO1A. Inducible FOXC1 expression with protein synthesis blockade (to identify direct targets), ChIP at the FOXO1A promoter, siRNA and morpholino knockdown, cell viability/death assays under oxidative stress Human molecular genetics High 17993506
2006 FGF19 is a direct transcriptional target of FOXC1 in the eye. FOXC1 positively regulates FGF19 expression in corneal and periocular mesenchymal cells; FGF19 then signals through FGFR4 to promote MAPK phosphorylation in the developing and mature cornea. Chromatin enrichment (nickel-agarose), cell-culture overexpression/knockdown, zebrafish embryo experiments, MAPK phosphorylation assays Human molecular genetics High 17000708
2008 Human p32 protein interacts with FOXC1 via the forkhead domain; p32 inhibits FOXC1-mediated transcriptional activation in a dose-dependent manner without affecting FOXC1 DNA-binding. The disease-causing F112S FOXC1 mutation impairs interaction with p32, suggesting disrupted protein-protein interaction as a disease mechanism. Yeast two-hybrid screen, Ni2+ pull-down, co-immunoprecipitation, immunofluorescence colocalization, EMSA, dual-luciferase reporter assays in human trabecular meshwork cells Investigative ophthalmology & visual science High 18676636
2003 The L86F missense mutation in helix 1 of the FOXC1 forkhead domain does not affect nuclear localization but reduces DNA binding and significantly reduces transcriptional transactivation. The analogous L86P mutation more severely disrupts function, establishing the importance of helix 1 for FOXC1 structure and activity. Site-directed mutagenesis, nuclear localization assays, DNA-binding assays, transactivation reporter assays Investigative ophthalmology & visual science Medium 14578375
2015 FOXC1 activates Smoothened-independent Hedgehog signaling in basal-like breast cancer by direct physical interaction between the N-terminal domain of FOXC1 (aa 1-68) and an internal region (aa 898-1168) of Gli2. This interaction enhances Gli2's DNA-binding and transcription-activating capacity, promoting cancer stem cell properties. Domain-mapping pulldown assays, reporter assays, ChIP, FOXC1 knockdown/overexpression with Gli2 rescue, xenograft models Cell reports High 26565916
2017 FOXC1 directly binds the WNT5A promoter to activate its expression. WNT5A then activates NF-κB signaling to induce MMP7 expression, constituting a non-canonical WNT5A-NF-κB-MMP7 pathway essential for FOXC1-driven invasiveness in triple-negative breast cancer. enChIP-MS identified heterogeneous nuclear ribonucleoproteins as FOXC1-interacting proteins involved in WNT5A transcription. Pathway PCR array, ChIP (direct binding to WNT5A promoter), enChIP coupled with mass spectrometry, WNT5A and MMP7 KO/OE rescue experiments, xenograft lung metastasis models Oncogene High 29249801
2011 BRCA1 and GATA3 form a transcriptional co-repressor complex on the FOXC1 distal promoter to repress FOXC1 expression; BRCA1 requires GATA3 for recruitment to this region. A mutant BRCA1 protein cannot localize to the FOXC1 promoter or repress FOXC1 expression. This repression is not FOXC1-specific, as FOXC2, CXCL1, and p-cadherin are co-repressed similarly. Co-immunoprecipitation of BRCA1-GATA3, ChIP at the FOXC1 promoter, reporter assays, mutant BRCA1 functional analysis Oncogene High 22120723
2015 FOXC1 interacts physically and functionally with Gli2 to stimulate expression of Ihh target genes (PTHrP and Col10a1) during endochondral ossification. An Axenfeld-Rieger syndrome-associated missense mutation in FOXC1 impairs the Gli2-FOXC1 association and Ihh function. Loss of Foxc1 function in Foxc1(ch/ch) mice delays endochondral ossification and reduces Ihh target gene expression. In vivo microarray, co-immunoprecipitation of FOXC1-Gli2, dominant-negative FOXC1 overexpression, Foxc1 hypomorphic mouse model, ARS missense mutation functional analysis Nature communications High 25808752
2014 Foxc1 is preferentially expressed in CAR (CXCL12-abundant reticular) mesenchymal progenitor cells in bone marrow. Conditional deletion of Foxc1 in mesenchymal or CAR cells depletes haematopoietic stem/progenitor cells, reduces CXCL12 and stem cell factor expression in CAR cells, and causes adipocytic conversion of marrow, establishing Foxc1 as a transcriptional regulator of the haematopoietic stem cell niche. Conditional and inducible Cre-mediated Foxc1 knockout in mice, flow cytometry of HSPCs, qPCR/in situ hybridization for CXCL12/SCF, histological analysis Nature High 24590069
2016 Foxc1 reinforces quiescence in self-renewing hair follicle stem cells by transcriptionally activating Nfatc1 and BMP signaling, two key quiescence-governing mechanisms. Deletion of Foxc1 in activated (but not quiescent) stem cells causes failure to re-establish quiescence. Conditional Foxc1 deletion (Cre-lox) in hair follicle SCs, lineage tracing, gene expression analysis (Nfatc1, BMP targets), in vivo hair cycle phenotyping Science (New York, N.Y.) High 26912704
2014 SDF1α is a direct transcriptional target of Foxc1 expressed in head mesenchyme. Loss of Foxc1 reduces SDF1α levels, impairing cerebellar radial glial proliferation and Purkinje cell migration; SDF1α rescues the Foxc1-/- cerebellar phenotype. Conditional deletion of SDF1α receptor Cxcr4 in neural progenitors (Nes-Cre) mimics the Foxc1-/- cerebellar phenotype. Foxc1-/- mouse cerebellar phenotyping, in vitro radial glial mitogen and chemoattractant assays with SDF1α, conditional Cxcr4 knockout, SDF1α rescue experiment eLife High 25513817
2016 FOXC1 and FOXC2 are essential regulators of lymphangiogenesis via control of ERK signaling in lymphatic endothelial cells. LEC-specific deletion of Foxc1 and/or Foxc2 causes aberrant Ras regulator expression and ERK hyperactivation; pharmacological ERK inhibition in utero rescues the abnormal lymphatic phenotype. LEC-specific conditional Foxc1/Foxc2 knockout mice, ERK phosphorylation analysis, gene expression profiling, in utero ERK inhibitor treatment rescue The Journal of clinical investigation High 27214551
2014 foxc1b-expressing mesenchymal cells differentiate into acta2-expressing vascular smooth muscle cells in zebrafish head. foxc1 is expressed upstream of acta2 and is required for acta2 expression; loss of foxc1 function blocks vSMC differentiation without affecting pericytes. Timelapse live imaging, CRISPR/morpholino loss-of-function, RNA-seq gene expression profiling, marker co-expression analysis Developmental biology High 31199900
2020 FOXC1 and FOXC2 are induced by laminar (not oscillatory) shear stress in lymphatic endothelial cells. FOXC1 KD increases focal adhesions and actin stress fibers whereas FOXC2 KD disrupts cell junctions, both mediated by increased ROCK activation. ROCK inhibition rescues FOXC1-dependent cytoskeletal changes in vitro and FOXC2-dependent valve degeneration in vivo. Shear stress cell culture experiments, inducible endothelial-specific Foxc1/Foxc2 deletion in mice, siRNA knockdown, ROCK inhibitor rescue, cytoskeletal/junctional marker analysis eLife High 32510325
2013 Foxc1 and Fgf8 genetically interact in jaw patterning; reduction of Fgf8 dosage modifies the syngnathic phenotype of Foxc1 null mice. Loss of Foxc1 causes ectopic osteoblast development in the maxillary prominence leading to bony fusion (syngnathia), establishing Foxc1 as a regulator of neural crest-derived osteogenic patterning in jaw development. Foxc1 null mouse phenotyping, Fgf8 compound genetic interaction, histology, in situ hybridization for osteogenic markers PLoS genetics Medium 24385915
2005 FGF2 regulates Foxc1 expression in calvarial mesenchyme, and Foxc1 integrates BMP and FGF signaling pathways (required for BMP regulation of Msx2) independently of Twist or Noggin during calvarial bone development. Foxc1 null mouse analysis, FGF2 treatment of calvarial cells, expression analysis of Fgf receptors/Twist/Noggin/Msx2, osteogenic differentiation assays Developmental dynamics Medium 15906377
2004 Foxc1 and Foxc2 regulate paraxial versus intermediate mesoderm cell fate specification. Mouse embryos lacking both genes show expansion of intermediate mesoderm markers into the paraxial domain and ectopic mesonephric tubules. Gain-of-function of Foxc1/Foxc2 in chick prospective intermediate mesoderm drives paraxial fate (Pax7, Paraxis expression). Foxc1/Foxc2 double knockout mouse embryo analysis, chick in ovo gain-of-function electroporation, in situ hybridization for mesoderm markers Developmental biology High 15196959
2002 FOXC1 is a TGF-β1-responsive gene; TGF-β1 upregulates FOXC1 transcription. Ectopic FOXC1 expression in HeLa cells (which lack FOXC1) restores TGF-β1-mediated G0/G1 cell cycle arrest, placing FOXC1 as a downstream mediator of TGF-β1 growth-inhibitory signaling. Gene cloning from TGF-β1-treated cells, FOXC1 cDNA transfection rescue assay in HeLa cells, cell cycle analysis (G0/G1 arrest readout) Genomics Medium 12408963
2017 NF-κB is a pivotal mediator of EGF/EGFR-induced FOXC1 transcription; NF-κB-p65 enhances FOXC1 promoter activity and directly binds to the FOXC1 promoter region. Loss or inhibition of NF-κB diminishes FOXC1 expression in basal-like breast cancer cells. Luciferase reporter assays, EMSA, biotinylated oligonucleotide precipitation assay, ChIP at the FOXC1 promoter, NF-κB knockdown/inhibition Cell communication and signaling : CCS Medium 28629477
2018 FOXC1 directly binds the beta-catenin promoter to activate its transcription, as demonstrated by luciferase reporter and ChIP assays. Overexpression of beta-catenin reverses CSC-like property inhibition by FOXC1 knockdown, and beta-catenin knockdown attenuates FOXC1-induced CSC properties in NSCLC. Luciferase reporter assay, ChIP at beta-catenin promoter, FOXC1 KD/OE with beta-catenin rescue/KD epistasis, sphere-formation and tumorigenicity assays Journal of experimental & clinical cancer research : CR Medium 30189871
2018 FOXC1 directly binds the promoter region of the FBP1 gene and negatively regulates its transcriptional activity, reprogramming glycolytic metabolism (Warburg effect) to promote colorectal cancer cell proliferation. ChIP at FBP1 promoter, luciferase reporter assay, FOXC1 OE/KD with metabolic readouts (glucose consumption, lactate production), in vivo xenograft Oncogene Medium 30171256
2019 p38 MAPK phosphorylates FOXC1 at Ser241 and Ser272, maintaining FOXC1 protein stability by inhibiting ubiquitination-mediated degradation. FOXC1 is dephosphorylated by protein phosphatase 2A. Stabilized FOXC1 transcriptionally activates MMP10, SOX4, and SOX13 to promote colorectal cancer metastasis. Site-directed mutagenesis of Ser241/272, co-immunoprecipitation of p38-FOXC1, ubiquitination assays, PP2A dephosphorylation assay, ChIP at MMP10 promoter, in vitro and in vivo metastasis assays The Journal of pathology High 31650548
2021 FOXC1 and RUNX1 physically interact through their Forkhead and Runt domains, respectively, and co-occupy primed/active enhancers near differentiation genes in AML. FOXC1 stabilizes association of RUNX1, HDAC1, and Groucho repressor TLE3 at these enhancers to suppress their activity; FOXC1 knockdown causes loss of repressor proteins, gain of CEBPA binding, enhancer acetylation, and upregulation of differentiation genes including KLF2, while redistributing RUNX1/TLE3/HDAC1 from enhancers to promoters to repress self-renewal genes MYC and MYB. Integrated proteomics (mass spectrometry), ChIP-seq, ATAC-seq, Co-IP of FOXC1-RUNX1 with domain mapping, FOXC1 KD with transcriptomic and epigenomic readouts Cell reports High 34551306
2016 FOXC1 expression is regulated by BMP4-induced SMAD activity; SMAD proteins bind the mouse Foxc1 promoter approximately 300 bp upstream of the transcription start site. In progenitor cells, Foxc1 is required for BMP4-induced osteogenic differentiation; its knockdown in preosteoblasts (MC3T3) accelerates osteoblast differentiation and mineralization. BMP4 treatment, ChIP demonstrating SMAD binding to Foxc1 promoter, luciferase reporter of SMAD-responsive Foxc1 promoter fragment, siRNA knockdown with osteogenic differentiation assays Journal of cellular biochemistry Medium 26666591
2021 FOXC1 governs corneal epithelial fate by directly binding lineage-specific open promoters or enhancers marked by H3K4me2. FOXC1 depletion activates the keratinization pathway and reprograms corneal epithelial cells into skin-like cells, disrupts collagen metabolism and interferon signaling, and causes loss of IRF1 and PAX6 expression. ATAC-seq, H3K4me2 ChIP-seq, FOXC1 KD in corneal epithelial cells and patient samples, transcriptomic analysis Signal transduction and targeted therapy High 33414365
2021 Foxc1 expression in chondrocytes is directly regulated by SOX9 activity. Foxc1 promotes chondrocyte differentiation; chondrocyte-specific deletion of Foxc1 and Foxc2 disrupts growth plate organization, reduces chondrocyte proliferation, and delays hypertrophy, with genome-wide disruption of chondrogenesis/ossification gene expression. SOX9 regulation of Foxc1 promoter, conditional chondrocyte-specific Foxc1/Foxc2 knockout in mice, skeletal phenotyping, differential gene expression (RNA-seq), in vitro chondrogenesis assays in ESCs The Journal of biological chemistry High 34331943
2021 Foxc1 promotes chondrogenesis in the face by establishing chromatin accessibility at cartilage-associated gene enhancers co-enriched for Sox9 and Fox binding motifs. In zebrafish lacking Foxc1 paralogs, there is a global decrease in chondrocyte chromatin accessibility coincident with loss of dorsal facial cartilages; transgenesis confirmed that Foxc1-dependent elements function as stage- and region-specific enhancers. ATAC-seq during zebrafish neural crest-to-chondrocyte differentiation, Foxc1 mutant zebrafish analysis, transgenic enhancer reporter assays eLife High 33501917
2017 FOXC1 loss of function in zebrafish reduces retinal ganglion cell number in the retinal ganglion cell layer and produces a thinner optic nerve. Foxc1 is required for expression of atoh7, a gene necessary for RGC differentiation; markers of differentiated RGCs (pou4f2) are downregulated without increased cell death, indicating a differentiation rather than survival defect. CRISPR-induced foxc1b null mutation, morpholino foxc1a knockdown, cell counting in RGC layer, optic nerve thickness measurement, atoh7 and pou4f2 expression analysis Vision research Medium 30684501
2021 FOXC1 promotes HCC proliferation and metastasis by transcriptionally upregulating DNMT3B, which induces DNA hypermethylation of the CTH (cystathionine γ-lyase) promoter to silence CTH, thereby inhibiting cysteine metabolism and increasing ROS levels. ROS in turn activates the ERK1/2-ELK1 pathway to further upregulate FOXC1, forming a positive feedback loop. Amino acid arrays, ROS measurement assays, bisulfite genomic sequencing, methylation-specific PCR, ChIP (FOXC1 on DNMT3B promoter), luciferase reporter, CTH overexpression/knockdown rescue, in vivo xenograft metastasis Journal of experimental & clinical cancer research : CR High 33522955
2023 FOXC1 and FOXC2 directly bind regulatory elements of CXCL12 and RSPO3 loci in blood endothelial cells and lymphatic endothelial cells, respectively. EC/LEC-specific deletion of Foxc1 and/or Foxc2 impairs vascular regrowth after intestinal ischemia-reperfusion and reduces CXCL12 (in BECs) and Wnt activator RSPO3 (in LECs), thereby suppressing Wnt/ISC activation; CXCL12 and RSPO3 treatment rescues intestinal damage in respective mutants. EC/LEC-specific conditional Foxc1/Foxc2 knockout mice, intestinal I/R injury model, ChIP at CXCL12 and RSPO3 loci, cytokine/growth factor rescue experiments, Wnt signaling readouts in ISCs EMBO reports High 37154714
2024 FOXC1 in vascular endothelial cells transcriptionally regulates Slc3a2 and Slc7a5 (encoding the CD98/LAT1/4F2hc amino acid transporter), which controls intracellular amino acid transport and mTOR activation. EC-specific Foxc1 loss reduces mTOR activity and impairs retinal vascular growth; mTOR agonist MHY-1485 rescues perturbed angiogenesis. Foxc1 is also required for pericyte function in blood-retina barrier formation. EC-specific conditional Foxc1 KO mice, retinal flat-mount vascular analysis, Slc3a2/Slc7a5 expression analysis, mTOR activity assay, mTOR agonist rescue, oxygen-induced retinopathy model, pericyte marker analysis Nature communications High 38755144
2015 Hypo- and hypermorphic FOXC1 mutations cause dominant glaucoma with remarkable phenotypic variability. Truncating mutations retaining the N-terminal activation domain (p.Y47X, p.Q106X) exhibit hyperactivity (gain-of-function); p.I126S causes partial loss-of-function through impaired protein stability, DNA binding, phosphorylation, and altered subcellular distribution. Transcriptional activity assays, protein stability assays, DNA-binding assays, subcellular localization analysis, phosphorylation analysis of patient-derived mutations PloS one Medium 25786029
2019 FOXC1 transcriptionally activates TLR3 and TLR4 promoters by direct binding (shown by ChIP), with FOXC1 overexpression/knockdown bidirectionally regulating TLR3/4 mRNA and protein levels. FOXC1 overexpression has pro-inflammatory effects and worsens infarct size and heart function in a mouse myocardial ischemia model. In silico TF screening, dual-luciferase reporter for TLR3/4 promoters, ChIP of FOXC1 at TLR3/4 promoters, FOXC1 OE/KD, in vivo mouse MI model with functional cardiac readouts Journal of cellular and molecular medicine Medium 31517441
2021 FOXC1 negatively regulates DKK1 expression by binding to its promoter, thereby de-repressing the Wnt signaling pathway. FOXC1 can also form a complex with unphosphorylated β-catenin in the cytoplasm, facilitating β-catenin nuclear translocation to regulate c-MYC expression and promote gastric cancer cell proliferation. ChIP at DKK1 promoter, Co-IP of FOXC1-β-catenin complex, luciferase reporter, FOXC1 KD/OE with Wnt pathway readouts and proliferation assays Frontiers in cell and developmental biology Medium 33987183
2023 Foxc1 functions as a transcriptional activator of Ubqln1 (ubiquilin-1); Sirt1 promotes Foxc1 expression by deacetylating EZH2 and inhibiting its activity toward the Foxc1 locus. This Sirt1/Foxc1/Ubqln1 axis regulates proteostasis during cerebral ischemia-reperfusion injury. Co-IP, ChIP, dual-luciferase reporter assays, siRNA knockdown of Foxc1/Sirt1/Ubqln1, OGD/R and MCAO/R ischemia models International immunopharmacology Medium 38452414
2023 FOXC1 overexpression inhibits microglial migration and neuroinflammation in sepsis-associated encephalopathy by increasing IκBα expression, thereby suppressing NF-κB (p65) activation and downstream pro-inflammatory cytokines (IL-1β, TNF-α); IκBα inhibition reverses the Foxc1-mediated effects in vitro. FOXC1 OE/KD in BV-2 microglia (LPS model), CLP mouse surgery model, Morris Water Maze cognitive assessment, Western blot (IκBα, p65, cytokines), IκBα inhibitor rescue Molecular medicine reports Medium 35103290
2017 FOXC1 directly binds an upstream regulatory region of Fgfr1 and activates an Fgfr1 promoter element to specifically induce expression of the FGFR1-IIIc isoform following TGFβ1-induced EMT, promoting three-dimensional invasive migration that can be rescued by FGFR1 re-expression in FOXC1-knockdown cells. RNA-seq (isoform analysis), ChIP at Fgfr1 upstream region, Fgfr1 promoter luciferase reporter, FOXC1 KD/OE, FGFR1 rescue experiment, 3D migration assay Molecular cancer research : MCR Medium 28684636
2020 FOXC1 mediates LINC00301 expression in NSCLC by binding to the LINC00301 promoter (validated by ChIP). LINC00301 in turn binds EZH2 (at EZH2 aa 612-727, LINC00301 nt 83-123), causing H3K27me3 at the EAF2 promoter to silence EAF2, which stabilizes pVHL and thereby controls HIF1α levels. ChIP (FOXC1 on LINC00301 promoter), RNA pull-down, protein domain mapping, RIP, EMSA, Alpha protein-protein interaction assay, luciferase reporter Genome medicine Medium 32878637
2024 Telomere shortening in cardiomyocytes leads to proximal telomeric chromatin decompaction (reduced TAD insulation), increasing FOXC1 accessibility and expression. Overexpression of FOXC1 in hiPSC-CMs induces aging markers, mitochondrial dysfunction, and contractile dysfunction; FOXC1 knockdown rescues these phenotypes. FOXC1 protein levels are negatively correlated with telomere lengths in heart failure patient cardiomyocytes. Hi-C (TAD analysis), RNA-seq, isogenic hiPSC-CM model with short vs normal telomeres, FOXC1 OE/KD with mitochondrial function and contractility readouts, patient sample FOXC1/telomere correlation Nucleic acids research Medium 38634789
2016 Foxc1 regulates early cardiomyogenesis during a specific differentiation window (D4-D6) in ESC-derived cardiomyocytes, controlling early cardiomyogenic factors Isl1, Mef2c, and Nkx2.5. A novel direct Foxc1 target, Myh7 (MHCβ), was identified, establishing Foxc1 as a regulator of both cardiomyogenic specification and functional cardiomyocyte properties. Foxc1 KD/OE in ESC differentiation model, whole-transcriptome RNA-seq, electrophysiological characterization of ESC-derived cardiomyocytes, Myh7 as direct target (RNA-seq + expression analysis) Stem cells (Dayton, Ohio) Medium 26824887
2025 Serine deprivation activates ERK1/2-ELK1 signaling to upregulate FOXC1 in colorectal cancer cells. Elevated FOXC1 promotes transcription of serine synthesis pathway enzymes PHGDH, PSAT1, and PSPH, supporting serine biosynthesis and 5-FU resistance through effects on purine metabolism and DNA damage repair. ChIP (FOXC1 at PHGDH/PSAT1/PSPH promoters), luciferase reporter, serine deprivation cell culture model, ERK1/2-ELK1 pathway inhibitor studies, 5-FU resistance assays Cell communication and signaling : CCS Medium 39773485
2020 EZH2 represses FOXC1 expression by mediating H3K27me3 at the FOXC1 promoter in breast cancer cells. OGT (O-GlcNAc transferase) knockdown alters EZH2 binding to the FOXC1 promoter in a cell-context-dependent manner and is associated with increased histone H3 acetylation at the FOXC1 locus, modulating FOXC1 expression. ChIP (EZH2 and OGT at FOXC1 promoter), EZH2 inhibitor treatment (DZNeP, GSK343), OGT siRNA knockdown, H3K27me3 and H3 acetylation analysis Acta pharmacologica Sinica Medium 33057161
2023 Icaritin inhibits endometrial cancer cell proliferation by suppressing O-GlcNAc modification (O-GlcNAcylation) of FOXC1, thereby reducing FOXC1 protein stability and activity. Western blot for O-GlcNAcylated FOXC1, icaritin treatment, FOXC1 KD/OE in cell proliferation/invasion assays, xenograft tumor model Phytomedicine Low 37683586

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1998 The forkhead transcription factor gene FKHL7 is responsible for glaucoma phenotypes which map to 6p25. Nature genetics 369 9620769
1998 Mutations of the forkhead/winged-helix gene, FKHL7, in patients with Axenfeld-Rieger anomaly. American journal of human genetics 272 9792859
2014 Foxc1 is a critical regulator of haematopoietic stem/progenitor cell niche formation. Nature 209 24590069
2017 FOXC1: an emerging marker and therapeutic target for cancer. Oncogene 137 28288141
2015 FOXC1 Activates Smoothened-Independent Hedgehog Signaling in Basal-like Breast Cancer. Cell reports 128 26565916
2020 FOXC1-mediated LINC00301 facilitates tumor progression and triggers an immune-suppressing microenvironment in non-small cell lung cancer by regulating the HIF1α pathway. Genome medicine 125 32878637
2016 Foxc1 reinforces quiescence in self-renewing hair follicle stem cells. Science (New York, N.Y.) 118 26912704
2006 Functional interactions between FOXC1 and PITX2 underlie the sensitivity to FOXC1 gene dose in Axenfeld-Rieger syndrome and anterior segment dysgenesis. Human molecular genetics 118 16449236
2012 PITX2 and FOXC1 spectrum of mutations in ocular syndromes. European journal of human genetics : EJHG 117 22569110
2014 Mutation of FOXC1 and PITX2 induces cerebral small-vessel disease. The Journal of clinical investigation 109 25250569
2000 Axenfeld-Rieger syndrome resulting from mutation of the FKHL7 gene on chromosome 6p25. European journal of human genetics : EJHG 96 10713890
2002 FOXC1 transcriptional regulation is mediated by N- and C-terminal activation domains and contains a phosphorylated transcriptional inhibitory domain. The Journal of biological chemistry 91 11782474
2007 FOXC1 is required for cell viability and resistance to oxidative stress in the eye through the transcriptional regulation of FOXO1A. Human molecular genetics 90 17993506
2004 The forkhead genes, Foxc1 and Foxc2, regulate paraxial versus intermediate mesoderm cell fate. Developmental biology 87 15196959
2018 FOXC1 induces cancer stem cell-like properties through upregulation of beta-catenin in NSCLC. Journal of experimental & clinical cancer research : CR 76 30189871
2011 BRCA1 and GATA3 corepress FOXC1 to inhibit the pathogenesis of basal-like breast cancers. Oncogene 74 22120723
2015 MicroRNA-138-5p regulates pancreatic cancer cell growth through targeting FOXC1. Cellular oncology (Dordrecht, Netherlands) 73 25875420
2017 FOXC1 in cancer development and therapy: deciphering its emerging and divergent roles. Therapeutic advances in medical oncology 71 29449899
2017 FOXC1-induced non-canonical WNT5A-MMP7 signaling regulates invasiveness in triple-negative breast cancer. Oncogene 67 29249801
2016 Foxc1 and Foxc2 deletion causes abnormal lymphangiogenesis and correlates with ERK hyperactivation. The Journal of clinical investigation 66 27214551
1995 Chromosomal localization of six human forkhead genes, freac-1 (FKHL5), -3 (FKHL7), -4 (FKHL8), -5 (FKHL9), -6 (FKHL10), and -8 (FKHL12). Genomics 65 8825632
2015 The transcription factor Foxc1 is necessary for Ihh-Gli2-regulated endochondral ossification. Nature communications 64 25808752
2014 The expression and functional role of a FOXC1 related mRNA-lncRNA pair in oral squamous cell carcinoma. Molecular and cellular biochemistry 64 24889262
2019 Inhibition of lung cancer growth and metastasis by DHA and its metabolite, RvD1, through miR-138-5p/FOXC1 pathway. Journal of experimental & clinical cancer research : CR 61 31783879
2021 FOXC1 promotes HCC proliferation and metastasis by Upregulating DNMT3B to induce DNA Hypermethylation of CTH promoter. Journal of experimental & clinical cancer research : CR 59 33522955
2017 FOXC1 promotes proliferation and epithelial-mesenchymal transition in cervical carcinoma through the PI3K-AKT signal pathway. American journal of translational research 55 28386355
2017 FOXC1, the new player in the cancer sandbox. Oncotarget 55 29487724
2009 The cooperative roles of Foxc1 and Foxc2 in cardiovascular development. Advances in experimental medicine and biology 55 20429416
2020 Shear stimulation of FOXC1 and FOXC2 differentially regulates cytoskeletal activity during lymphatic valve maturation. eLife 51 32510325
2018 The FOXC1/FBP1 signaling axis promotes colorectal cancer proliferation by enhancing the Warburg effect. Oncogene 51 30171256
2015 Diagnosis of Basal-Like Breast Cancer Using a FOXC1-Based Assay. Journal of the National Cancer Institute 51 26041837
2013 Interaction between Foxc1 and Fgf8 during mammalian jaw patterning and in the pathogenesis of syngnathia. PLoS genetics 49 24385915
1999 An interstitial deletion of 6p24-p25 proximal to the FKHL7 locus and including AP-2alpha that affects anterior eye chamber development. Journal of medical genetics 48 10507730
2015 MicroRNA-495 downregulates FOXC1 expression to suppress cell growth and migration in endometrial cancer. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 47 26198045
2002 Identification of FOXC1 as a TGF-beta1 responsive gene and its involvement in negative regulation of cell growth. Genomics 47 12408963
2006 Novel mutations of FOXC1 and PITX2 in patients with Axenfeld-Rieger malformations. Investigative ophthalmology & visual science 46 16936096
2019 foxc1 is required for embryonic head vascular smooth muscle differentiation in zebrafish. Developmental biology 44 31199900
2017 Foxc1 and Foxc2 in the Neural Crest Are Required for Ocular Anterior Segment Development. Investigative ophthalmology & visual science 42 28253399
2003 Mutation spectrum of FOXC1 and clinical genetic heterogeneity of Axenfeld-Rieger anomaly in India. Molecular vision 42 12592227
2014 Foxc1 dependent mesenchymal signalling drives embryonic cerebellar growth. eLife 41 25513817
2006 FGF19 is a target for FOXC1 regulation in ciliary body-derived cells. Human molecular genetics 39 17000708
2017 MicroRNA-374c-5p regulates the invasion and migration of cervical cancer by acting on the Foxc1/snail pathway. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 38 28810526
2016 FOXC1 promotes melanoma by activating MST1R/PI3K/AKT. Oncotarget 38 27533251
2019 Prevalence of FOXC1 Variants in Individuals With a Suspected Diagnosis of Primary Congenital Glaucoma. JAMA ophthalmology 35 30653210
2017 The long noncoding RNA FOXCUT promotes proliferation and migration by targeting FOXC1 in nasopharyngeal carcinoma. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 35 28635400
2016 miR-133 inhibits pituitary tumor cell migration and invasion via down-regulating FOXC1 expression. Genetics and molecular research : GMR 35 27050992
2012 Generation of conditional alleles for Foxc1 and Foxc2 in mice. Genesis (New York, N.Y. : 2000) 35 22522965
2011 IHG-1 promotes mitochondrial biogenesis by stabilizing PGC-1α. Journal of the American Society of Nephrology : JASN 35 21784897
2017 Phenotypic outcomes in Mouse and Human Foxc1 dependent Dandy-Walker cerebellar malformation suggest shared mechanisms. eLife 33 28092268
2017 Identification of EGF-NF-κB-FOXC1 signaling axis in basal-like breast cancer. Cell communication and signaling : CCS 33 28629477
2016 Foxc1 Regulates Early Cardiomyogenesis and Functional Properties of Embryonic Stem Cell Derived Cardiomyocytes. Stem cells (Dayton, Ohio) 33 26824887
2005 Foxc1 integrates Fgf and Bmp signalling independently of twist or noggin during calvarial bone development. Developmental dynamics : an official publication of the American Association of Anatomists 31 15906377
2016 Foxc1 Expression in Early Osteogenic Differentiation Is Regulated by BMP4-SMAD Activity. Journal of cellular biochemistry 30 26666591
2003 Identification and analysis of a novel mutation in the FOXC1 forkhead domain. Investigative ophthalmology & visual science 30 14578375
2021 Loss of FOXC1 contributes to the corneal epithelial fate switch and pathogenesis. Signal transduction and targeted therapy 29 33414365
2015 Hypo- and hypermorphic FOXC1 mutations in dominant glaucoma: transactivation and phenotypic variability. PloS one 29 25786029
2019 p38-regulated FOXC1 stability is required for colorectal cancer metastasis. The Journal of pathology 28 31650548
2018 Impact of OGT deregulation on EZH2 target genes FOXA1 and FOXC1 expression in breast cancer cells. PloS one 28 29864144
2022 Overexpression of Foxc1 ameliorates sepsis‑associated encephalopathy by inhibiting microglial migration and neuroinflammation through the IκBα/NF‑κB pathway. Molecular medicine reports 27 35103290
2021 Foxc1 establishes enhancer accessibility for craniofacial cartilage differentiation. eLife 27 33501917
2021 Enhancer recruitment of transcription repressors RUNX1 and TLE3 by mis-expressed FOXC1 blocks differentiation in acute myeloid leukemia. Cell reports 27 34551306
2008 Human p32 is a novel FOXC1-interacting protein that regulates FOXC1 transcriptional activity in ocular cells. Investigative ophthalmology & visual science 26 18676636
2023 Endothelial FOXC1 and FOXC2 promote intestinal regeneration after ischemia-reperfusion injury. EMBO reports 25 37154714
2020 MIR-138-5P inhibits the progression of prostate cancer by targeting FOXC1. Molecular genetics & genomic medicine 25 32107877
2004 FOXC1 gene deletion is associated with eye anomalies in ring chromosome 6. American journal of medical genetics. Part A 25 14708101
2002 Novel mutation in FOXC1 wing region causing Axenfeld-Rieger anomaly. Investigative ophthalmology & visual science 25 12454026
2017 FOXC1 Regulates FGFR1 Isoform Switching to Promote Invasion Following TGFβ-Induced EMT. Molecular cancer research : MCR 24 28684636
2021 Therapeutically Targeting Cancers That Overexpress FOXC1: A Transcriptional Driver of Cell Plasticity, Partial EMT, and Cancer Metastasis. Frontiers in oncology 23 34540690
2016 FOXC1 is associated with estrogen receptor alpha and affects sensitivity of tamoxifen treatment in breast cancer. Cancer medicine 23 28028927
2001 A novel (Pro79Thr) mutation in the FKHL7 gene in a Japanese family with Axenfeld-Rieger syndrome. American journal of ophthalmology 23 11589884
2021 FOXC1 Negatively Regulates DKK1 Expression to Promote Gastric Cancer Cell Proliferation Through Activation of Wnt Signaling Pathway. Frontiers in cell and developmental biology 22 33987183
2020 The Sp1/FOXC1/HOTTIP/LATS2/YAP/β-catenin cascade promotes malignant and metastatic progression of osteosarcoma. Molecular oncology 21 32634265
2014 IHG-1 increases mitochondrial fusion and bioenergetic function. Diabetes 21 25008184
2021 MicroRNA-495/TGF-β/FOXC1 axis regulates multidrug resistance in metaplastic breast cancer cells. Biochemical pharmacology 20 34298004
2019 Loss of foxc1 in zebrafish reduces optic nerve size and cell number in the retinal ganglion cell layer. Vision research 20 30684501
2017 Foxc1 and Foxc2 are necessary to maintain glomerular podocytes. Experimental cell research 20 28223138
2015 Rare FOXC1 variants in congenital glaucoma: identification of translation regulatory sequences. European journal of human genetics : EJHG 20 26220699
2012 FoxC1-dependent regulation of vascular endothelial growth factor signaling in corneal avascularity. Trends in cardiovascular medicine 20 22939989
2019 FOXC1 up-regulates the expression of toll-like receptors in myocardial ischaemia. Journal of cellular and molecular medicine 19 31517441
2014 Signals from the lens and Foxc1 regulate the expression of key genes during the onset of corneal endothelial development. Experimental cell research 19 24472616
2010 MYOC and FOXC1 gene analysis in primary congenital glaucoma. Molecular vision 19 21031026
2004 Axenfeld-Rieger anomaly: a novel mutation in the forkhead box C1 (FOXC1) gene in a 4-generation family. Archives of ophthalmology (Chicago, Ill. : 1960) 19 15477465
2024 Fisetin alleviates cerebral ischemia/reperfusion injury by regulating Sirt1/Foxc1/Ubqln1 pathway-mediated proteostasis. International immunopharmacology 18 38452414
2020 EZH2 regulates expression of FOXC1 by mediating H3K27me3 in breast cancers. Acta pharmacologica Sinica 18 33057161
2025 FOXC1-mediated serine metabolism reprogramming enhances colorectal cancer growth and 5-FU resistance under serine restriction. Cell communication and signaling : CCS 17 39773485
2024 FOXC1 regulates endothelial CD98 (LAT1/4F2hc) expression in retinal angiogenesis and blood-retina barrier formation. Nature communications 17 38755144
2022 FOXC1. Archives of pathology & laboratory medicine 17 34784418
2022 FOXC1‑mediated TRIM22 regulates the excessive proliferation and inflammation of fibroblast‑like synoviocytes in rheumatoid arthritis via NF‑κB signaling pathway. Molecular medicine reports 17 35946462
2020 Activation of the LINC00242/miR-141/FOXC1 axis underpins the development of gastric cancer. Cancer cell international 17 32587479
2019 Satb2 expression in Foxc1-promoted osteogenic differentiation of MC3T3-E1 cells is negatively regulated by microRNA-103-3p. Acta biochimica et biophysica Sinica 17 31089719
2024 Proximal telomeric decompaction due to telomere shortening drives FOXC1-dependent myocardial senescence. Nucleic acids research 16 38634789
2021 Long Non-Coding RNA LINC01929 Accelerates Progression of Oral Squamous Cell Carcinoma by Targeting the miR-137-3p/FOXC1 Axis. Frontiers in oncology 16 33968763
2021 Mechanistic Insights into Axenfeld-Rieger Syndrome from Zebrafish foxc1 and pitx2 Mutants. International journal of molecular sciences 16 34576164
2020 Disruption of foxc1 genes in zebrafish results in dosage-dependent phenotypes overlapping Axenfeld-Rieger syndrome. Human molecular genetics 16 32720677
2020 Gene-specific facial dysmorphism in Axenfeld-Rieger syndrome caused by FOXC1 and PITX2 variants. American journal of medical genetics. Part A 16 33231930
2016 Characterization of Kidney and Skeleton Phenotypes of Mice Double Heterozygous for Foxc1 and Foxc2. Cells, tissues, organs 16 27193493
2024 Remdesivir ameliorates ulcerative colitis-propelled cell inflammation and pyroptosis in acetic acid rats by restoring SIRT6/FoxC1 pathway. International immunopharmacology 15 38878489
2024 HIF1A-AS2 promotes the metabolic reprogramming and progression of colorectal cancer via miR-141-3p/FOXC1 axis. Cell death & disease 15 39227375
2023 Icaritin inhibits endometrial carcinoma cells by suppressing O-GlcNAcylation of FOXC1. Phytomedicine : international journal of phytotherapy and phytopharmacology 14 37683586
2021 Loss of Foxc1 and Foxc2 function in chondroprogenitor cells disrupts endochondral ossification. The Journal of biological chemistry 14 34331943

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