Affinage

FOXP4

Forkhead box protein P4 · UniProt Q8IVH2

Length
680 aa
Mass
73.5 kDa
Annotated
2026-06-09
77 papers in source corpus 27 papers cited in narrative 26 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

FOXP4 is a forkhead/winged-helix transcription factor of the FOXP subfamily that acts as a context-dependent activator or repressor to control epithelial, neuronal, immune, metabolic, and stem-cell programs during development (PMID:14516685, PMID:12617805, PMID:25027557). It assembles into homo- and heterodimers with FOXP1 and FOXP2, and distinct dimer combinations differentially tune shared neuronal target genes, establishing combinatorial FOXP coding as a core regulatory logic (PMID:25027557). This combinatorial principle recurs across tissues through genetic redundancy: only compound loss of FOXP1/2/4 in pancreatic islet endocrine cells reduces alpha-cell mass and glucagon secretion via cell-cycle mediators (PMID:26021489), compound FOXP1/4 loss in hair follicles destroys the stem-cell niche (PMID:35759955), and combined FOXP1/4 deletion in Tregs causes autoimmunity, with both factors binding the Il2ra (CD25) promoter (PMID:40794436). FOXP4 directly binds target promoters through its forkhead DNA-binding domain to drive distinct programs: it activates the thermogenic gene UCP1 in adipocytes downstream of HSF1 while acting as a repressor of the beige thermogenic program in differentiated cells (PMID:34384787, PMID:35297993), and it controls cortical neuron migration by maintaining N-cadherin-based adherens junctions in radial glia (PMID:36646976). In cancer it functions as a pro-malignant transcription factor, directly regulating EMT effectors Slug and Snail, Wnt-pathway components LEF-1 and PTK7, and additional targets including GPX4 (suppressing ferroptosis), SOX12, FBXW7, NDST2, CYP26B1, and MYC (PMID:30930991, PMID:31040716, PMID:38740744, PMID:40789053). Heterozygous missense variants in the FOXP4 forkhead domain abolish transcriptional repressor activity and can mislocalize the protein, causing an autosomal dominant neurodevelopmental disorder with speech/language delay and congenital abnormalities (PMID:33110267).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2002 Medium

    Establishing FOXP4 as a forkhead transcription factor and mapping its developmental expression defined where in the embryo it might act.

    Evidence cDNA characterization, IHC with specific antisera, and in situ hybridization across embryonic tissues

    PMID:12617805 PMID:14516685

    Open questions at the time
    • No direct target genes identified at this stage
    • No functional loss-of-function data
  2. 2010 Medium

    Showed FOXP4 has a tissue-specific maintenance role, being required to sustain Purkinje cell dendritic arborization rather than to initiate it.

    Evidence siRNA knockdown in organotypic cerebellar slice culture with dendritic morphology analysis

    PMID:20951773

    Open questions at the time
    • Direct transcriptional targets in Purkinje cells not identified
    • Mechanism linking FOXP4 to Bergmann glial alignment unresolved
  3. 2012 Medium

    Defined a non-essential but modulatory immune role, with FOXP4 dispensable for T cell development but required for full effector cytokine recall responses.

    Evidence CD4Cre conditional knockout with in vivo T. gondii and LCMV challenge and cytokine measurement

    PMID:22912696

    Open questions at the time
    • Target genes controlling cytokine recall not identified
    • Possible redundancy with other FOXPs not tested here
  4. 2014 Medium

    Established the combinatorial logic of FOXP regulation by showing FOXP4 dimerizes with FOXP1/2 and that specific dimers differentially regulate shared targets.

    Evidence Stable transfection of FOXP1/2/4 ORFs in HEK293 cells with qRT-PCR of target genes

    PMID:25027557

    Open questions at the time
    • Overexpression system may not reflect endogenous stoichiometry
    • Direct DNA binding by FOXP4 not shown in this study
  5. 2015 High

    Demonstrated functional redundancy among FOXP1/2/4 in vivo, where only compound deletion impairs alpha-cell proliferation and glucagon secretion through cell-cycle genes.

    Evidence Triple conditional knockout (Pax6-Cre) mice with physiological, hormone-secretion, and islet gene-expression readouts

    PMID:26021489

    Open questions at the time
    • Whether FOXP4 binds the cell-cycle gene promoters directly not shown
    • Individual FOXP4 contribution masked by redundancy
  6. 2019 Medium

    Identified FOXP4 as a direct driver of EMT in carcinoma by binding and activating Slug and Snail promoters.

    Evidence ChIP/qChIP and luciferase reporter assays with gain/loss-of-function in HCC and breast cancer cells

    PMID:30930991 PMID:31040716

    Open questions at the time
    • Upstream signals controlling FOXP4 in these tumors not defined
    • Single-lab evidence per cancer type
  7. 2019 Medium

    Extended FOXP combinatorial roles to behavior, showing FoxP4 contributes non-redundantly but partly overlapping with FoxP1/2 to vocal learning.

    Evidence Lentiviral knockdown in zebra finch Area X with quantified song analysis

    PMID:31641053

    Open questions at the time
    • Molecular targets underlying song deficits unknown
    • Mammalian relevance not directly tested
  8. 2020 High

    Established the disease mechanism: forkhead-domain missense variants cause loss of repressor activity and sometimes mislocalization, defining an autosomal dominant neurodevelopmental disorder.

    Evidence Luciferase repressor and localization assays across patient variants with clinical phenotyping of 8 individuals

    PMID:33110267

    Open questions at the time
    • In vivo neurodevelopmental targets driving the phenotype not identified
    • Genotype-phenotype correlation across the variant spectrum incomplete
  9. 2021 Medium

    Placed FOXP4 in thermogenic control, acting downstream of HSF1 to directly activate UCP1 in adipocytes.

    Evidence ChIP and luciferase assays, gain/loss-of-function in adipocytes, oxygen consumption, and in vivo thermogenic stimulation

    PMID:34384787

    Open questions at the time
    • Reconciling activator role here with later repressor role in differentiated beige cells
    • Full thermogenic target set not mapped
  10. 2022 Medium

    Resolved a stage-specific dual role in adipocytes: FOXP4 is required for early beige differentiation but represses the thermogenic program in differentiated beige cells.

    Evidence Beige adipocyte-specific conditional knockouts at progenitor and differentiated stages with cold exposure and gene expression

    PMID:35297993

    Open questions at the time
    • Mechanism switching FOXP4 between activator and repressor states unknown
    • Beige-versus-brown specificity determinants undefined
  11. 2022 Medium

    Showed FOXP4 partners with FOXP1 in a complex to maintain the hair follicle stem cell niche, with redundancy revealed only by double knockout.

    Evidence Reciprocal co-immunoprecipitation and single/double conditional knockout mice with niche and apoptosis analysis

    PMID:35759955

    Open questions at the time
    • Direct promoter targets (Fgf18, Bmp6) binding not confirmed
    • Whether FOXP4 acts as activator or repressor at these loci unclear
  12. 2023 Medium

    Defined a cell-biological mechanism in cortical development, placing FOXP4 upstream of N-cadherin adherens junctions required for radial migration.

    Evidence In utero shRNA/dominant-negative knockdown with N-cadherin overexpression rescue in mouse neocortex

    PMID:36646976

    Open questions at the time
    • Whether FOXP4 directly transcribes N-cadherin not shown
    • Link to human neurodevelopmental disorder phenotype not directly tested
  13. 2023 Medium

    Extended FOXP4's pro-proliferative role to human spermatogonial stem cells, marking and sustaining a stem subset.

    Evidence Conditional inactivation in human SSC lines with scRNA-seq, proliferation, and apoptosis assays

    PMID:36018067

    Open questions at the time
    • Direct transcriptional targets in SSCs not identified
    • In vivo human relevance limited to cell lines
  14. 2024 Medium

    Expanded the FOXP4 oncogenic target repertoire across tumor types, linking it directly to Wnt signaling, ferroptosis resistance, stemness, and growth/EMT regulators.

    Evidence ChIP/CUT&Tag, RNA-seq, luciferase, and rescue experiments identifying PTK7, GPX4, SOX12, FBXW7, NDST2, CYP26B1, and MYC across ovarian, colorectal, gastric, thyroid, hepatocellular, and esophageal cancers

    PMID:34590150 PMID:38293397 PMID:38740744 PMID:39047223 PMID:39429915 PMID:40228145 PMID:40789053

    Open questions at the time
    • Determinants of activator-versus-repressor choice at different promoters unknown
    • Each target largely supported by single-lab evidence
  15. 2025 High

    Demonstrated that FOXP1 and FOXP4 redundantly enforce Treg suppressive identity, jointly binding the Il2ra (CD25) promoter to prevent autoimmunity.

    Evidence Foxp3-Cre single and double conditional knockouts with immune phenotyping, cytokine assays, and Il2ra promoter-binding assays

    PMID:40794436

    Open questions at the time
    • FOXP4-specific (non-redundant) Treg targets not separated
    • Direct contribution of FOXP4 binding at CD25 versus FOXP1 not isolated

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved what molecular switch determines whether FOXP4 acts as a transcriptional activator or repressor at a given promoter, and how its tissue-specific cofactor partnerships dictate context-dependent outcomes.
  • No structural model of FOXP4 on DNA or in FOXP dimers
  • Cofactors converting FOXP4 between activation and repression unidentified
  • Genome-wide endogenous FOXP4 binding landscape across tissues not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 9 GO:0003677 DNA binding 7
Localization
GO:0005634 nucleus 2
Pathway
R-HSA-74160 Gene expression (Transcription) 7 R-HSA-1266738 Developmental Biology 4 R-HSA-162582 Signal Transduction 3 R-HSA-1643685 Disease 2 R-HSA-168256 Immune System 2
Partners
FOXP1FOXP2
Complex memberships
FOXP1/FOXP4 complexFOXP4/FOXP1/FOXP2 dimers

Evidence

Reading pass · 26 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 FOXP4 is a member of the Foxp subfamily of winged-helix (forkhead) transcription factors, encoding a 685-amino-acid protein similar to FOXP1 and FOXP2, expressed primarily in pulmonary epithelium (proximal and distal airway), neural tissues, and gut epithelial cells during embryonic development, as determined by immunohistochemical analysis with specific polyclonal antisera. Immunohistochemistry with specific polyclonal antisera; in situ hybridization; cDNA characterization Mechanisms of development / Gene expression patterns Medium 12617805 14516685
2014 FOXP4 forms homo- and heterodimers with FOXP1 and FOXP2, and specific combinations of FOXP1/2/4 dimers differentially regulate transcription of known FOXP2 target genes (including CER1, SFRP4, WISP2, PRICKLE1, NCOR2, SNW1, NEUROD2, PAX3, EFNB3, and SLIT1) involved in early neuronal development, as shown by stable transfection of open-reading frames into HEK293 cells and quantitative real-time PCR. Stable transfection of FOXP1/2/4 ORFs in HEK293 cells; qRT-PCR of target gene expression Journal of molecular neuroscience Medium 25027557
2010 Foxp4 is expressed specifically in migrating and mature Purkinje cells of the cerebellum and is required for the maintenance (but not initiation) of Purkinje cell dendritic arborization; siRNA-mediated knockdown at postnatal day 10 in organotypic cerebellar slice cultures caused highly impaired dendritic arbor organization and loss of associated Bergmann glial fiber radial alignment, without inducing apoptosis. siRNA knockdown in organotypic cerebellar slice culture; morphological analysis of dendritic arbors Neuroscience Medium 20951773
2012 Foxp4 is expressed in thymocytes and peripheral CD4+ and CD8+ T cells, increases following T cell activation, and is dispensable for T cell development, Foxp3+ Treg generation, and in vitro proliferative/effector responses; however, Foxp4-deficient CD4 T cells show reduced effector cytokine production during antigen-specific recall responses in vivo (Toxoplasma gondii and LCMV infection models), as assessed by CD4Cre-mediated conditional knockout. CD4Cre conditional knockout; flow cytometry; in vivo infection challenge with T. gondii and LCMV; cytokine measurement PloS one Medium 22912696
2015 Combined conditional knockout of FOXP1, FOXP2, and FOXP4 (but not individual knockouts) in pancreatic islet endocrine cells (using Pax6-Cre) causes hypoglycemia and hypoglucagonemia due to reduced alpha cell mass and impaired glucagon secretion; this is mediated through effects on cell cycle mediators (decreased Ccna2, Ccnb1, Ccnd2; increased Cdkn1a), while beta cell insulin secretion remains intact. Triple conditional knockout (Pax6-Cre) mice; glucose tolerance testing; radioimmunoassay; immunohistochemistry; gene expression in isolated islets; glucose-stimulated hormone secretion assays Diabetologia High 26021489
2019 FOXP4 transcriptionally regulates Slug (SNAI2), an EMT-associated transcription factor, to promote EMT in hepatocellular carcinoma cells; this was established by ChIP, qChIP, and luciferase reporter assays showing FOXP4 binding to the Slug promoter, and gain/loss-of-function experiments demonstrating FOXP4 controls migration and invasion through this mechanism. ChIP and qChIP assays; luciferase reporter assay; gain- and loss-of-function experiments in HCC cell lines Oncology letters Medium 30930991
2019 FOXP4 transcriptionally activates Snail (SNAI1) in breast cancer cells by binding to the Snail promoter (confirmed by ChIP, qChIP, and dual luciferase reporter assays), promoting EMT; overexpression of Snail partially rescued the inhibitory effect of FOXP4 knockdown on migration and invasion. ChIP assay; dual luciferase reporter assay; gain- and loss-of-function experiments Cancer management and research Medium 31040716
2019 Lentiviral knockdown of FoxP4 in Area X (striatal song nucleus) of juvenile male zebra finches impaired vocal production learning, with song deficits partly overlapping but distinct from those caused by FoxP1 or FoxP2 knockdown, indicating non-redundant but partially shared roles of FoxP1/2/4 in Area X-dependent vocal learning. Lentivirus-mediated knockdown in Area X of juvenile zebra finches; song analysis (spectral and temporal features); comparison across FoxP1/2/4 knockdown groups The Journal of neuroscience Medium 31641053
2020 Heterozygous missense variants in the FOXP4 forkhead box DNA-binding domain cause loss of transcriptional repressor activity (demonstrated by luciferase assays) and in some cases aberrant subcellular localization, leading to an autosomal dominant neurodevelopmental disorder with speech/language delays, growth abnormalities, congenital diaphragmatic hernia, cervical spine abnormalities, and ptosis. Luciferase transcriptional repressor assays; subcellular localization assays; clinical phenotyping of 8 individuals with de novo FOXP4 variants Genetics in medicine High 33110267
2021 FOXP4 directly activates UCP1 transcription in thermogenic adipocytes, as demonstrated by chromatin immunoprecipitation and luciferase assays; FoxP4 is induced by heat shock factor protein 1 (HSF1) via a heat shock response element in the FoxP4 proximal promoter, and gain/loss-of-function studies show FoxP4 regulates multiple brown and beige fat genes and affects oxygen consumption in isolated adipocytes. Chromatin immunoprecipitation (ChIP); luciferase reporter assay; gain- and loss-of-function in adipocytes; oxygen consumption assay; in vivo thermogenic stimulation Journal of lipid research Medium 34384787
2022 FOXP4 deficiency in beige adipocyte progenitors impairs early beige cell differentiation, whereas ablation of Foxp4 in differentiated adipocytes potentiates thermogenesis capacity upon cold exposure by de-repressing the thermogenic program; this effect is specific to beige adipocytes and not brown adipocytes, establishing FOXP4 as a transcriptional repressor of the beige thermogenic program in differentiated cells. Beige adipocyte-specific conditional knockout mice; cold exposure experiments; gene expression analysis Development Medium 35297993
2022 FOXP4 promotes proliferation and inhibits squamous differentiation of atypical cervical cells via an ELF3-dependent pathway; FOXP4 knockdown in W12 (HPV16+) cells attenuated proliferation and induced squamous differentiation in monolayer and organotypic raft cultures through regulation of ELF3. FOXP4 siRNA knockdown in W12 cells; monolayer and organotypic raft culture; proliferation assays; gene expression analysis; ELF3 pathway analysis Cancer science Medium 35838233
2022 FOXP1 and FOXP4 form a complex in vivo in hair follicle cells; combined (but not individual) conditional knockout of Foxp1/4 in hair follicles causes additive defects including precocious HFSC activation, hair cycling acceleration, hair loss, downregulation of Fgf18 and Bmp6 in bulge cells, and apoptosis of K6+ inner bulge cells (the stem cell niche), leading to destruction of the niche. Co-immunoprecipitation (complex formation in vitro and in vivo); hair follicle-specific conditional knockout mice (single and double KO); immunofluorescence; apoptosis assays; gene expression Stem cells Medium 35759955
2023 FOXP4 knockdown or dominant-negative inhibition in the mouse neocortex abolishes apical condensation of N-cadherin in radial glial cells (RGCs), disrupts adherens junction integrity in the ventricular zone, impedes radial migration of cortical neurons, and causes ectopic neurogenesis; N-cadherin overexpression in RGCs rescues the migration and differentiation defects, placing FOXP4 upstream of N-cadherin-based adherens junctions in cortical development. In utero shRNA knockdown and dominant-negative inhibition; N-cadherin overexpression rescue; immunofluorescence; cortical neuron migration analysis in mice Neuroscience bulletin Medium 36646976
2023 FOXP4 promotes proliferation of human spermatogonial stem cells (SSCs); conditional inactivation of FOXP4 in human SSC lines suppressed proliferation and significantly activated apoptosis, and FOXP4 expression was found to specifically mark a subset of spermatogonia with stem cell potential. Conditional inactivation in human SSC lines; single-cell RNA sequencing analysis; proliferation and apoptosis assays Asian journal of andrology Medium 36018067
2024 FOXP4 directly induces PTK7 (a Wnt co-receptor) transcription in ovarian cancer, thereby activating the Wnt/β-catenin signaling pathway and promoting malignant phenotype; RNA sequencing in FOXP4-deficient cells identified PTK7 as a downstream target, and restoration of PTK7 reversed the effects of FOXP4 loss. RNA sequencing in FOXP4-deficient cells; ChIP or functional promoter assays (inferred from 'directly induces'); PTK7 rescue experiments; Wnt pathway activity assays Cell death & disease Medium 38740744
2024 FOXP4 is a direct transcriptional target of YAP1 in gastric cancer; FOXP4 upregulation by YAP1 maintains cancer stemness by transcriptionally activating SOX12; FOXP4 loss impairs spheroid formation and stemness marker expression, while a small-molecule screen identified 42-(2-tetrazolyl) rapamycin as a FOXP4 inhibitor. RNA sequencing; functional loss-of-function (spheroid, stemness markers); small-molecule inhibitor screen; ChIP or transcriptional assays for YAP1→FOXP4 and FOXP4→SOX12; in vivo xenograft Cancer research Medium 39047223
2024 FOXP4 promotes radioresistance in colorectal cancer by transcriptionally activating GPX4 (glutathione peroxidase 4) via binding to the GPX4 promoter through its forkhead domain, thereby suppressing ferroptosis; doxorubicin promotes FOXP4 ubiquitination and degradation, reducing GPX4 expression and increasing radiosensitivity. Patient-derived organoids (PDOs); transcriptome analysis; ChIP/CUT&Tag for FOXP4 binding to GPX4 promoter; ubiquitination assay; clonogenic and xenograft assays Advanced science Medium 40789053
2021 FOXP4 transcriptionally activates β-catenin (CTNNB1) expression in esophageal squamous cell carcinoma cells, thereby promoting Wnt/β-catenin signaling and malignant progression; this was established by luciferase reporter assay showing FOXP4 drives β-catenin transcription. Luciferase reporter assay; shRNA knockdown; gene expression analysis; RIP and ChIP (for FOXP4-AS1 mechanism upstream) Frontiers in oncology Low 34970490
2021 FOXP4 directly binds the promoter of LEF-1 and activates Wnt signaling in laryngeal squamous cell carcinoma, as confirmed by chromatin immunoprecipitation and luciferase reporter assays; FOXP4 also participates in EMT regulation in this cancer type. ChIP assay; luciferase reporter assay; gain- and loss-of-function experiments; microarray for differential transcript identification Molecular medicine reports Medium 34590150
2024 FOXP4 transcriptionally represses FBXW7 (a tumor suppressor) in thyroid cancer cells; FOXP4 protein binding to the FBXW7 promoter was confirmed by ChIP assay, and FBXW7 overexpression reversed the effects of FOXP4-driven proliferation, migration, and EMT. ChIP assay; luciferase reporter; gain- and loss-of-function; xenograft Heliyon Medium 38293397
2024 FOXP4 transcriptionally activates NDST2 in hepatocellular carcinoma cells after incomplete thermal ablation (heat stress), as validated by CUT&Tag experiments showing FOXP4 binding to NDST2 regulatory regions; suppression of FOXP4 reduced NDST2 expression and weakened cancer cell progression. CUT&Tag; qRT-PCR; Western blotting; CCK-8, scratch, Transwell assays Journal of hepatocellular carcinoma Medium 39429915
2025 Combined but not individual deficiency of FOXP1 and FOXP4 in FOXP3+ committed Tregs causes lymphoproliferation, inflammation, autoimmunity, and early lethality; loss of both proteins results in an activated/effector phenotype with compromised Treg suppressive function, enhanced germinal center response, and proinflammatory cytokine production; FOXP1 and FOXP4 both bind to Il2ra (CD25) promoter regions to regulate CD25 expression, as demonstrated by promoter-binding assays. Foxp3-Cre conditional knockout of Foxp1, Foxp4, or both; flow cytometry; cytokine assays; promoter-binding assays for Il2ra; immune phenotyping JCI insight High 40794436
2024 An enhancer variant (rs10223516) in FOXP4 modulates FOXP4 expression through long-range chromatin interaction; the T allele shows higher enhancer activity; upregulated FOXP4 promotes ESCC development in vivo; ChIP-seq and RNA-seq revealed FOXP4 transcriptionally activates CYP26B1 and MYC in ESCC cells. Three-stage case-control GWAS; functional enhancer assay; in vivo ESCC model; ChIP-seq; RNA-seq Cancer research Medium 40228145
2024 METTL14-induced N6-methyladenosine (m6A) modification of FOXP4 mRNA in HBV-infected hepatocellular carcinoma cells enhances FOXP4 mRNA stability and increases FOXP4 mRNA levels; HBV gene expression activates the PI3K/AKT pathway via modulation of FOXP4 mRNA stability. m6A modification analysis; mRNA stability assays; PI3K/AKT pathway analysis; HBV infection model Journal of Cancer Low 39513116
2024 FOXP4 is highly expressed in anterior segment structures relevant to the drainage angle (periocular mesenchyme, iris, ciliary body, cornea) in embryonic mouse eyes; a FOXP4 missense variant (p.Q478R) in the forkhead domain acts as a hypomorphic allele retaining transcriptional activity but frequently mislocalizing to cytosolic aggregates, suggesting protein instability underlies pathogenicity in angle closure glaucoma. Exome sequencing; immunostaining in embryonic mouse eyes; YFP-tagged mutant/WT protein expression in HEK-293T and ARPE-19 cells for nuclear localization; SRPX2-luciferase reporter for transcriptional activity Investigative ophthalmology & visual science Medium 40637512

Source papers

Stage 0 corpus · 77 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2019 LncRNA FOXP4-AS1 is activated by PAX5 and promotes the growth of prostate cancer by sequestering miR-3184-5p to upregulate FOXP4. Cell death & disease 104 31209207
2018 CircRNA ZNF609 functions as a competitive endogenous RNA to regulate FOXP4 expression by sponging miR-138-5p in renal carcinoma. Journal of cellular physiology 97 30478938
2019 Circular RNA circABCC4 as the ceRNA of miR-1182 facilitates prostate cancer progression by promoting FOXP4 expression. Journal of cellular and molecular medicine 71 31270953
2002 Foxp4: a novel member of the Foxp subfamily of winged-helix genes co-expressed with Foxp1 and Foxp2 in pulmonary and gut tissues. Mechanisms of development 70 14516685
2003 FoxP4, a novel forkhead transcription factor. Biochimica et biophysica acta 66 12818433
2021 Exosomal miR-101-3p and miR-423-5p inhibit medulloblastoma tumorigenesis through targeting FOXP4 and EZH2. Cell death and differentiation 65 34294888
2015 MicroRNA-338-3p inhibits cell proliferation in hepatocellular carcinoma by target forkhead box P4 (FOXP4). International journal of clinical and experimental pathology 57 25755720
2016 Up-regulation of microRNA-491-5p suppresses cell proliferation and promotes apoptosis by targeting FOXP4 in human osteosarcoma. Cell proliferation 55 27704627
2014 Transcriptional regulation by FOXP1, FOXP2, and FOXP4 dimerization. Journal of molecular neuroscience : MN 55 25027557
2008 Expression of Foxp4 in the developing and adult rat forebrain. Journal of neuroscience research 52 18561326
2020 LncRNA FOXP4-AS1 Is Involved in Cervical Cancer Progression via Regulating miR-136-5p/CBX4 Axis. OncoTargets and therapy 49 32256085
2019 Upregulation of FoxP4 in HCC promotes migration and invasion through regulation of EMT. Oncology letters 49 30930991
2015 The FOXP1, FOXP2 and FOXP4 transcription factors are required for islet alpha cell proliferation and function in mice. Diabetologia 48 26021489
2018 FOXP4-AS1 participates in the development and progression of osteosarcoma by downregulating LATS1 via binding to LSD1 and EZH2. Biochemical and biophysical research communications 47 29859193
2015 FOXP4 modulates tumor growth and independently associates with miR-138 in non-small cell lung cancer cells. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 40 25994569
2018 Circular RNA circMYO9B facilitates breast cancer cell proliferation and invasiveness via upregulating FOXP4 expression by sponging miR-4316. Archives of biochemistry and biophysics 35 29702064
2012 Replication and fine mapping for association of the C2orf43, FOXP4, GPRC6A and RFX6 genes with prostate cancer in the Chinese population. PloS one 35 22662242
2020 CircRNA_0058063 functions as a ceRNA in bladder cancer progression via targeting miR-486-3p/FOXP4 axis. Bioscience reports 34 32181485
2012 Foxp4 is dispensable for T cell development, but required for robust recall responses. PloS one 34 22912696
2020 Exosomal miR-3180-3p inhibits proliferation and metastasis of non-small cell lung cancer by downregulating FOXP4. Thoracic cancer 33 33350095
2002 Foxp4: a novel member of the Foxp subfamily of winged-helix genes co-expressed with Foxp1 and Foxp2 in pulmonary and gut tissues. Gene expression patterns : GEP 33 12617805
2015 Differential coexpression of FoxP1, FoxP2, and FoxP4 in the Zebra Finch (Taeniopygia guttata) song system. The Journal of comparative neurology 30 25556631
2020 YY1-induced upregulation of FOXP4-AS1 and FOXP4 promote the proliferation of esophageal squamous cell carcinoma cells. Cell biology international 29 32159250
2010 Foxp4 is essential in maintenance of Purkinje cell dendritic arborization in the mouse cerebellum. Neuroscience 27 20951773
2019 Differential Song Deficits after Lentivirus-Mediated Knockdown of FoxP1, FoxP2, or FoxP4 in Area X of Juvenile Zebra Finches. The Journal of neuroscience : the official journal of the Society for Neuroscience 26 31641053
2019 Upregulation of FOXP4 in breast cancer promotes migration and invasion through facilitating EMT. Cancer management and research 25 31040716
2020 Heterozygous variants that disturb the transcriptional repressor activity of FOXP4 cause a developmental disorder with speech/language delays and multiple congenital abnormalities. Genetics in medicine : official journal of the American College of Medical Genetics 23 33110267
2021 CircRPPH1 serves as a sponge for miR-296-5p to enhance progression of breast cancer by regulating FOXP4 expression. American journal of translational research 21 34377235
2019 The carcinogenic complex lncRNA FOXP4-AS1/EZH2/LSD1 accelerates proliferation, migration and invasion of gastric cancer. European review for medical and pharmacological sciences 21 31646567
2021 LncRNA FOXP4-AS1 Promotes Progression of Ewing Sarcoma and Is Associated With Immune Infiltrates. Frontiers in oncology 18 34765540
2021 LncRNA FOXP4-AS1 Promotes the Progression of Esophageal Squamous Cell Carcinoma by Interacting With MLL2/H3K4me3 to Upregulate FOXP4. Frontiers in oncology 18 34970490
2020 LncRNA SNHG16 Promotes the Progression of Laryngeal Squamous Cell Carcinoma by Mediating miR-877-5p/FOXP4 Axis. OncoTargets and therapy 18 32547087
2019 Upregulation of long non-coding RNA FOXP4-AS1 and its regulatory network in hepatocellular carcinoma. OncoTargets and therapy 18 31695417
2024 FOXP4-AS1 promotes CD8+ T cell exhaustion and esophageal cancer immune escape through USP10-stabilized PD-L1. Immunologic research 17 38687433
2021 miR-491-5p inhibits the proliferation and migration of A549 cells by FOXP4. Experimental and therapeutic medicine 16 33936279
2024 FOXP4-mediated induction of PTK7 activates the Wnt/β-catenin pathway and promotes ovarian cancer development. Cell death & disease 15 38740744
2019 LncRNA LOC105372579 promotes proliferation and epithelial-mesenchymal transition in hepatocellular carcinoma via activating miR-4316/FOXP4 signaling. Cancer management and research 15 31114338
2024 FOXP4 Is a Direct YAP1 Target That Promotes Gastric Cancer Stemness and Drives Metastasis. Cancer research 14 39047223
2021 Long non-coding RNA FOXP4-AS1 facilitates the biological functions of hepatocellular carcinoma cells via downregulating ZC3H12D by mediating H3K27me3 through recruitment of EZH2. Cell biology and toxicology 14 34545456
2019 MiR-3196, a p53-responsive microRNA, functions as a tumor suppressor in hepatocellular carcinoma by targeting FOXP4. American journal of cancer research 13 31911853
2015 Association of FOXP4 Gene with Prostate Cancer and the Cumulative Effects of rs4714476 and 8q24 in Chinese Men. Clinical laboratory 13 26642711
2021 circ‑0000212 promotes cell proliferation of colorectal cancer by sponging miR‑491 and modulating FOXP4 expression. Molecular medicine reports 11 33649850
2020 MicroRNA-4651 represses hepatocellular carcinoma cell growth and facilitates apoptosis via targeting FOXP4. Bioscience reports 10 32436934
2023 FOXP4 promotes proliferation of human spermatogonial stem cells. Asian journal of andrology 9 36018067
2020 Long noncoding RNA LINC00858 promotes the proliferation, migration and invasion of gastric cancer cells via the miR-363-3p/FOXP4 axis. European review for medical and pharmacological sciences 9 33015780
2019 MicroRNA-299-3p/FOXP4 Axis Regulates the Proliferation and Migration of Oral Squamous Cell Carcinoma. Technology in cancer research & treatment 9 31500519
2022 FOXP4-AS1 Inhibits Papillary Thyroid Carcinoma Proliferation and Migration Through the AKT Signaling Pathway. Frontiers in oncology 8 35720005
2021 FOXP4 promotes laryngeal squamous cell carcinoma progression through directly targeting LEF‑1. Molecular medicine reports 8 34590150
2023 LncRNA FOXP4-AS1 silencing inhibits metastasis and epithelial-mesenchymal transition in nasopharyngeal carcinoma via miR-136-5p/MAPK1. Anti-cancer drugs 7 36961080
2022 LncRNA FOXP4-AS promotes the progression of non-small cell lung cancer by regulating the miR-3184-5p/EIF5A axis. Journal of tissue engineering and regenerative medicine 7 34921595
2022 FOXP4 inhibits squamous differentiation of atypical cells in cervical intraepithelial neoplasia via an ELF3-dependent pathway. Cancer science 7 35838233
2024 Emerging roles of long non-coding RNA FOXP4-AS1 in human cancers: From molecular biology to clinical application. Heliyon 6 39539976
2022 Downregulation of microRNA‑423‑5p suppresses TGF‑β1‑induced EMT by targeting FOXP4 in airway fibrosis. Molecular medicine reports 6 35642665
2021 LncRNA RP11-116G8.5 promotes the progression of lung squamous cell carcinoma through sponging miR-3150b-3p/miR-6870-5p to upregulate PHF12/FOXP4. Pathology, research and practice 6 34500373
2021 The forkhead box transcription factor FoxP4 regulates thermogenic programs in adipocytes. Journal of lipid research 5 34384787
2024 Prediction and assessment of deleterious and disease causing nonsynonymous single nucleotide polymorphisms (nsSNPs) in human FOXP4 gene: An in - silico study. Heliyon 4 38994097
2022 Foxp1 and Foxp4 Deletion Causes the Loss of Follicle Stem Cell Niche and Cyclic Hair Shedding by Inducing Inner Bulge Cell Apoptosis. Stem cells (Dayton, Ohio) 4 35759955
2018 Retracted: Long Noncoding RNA SOX2OT Accelerates the Carcinogenesis of Wilms' Tumor Through ceRNA Through miR-363/FOXP4 Axis. DNA and cell biology 4 30481065
2025 Normal Treg homeostasis and suppressive function require both FOXP1 and FOXP4. JCI insight 3 40794436
2024 Polymorphisms of IFN signaling genes and FOXP4 influence the severity of COVID-19. BMC infectious diseases 3 38429664
2024 Androgen-responsive FOXP4 is a target for endometrial carcinoma. Communications biology 3 38890503
2024 Incomplete Thermal Ablation-Induced FOXP4-Mediated Promotion of Malignant Progression in Liver Cancer via NDST2. Journal of hepatocellular carcinoma 3 39429915
2024 METTL14 Induced N6-Methyladenosine Modification of FOXP4 mRNA in HBV-HCC. Journal of Cancer 3 39513116
2023 Inhibition of Foxp4 Disrupts Cadherin-based Adhesion of Radial Glial Cells, Leading to Abnormal Differentiation and Migration of Cortical Neurons in Mice. Neuroscience bulletin 3 36646976
2022 The FOXP4-AS1/miR-3130-3p/SP4 feedback loop is associated with prostate cancer. Cellular and molecular biology (Noisy-le-Grand, France) 3 37114256
2025 Patients-Derived Organoids Sequencing-based FOXP4 Facilitates Radioresistance by Transcriptionally Modifying GPX4 to Regulate ferroptosis in Colorectal Cancer. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 2 40789053
2024 Transcription factor FOXP4 inversely governs tumor suppressor genes and contributes to thyroid cancer progression. Heliyon 2 38293397
2024 Participation of Long Noncoding RNA FOXP4-AS1 in the Development and Progression of Endometrioid Carcinoma with Epigenetically Silencing DUSP5. Cancer biotherapy & radiopharmaceuticals 2 38512300
2022 FOXP4 differentially controls cold-induced beige adipocyte differentiation and thermogenesis. Development (Cambridge, England) 2 35297993
2026 LncRNA FOXP4-AS1 facilitates colorectal cancer invasion and migration by enhancing USP7 interaction with ZEB1. Scientific reports 1 41486276
2025 LINC00917 Promotes Bone Metastasis of Breast Cancer by Targeting the miR-491-5p/FOXP4 Axis. Breast cancer (Dove Medical Press) 1 41133246
2023 The essential role of forkhead box P4 (FOXP4) in thyroid cancer: a study related to The Cancer Genome Atlas and experimental data. Endocrine connections 1 36752821
2023 Hsa_circ_0017956 Acts as miR-758-3p Sponge to Facilitate the Progression of Non-small-Cell Lung Cancer by Regulating FOXP4 Expression. Molecular biotechnology 1 36763305
2022 Corrigendum: LncRNA FOXP4-AS1 promotes the progression of esophageal squamous cell carcinoma by interacting with MLL2/H3K4me3 to upregulate FOXP4. Frontiers in oncology 1 36313704
2026 FOXP4-AS1 suppresses papillary thyroid carcinoma progression by binding to Lactate Dehydrogenase A and suppressing its expression, with implications for metabolic-targeted therapy. Cellular signalling 0 42066828
2025 Cis-Regulatory Alterations in FOXP4 Modulate Esophageal Cancer Susceptibility Induced by Chronic Alcohol Exposure. Cancer research 0 40228145
2025 FOXP4 Variants Are Associated With Plateau Iris and Angle Closure Glaucoma. Investigative ophthalmology & visual science 0 40637512

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