Affinage

CDX1

Homeobox protein CDX-1 · UniProt P47902

Length
265 aa
Mass
28.1 kDa
Annotated
2026-04-28
100 papers in source corpus 45 papers cited in narrative 45 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

CDX1 is a homeodomain transcription factor that governs anterior-posterior vertebral patterning and intestinal epithelial differentiation by directly regulating Hox gene expression boundaries and intestinal-specific gene programs. CDX1 is transcriptionally activated by Wnt/β-catenin signaling through LEF/TCF response elements (the dominant input in vivo) and by retinoic acid through a proximal RARE, and sustains its own expression via an autoregulatory loop requiring a CDX1–LEF1 protein complex (PMID:10934025, PMID:14660544, PMID:17537796, PMID:15143193). It transactivates target genes (including KRT20, G6Pase, SALL4, KLF5, and miR-215) through its homeodomain-mediated interaction with TBP/TFIID and cooperation with partners such as C/EBPα and EHF, while restraining proliferation by inhibiting β-catenin/TCF transcriptional output, reducing cyclin D1/D2, and activating differentiation-promoting microRNAs (PMID:12954759, PMID:17158164, PMID:10660624, PMID:25775580, PMID:35606410). CDX1 functions as a tumor suppressor in the distal colon, where its promoter is frequently silenced by CpG hypermethylation in colorectal carcinoma, and its loss cooperates with APC and Cdx2 mutations to promote tumorigenesis (PMID:14704280, PMID:25320087).

Mechanistic history

Synthesis pass · year-by-year structured walk · 18 steps
  1. 1995 High

    Establishing that CDX1 is a direct upstream regulator of Hox gene expression and vertebral identity resolved how caudal-type homeobox genes connect to the vertebral patterning code.

    Evidence Cdx1 knockout mice showing anterior homeotic transformations and posterior shifts in Hox expression, plus in vitro transactivation of Hoxa-7

    PMID:7585967

    Open questions at the time
    • Mechanism by which Cdx1 selects specific Hox targets was unknown
    • Redundancy with other Cdx family members was untested
    • Whether Cdx1 regulates Hox genes directly via enhancer binding in vivo was not shown
  2. 2000 High

    Identification of Wnt/β-catenin and retinoic acid as the two major upstream signals directly activating Cdx1 transcription placed Cdx1 at a signaling node integrating morphogen inputs for axial patterning.

    Evidence Functional LEF/TCF elements and an atypical RARE in the Cdx1 promoter demonstrated by reporter assays, EMSA, and Tcf4-null mice losing Cdx1 expression

    PMID:10934025 PMID:10938132

    Open questions at the time
    • Relative contribution of Wnt vs RA to Cdx1 expression in vivo was unclear
    • Whether the two inputs are independent or synergistic in vivo was unresolved
  3. 2000 High

    Demonstrating that Cdx1 inhibits intestinal cell proliferation by specifically reducing cyclin D1/D2 established a growth-suppressive function distinct from its patterning role.

    Evidence Inducible Cdx1 expression in IEC-6 cells causing G0/G1 arrest with specific cyclin D1/D2 reduction

    PMID:10660624

    Open questions at the time
    • Whether cyclin D reduction is transcriptionally direct was unknown
    • Relationship to Cdx1's effect on β-catenin/TCF output was not connected
  4. 2001 High

    Genetic epistasis showing Wnt3a is required upstream of Cdx1 for vertebral patterning, combined with demonstration of Wnt/RA synergy and Cdx1 autoregulation, defined the signaling hierarchy and positive feedback loop maintaining Cdx1 expression.

    Evidence Wnt3a mutant mice with reduced Cdx1 and homeotic transformations; LEF/TCF mutagenesis and autoregulatory loop demonstration in reporters

    PMID:11335109 PMID:11784033

    Open questions at the time
    • Molecular mechanism of autoregulation (direct or indirect) was unresolved
    • Whether autoregulation requires a Cdx1–LEF1 protein complex was unknown
  5. 2002 High

    Compound Cdx1/Cdx2 mutant analysis demonstrated cooperative, dose-dependent regulation of Hox boundaries along the entire vertebral column, establishing functional redundancy between the two paralogues.

    Evidence Cdx1/Cdx2 double-mutant mice with extensive homeotic transformations exceeding single mutants

    PMID:11959827

    Open questions at the time
    • Whether functional equivalence is complete or context-dependent was untested
    • Cdx4 contribution was not assessed
  6. 2002 Medium

    Showing that CDX1 inhibits β-catenin/TCF transcriptional activity without affecting β-catenin levels identified a negative feedback mechanism whereby a Wnt target gene restrains the Wnt output.

    Evidence Stable CDX1 expression in multiple colon cancer lines reducing TCF reporter activity dose-dependently; APC(Min/+) polyp analysis

    PMID:15215241

    Open questions at the time
    • Direct molecular mechanism of β-catenin/TCF inhibition by CDX1 was not identified
    • Whether this involves direct protein interaction with TCF was unknown
  7. 2003 High

    Mapping the CDX1 homeodomain–TBP interaction and its requirement for G6Pase transactivation revealed how CDX1 mechanistically engages the basal transcription machinery, distinguishing it from CDX2.

    Evidence EMSA, TATA-box mutagenesis, and luciferase reporters showing CDX1-specific TBP-dependent transactivation of G6Pase

    PMID:12954759

    Open questions at the time
    • Structural basis of CDX1–TBP interaction was not determined
    • Whether Mediator engagement is direct was unresolved
  8. 2004 High

    Discovery that CDX1 autoregulation operates through a direct CDX1 homeodomain–LEF1 B-box protein complex explained how Wnt signaling and CDX1 converge on a single cis-regulatory element for self-sustaining expression.

    Evidence Co-IP and GST pulldown mapping the interaction domains; compound Cdx1/Wnt3a(vt) mutant genetic validation

    PMID:15143193

    Open questions at the time
    • Whether the CDX1–LEF1 complex recruits distinct co-activators was unknown
    • Crystal structure of the complex was not obtained
  9. 2004 High

    Demonstrating frequent CDX1 promoter hypermethylation in colorectal cancer with demethylation-dependent rescue of expression established an epigenetic silencing mechanism contributing to loss of CDX1 tumor suppression.

    Evidence Bisulfite sequencing and MSP across 37 CRC cell lines; 5-aza-2′-deoxycytidine restoring CDX1 expression

    PMID:14704280

    Open questions at the time
    • Causal relationship between methylation-driven CDX1 loss and tumor progression was not established in vivo
    • Which DNA methyltransferases are responsible was not identified
  10. 2004 High

    Transgenic CDX1 expression in stomach generating complete intestinal metaplasia including Paneth cells showed CDX1 is sufficient to specify all four intestinal cell lineages, revealing a broader differentiation capacity than CDX2.

    Evidence Gastric CDX1-transgenic mice producing goblet, absorptive, enteroendocrine, and Paneth cells

    PMID:15361487

    Open questions at the time
    • Whether CDX1 directly activates lineage-specific transcription factors was unknown
    • Mechanism distinguishing CDX1 from CDX2 in Paneth cell specification was unresolved
  11. 2007 High

    In vivo cis-regulatory dissection showed that the LEF/TCF response element is the primary determinant of Cdx1 expression and vertebral patterning, subordinating the RARE input, and that RA responsiveness itself requires intact Wnt signaling through the LRE.

    Evidence LRE-null and RARE-null knock-in mice; LRE mutation phenocopies Cdx1-null while RARE mutation produces limited defects

    PMID:17537796

    Open questions at the time
    • How Wnt signaling gates RA responsiveness mechanistically was not explained
    • Other cis-elements contributing to Cdx1 regulation in non-mesodermal tissues were not tested
  12. 2007 High

    Identifying CDX1 as a physical interactor and inhibitor of SMAD3-dependent transcription (opposing CDX2's stimulatory effect) revealed a mechanism for differential TGF-β/BMP pathway modulation by the two CDX paralogues.

    Evidence Reciprocal co-IP and GST pulldown; 10-fold inhibition of SMAD3/SMAD4 reporters by CDX1; DSS colitis in Cdx1-null mice

    PMID:17595234

    Open questions at the time
    • Whether SMAD3 interaction competes with TBP binding was unknown
    • In vivo relevance to intestinal homeostasis beyond inflammation was not established
  13. 2009 High

    Cdx2 knock-in at the Cdx1 locus fully rescuing the skeletal phenotype demonstrated complete functional equivalence for vertebral patterning, proving that expression differences rather than protein divergence account for non-redundancy.

    Evidence Cdx2 knock-in replacing Cdx1 coding region; complete rescue of vertebral transformations and Hox expression

    PMID:19328777

    Open questions at the time
    • Whether equivalence extends to intestinal functions was not tested
    • Endogenous Cdx2 was still present, complicating interpretation of dosage effects
  14. 2010 High

    Showing Cdx1 directly represses Mafb in the hindbrain expanded CDX1's known role from mesoderm to neural tube patterning and identified a direct transcriptional repression target.

    Evidence ChIP demonstrating Cdx1 binding to Mafb regulatory sequences; altered Mafb expression in Cdx1-null neural tube

    PMID:21098558

    Open questions at the time
    • Mechanism of repression (co-repressor recruitment) was not identified
    • Other neural tube targets of Cdx1 were not catalogued
  15. 2012 High

    Identification of SALL4 and KLF5 as direct CDX1 target genes that convert gastric cells into intestinal stem-like progenitors provided a molecular mechanism for CDX1-driven intestinal metaplasia.

    Evidence Microarray, siRNA knockdown of SALL4/KLF5 suppressing CDX1-induced intestinal markers in gastric cells; confirmation in human/mouse IM specimens

    PMID:23112162

    Open questions at the time
    • Whether CDX1 binds SALL4 and KLF5 promoters directly (ChIP) was not shown
    • Sufficiency of SALL4/KLF5 without CDX1 was not tested
  16. 2014 High

    Compound Cdx1/Cdx2 loss in APC(Min/+) mice increasing distal colon tumors provided definitive in vivo genetic evidence that CDX1 functions as a tumor suppressor cooperating with CDX2 specifically in the distal colon.

    Evidence Conditional somatic Cdx2 deletion combined with Cdx1-null in APC(Min/+) mice; site-specific tumor quantification

    PMID:25320087

    Open questions at the time
    • Molecular targets mediating tumor suppression in distal colon were not identified
    • Whether CDX1 tumor suppression is cell-autonomous was not established
  17. 2015 High

    Demonstrating CDX1 directly activates miR-215 transcription, which in turn represses stemness genes including BMI1, connected CDX1's differentiation-promoting activity to a microRNA effector pathway in colorectal cancer stem cells.

    Evidence ChIP-PCR and promoter luciferase for CDX1 binding to miR-215 locus; miR-215 overexpression/knockdown in FACS-enriched cancer stem cells

    PMID:25775580

    Open questions at the time
    • Full spectrum of miR-215 targets downstream of CDX1 was not determined
    • In vivo validation of CDX1–miR-215–BMI1 axis in tumors was lacking
  18. 2022 High

    Identification of EHF as a direct CDX1 interaction partner that cooperatively drives colonic differentiation gene expression (VIL1) and suppresses tumorigenesis revealed a new co-factor axis for CDX1 function in the colon.

    Evidence Co-IP of EHF–CDX1 via PNT domain; VIL1 promoter co-activation; compound Ehf/Cdx1 knockout mice with disrupted colonic differentiation and enhanced tumor progression

    PMID:35606410

    Open questions at the time
    • Genome-wide co-occupancy of EHF and CDX1 was not mapped
    • Whether EHF modifies CDX1's interaction with TBP/Mediator is unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • A genome-wide map of direct CDX1 chromatin occupancy in normal intestinal epithelium is lacking, and the structural basis for CDX1's selective interactions with TBP, LEF1, SMAD3, and EHF — and how these are coordinated on different promoters — remains unresolved.
  • No genome-wide ChIP-seq for CDX1 in normal intestine has been reported
  • No crystal or cryo-EM structure of CDX1 or its complexes exists
  • How CDX1 selectively activates versus represses targets (e.g., Hox activation vs Mafb repression) is mechanistically unexplained

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 7 GO:0003677 DNA binding 6
Localization
GO:0005634 nucleus 3
Pathway
R-HSA-1266738 Developmental Biology 6 R-HSA-74160 Gene expression (Transcription) 6 R-HSA-162582 Signal Transduction 5 R-HSA-1643685 Disease 3
Partners
CEBPACTNNB1EHFLEF1SMAD3TBP

Evidence

Reading pass · 45 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 Cdx1 loss-of-function (homologous recombination knockout) causes anterior homeotic transformations of vertebrae, accompanied by posterior shifts of Hox gene expression domains in somitic mesoderm; putative Cdx1-binding sites were identified in Hox gene control regions, and in vitro transactivation of Hoxa-7 was demonstrated, indicating direct regulation of Hox genes by Cdx1. Homologous recombination knockout, in vitro transactivation assay, in situ hybridization of Hox expression domains Cell High 7585967
2000 Cdx1 is a direct transcriptional target of the Wnt/β-catenin signaling pathway; the Cdx1 promoter contains functional TCF/LEF-binding motifs that bind TCF/LEF1/β-catenin complexes and mediate β-catenin-dependent transactivation; Tcf4-deficient mice lose Cdx1 protein in small intestinal epithelium. Promoter reporter assays, gel shift (EMSA), in vivo Wnt stimulation of ES cells and embryonic endoderm, Tcf4 knockout mice analysis Development (Cambridge, England) High 10934025
2000 Cdx1 is a direct retinoic acid (RA) target gene; an atypical RA response element (RARE) in the proximal Cdx1 promoter mediates retinoid-dependent activation, providing an indirect pathway for RA to regulate Hox gene expression and vertebral patterning. Promoter reporter assay, RARE mutagenesis, retinoid receptor loss-of-function genetics Molecular and cellular biology High 10938132
2000 Cdx1 inhibits intestinal epithelial cell proliferation by arresting cells in G0/G1, specifically by reducing cyclin D1 and D2 protein levels (without affecting other cyclins or CDK inhibitors), leading to increased hypophosphorylated Rb and p130. Stable transfection and adenoviral induction of Cdx1 in rat IEC-6 cells, flow cytometry, Western blot for cyclins and Rb The Journal of biological chemistry High 10660624
2001 Cdx1 expression is regulated by both Wnt3a signaling (through functional LEF/TCF response elements) and by an autoregulatory loop; Wnt3a and retinoic acid synergize strongly to activate Cdx1, and Cdx1 positively regulates its own expression. LEF/TCF motif mutagenesis in promoter reporters, Wnt3a hypomorph (vestigial tail) mouse analysis, EMSA, compound mutant analysis Developmental biology High 11784033
2001 CDX1 regulates PAP I (Pancreatitis Associated Protein I) gene expression in intestinal cells by directly binding the PAP I promoter; PAP I acts downstream of Cdx1 to promote intestinal cell proliferation via an autocrine/paracrine mechanism. Stable transfection of Cdx1 in IEC-6 cells, promoter binding/deletion assay, adenoviral PAP I expression, antisense inhibition of PAP I European journal of cell biology Medium 11302520
2001 Wnt-3a is required for Cdx1 expression in the primitive streak and tail bud; Wnt-3a mutant mice show reduced Cdx1 expression concomitant with vertebral homeotic transformations, placing Wnt-3a upstream of Cdx1 in the axial patterning pathway. Wnt-3a mutant mouse analysis, in situ hybridization, skeletal phenotyping Mechanisms of development High 11335109
2001 Cdx1 expression in intestinal epithelial IEC-6 cells activates Ras (increased GTP-bound Ras), modulates Cdc42 and RhoA activities, and accumulates PI3-kinase products; combined inhibition of Ras/Rho and PI3K signaling blocked Cdx1-induced anchorage-independent growth and tumorigenesis. Stable transfection, Ras GTP-loading assay, Rho activity assay, PI3K lipid product measurement, pharmacological inhibition, soft agar and xenograft assays Oncogene Medium 11464284
2002 Cdx1 and Cdx2 double mutants show cooperative anterior homeotic transformations along an extensive vertebral column length and more extensive posterior shifts of Hox gene expression boundaries than single mutants, demonstrating that Cdx1 and Cdx2 cooperate in instructing vertebral progenitors through regulation of Hox gene rostral expression boundaries. Compound mouse mutant analysis, skeletal phenotyping, Hox gene expression analysis by in situ hybridization Development (Cambridge, England) High 11959827
2002 Oncogenic β-catenin/Tcf4 stimulates Cdx1 promoter activity and endogenous Cdx1 mRNA expression in colon cancer cells; conversely, CDX2 inhibits basal and β-catenin-stimulated Cdx1 promoter activity through its homeodomain, acting independently of canonical CDX-binding sites and TCF elements. Transient transfection of promoter reporters, expression of constitutively active β-catenin, CDX2 homeodomain mutant analysis FEBS letters Medium 11997022
2002 CDX1 and Cdx2 expression reduces colon cancer cell proliferation, in part by inhibiting β-catenin/TCF transcriptional activity in a dose-dependent manner without altering β-catenin protein levels or intracellular distribution. Stable transfection of Cdx1/Cdx2 in multiple colon cancer cell lines, TCF reporter assays, β-catenin localization/Western blot, APC(Min/+) mouse polyp analysis The Journal of biological chemistry Medium 15215241
2002 Cdx1 regulates apolipoprotein B mRNA editing during intestinal development; Cdx1 overexpression in IEC-6 cells increases apoB mRNA editing more than 10-fold, associated with upregulation of the editing complex component ACF. Stable transfection of Cdx1 in IEC-6 cells, apoB mRNA editing quantification, ACF protein measurement The Journal of biological chemistry Medium 12493769
2002 Cdx1 belongs to the p53-p21(WAF)-Bcl-2 network in intestinal epithelial cells: wild-type p53 inhibits the Cdx1 promoter; Cdx1 inhibits p21(WAF) promoter activity by binding its TATA-box and activates Bcl-2 promoter P2 through a CDX-binding site, increasing apoptotic resistance. Promoter reporter assays, EMSA for Cdx1 binding to p21 TATA-box, Bcl-2 Western blot, apoptosis resistance assays in IEC-6/SW480 cells Biochemical and biophysical research communications Medium 12270138
2003 CDX1 directly binds the TATA-box region of the glucose-6-phosphatase (G6Pase) promoter and transactivates it, unlike CDX2 which binds but does not transactivate; CDX1-specific transactivation requires interaction with the TATA-binding protein (TBP). EMSA, mutagenesis of CDX-binding sites and TATA box, luciferase reporter assays in HepG2 and CaCo2 cells, stable transfection in IEC-6 cells Nucleic acids research High 12954759
2003 CDX1 is a key mediator of Barrett's metaplasia: conjugated bile salts, TNF-α and IL-1β increase CDX1 mRNA expression primarily through NF-κB signaling, but only when the CDX1 promoter is unmethylated or partially methylated. CDX1 mRNA quantification in esophageal cell lines after cytokine/bile salt treatment, NF-κB inhibition, promoter methylation analysis Proceedings of the National Academy of Sciences of the United States of America Medium 15894614
2003 Retinoic acid regulates Cdx1 expression through an atypical RARE in the proximal promoter; RARE-null mutant mice show reduced Cdx1 expression, vertebral homeotic transformations and altered Hox gene expression, demonstrating that the RARE is required for a subset of Cdx1 function governing vertebral patterning in vivo. RARE mutagenesis knock-in mice, skeletal phenotyping, Hox gene expression analysis Development (Cambridge, England) High 14660544
2004 Cdx1 autoregulation is mediated by a novel complex between the Cdx1 homeodomain and the B box of LEF1; Cdx1 and LEF1 synergize to activate transcription from LEF/TCF response elements in the Cdx1 promoter, and compound Cdx1/Wnt3a(vt) mutants demonstrate convergence of these pathways on Cdx1 expression and vertebral patterning in vivo. Co-immunoprecipitation, GST pulldown mapping of homeodomain-B box interaction, luciferase reporter assays, Cdx1/Wnt3a compound mutant mouse analysis Molecular and cellular biology High 15143193
2004 CDX1 promoter silencing in colorectal carcinoma is primarily due to promoter CpG hypermethylation; treatment with the demethylating agent 5-aza-2'-deoxycytidine restores CDX1 expression in cell lines with partially methylated promoters. Promoter methylation analysis by bisulfite sequencing, methylation-specific PCR, 5-aza-2'-deoxycytidine treatment, RT-PCR expression analysis in 37 CRC cell lines Proceedings of the National Academy of Sciences of the United States of America High 14704280
2004 Cdx1 or Cdx2 expression in COLO 205 colon cancer cells induces E-cadherin-dependent cell-cell adhesion and compaction without changing E-cadherin protein levels; adhesion was Ca2+-dependent and blocked by an anti-E-cadherin antibody, and was associated with acquisition of columnar morphology and differentiation markers. Stable transfection, E-cadherin blocking antibody, Ca2+ chelation, electron microscopy, immunofluorescence American journal of physiology. Gastrointestinal and liver physiology Medium 14977637
2004 CDX1 expression in transgenic mice induces gastric intestinal metaplasia comprising all four intestinal epithelial cell types (including Paneth cells absent from Cdx2-induced metaplasia), with diffuse proliferating cell nuclear antigen (PCNA)-positive cells, demonstrating distinct differentiation and proliferation programs compared to CDX2. Transgenic mouse model with gastric Cdx1 expression, histology, immunohistochemistry for PCNA and cell-type markers Gut High 15361487
2005 COUP-TF transcription factors antagonize RA-induced Cdx1 expression by competing with RXR-RAR heterodimers for binding to the Cdx1 RARE, providing a repressor mechanism that restricts Cdx1 to the caudal embryo. Promoter reporter competition assays, EMSA, in situ hybridization of COUP-TF and Cdx1 expression domains The Journal of biological chemistry Medium 15677473
2006 CDX1 physically interacts with the TATA-binding protein (TBP) via its homeodomain, and is connected to TFIID and Mediator complex members; the N-terminal domain is additionally required for transactivation; the C-terminal domain determines the functional specificity between CDX1 and CDX2 through intramolecular interactions. Co-immunoprecipitation, altered-specificity TBP mutant transcription assay, domain swapping between CDX1 and CDX2, luciferase reporters Nucleic acids research High 17158164
2007 Wnt signaling is the primary pathway governing Cdx1 expression in vivo; LRE (LEF/TCF response element)-null mutant mice phenocopy Cdx1-null vertebral defects (affecting the entire cervical region), while RARE-null mutants have limited defects; LRE mutation also ablates RA-induced Cdx1 expression, demonstrating that Wnt signaling is required for RA responsiveness of Cdx1. Knock-in mice with LRE and/or RARE mutations, skeletal phenotyping, RA treatment, Cdx1 expression analysis Development (Cambridge, England) High 17537796
2007 CDX1 protein physically interacts with SMAD3 (independently of SMAD4) and inhibits SMAD3/SMAD4-dependent transcription approximately 10-fold, whereas CDX2 interacts with SMAD3 and stimulates its activity approximately 5-fold; this differential interaction explains distinct effects on intestinal inflammation outcome in Cdx1-/- vs Cdx2+/- mice. Co-immunoprecipitation, GST-pulldown, luciferase reporter assays, DSS-induced colitis model in Cdx1-/- and Cdx2+/- mice Gut High 17595234
2007 Cdx1 or Cdx2 expression in COLO 205 cells induces E-cadherin-dependent cell-cell adhesion by reducing tyrosine phosphorylation of β-catenin and p120-catenin; restoring β- and p120-catenin tyrosine phosphorylation (by knocking down PTP1B) reversed Cdx-induced cell-cell adhesion. Stable transfection, co-immunoprecipitation of E-cadherin/catenin complexes, phospho-tyrosine immunoprecipitation, PTP1B siRNA knockdown, migration/invasion assays American journal of physiology. Gastrointestinal and liver physiology Medium 17463179
2008 CDX1 expression in cultured esophageal squamous epithelial cells induces Cdx2 protein production; in a Barrett's rat model (esophago-jejunal anastomosis), bile acids dose-dependently increase Cdx1 promoter activity and protein expression, suggesting a sequential Cdx1→Cdx2 cascade in Barrett's epithelium development. Cdx1 promoter luciferase assay, bile acid treatment, transfection of Cdx1 expression vector in esophageal cells, rat Barrett's model, immunohistochemistry Gut Medium 19136512
2008 CDX1 transactivates the COX-2 gene in gastric cancer cells; bile acid induction of CDX1 is mediated by the orphan nuclear receptor SHP, establishing a SHP→CDX1→COX-2 sequential transcriptional cascade. Transfection/overexpression assays, COX-2 reporter, CDX1 promoter assay, SHP siRNA/overexpression Carcinogenesis Medium 18775915
2008 Cdx1 and c-Myc cooperate to induce mucin production and changes in keratin expression characteristic of Barrett's esophagus in immortalized human esophageal keratinocytes grown in organotypic culture. Organotypic culture, microarray-guided candidate identification, retroviral expression of Cdx1 and c-Myc, immunostaining for mucins and keratins PloS one Medium 18953412
2009 CDX1 directly regulates Keratin 20 (KRT20) expression: CDX1 binds CDX elements within 246 bp upstream of the KRT20 transcription start site (confirmed by ChIP), and deletion/mutation of these elements abolishes CDX1-dependent KRT20 promoter activity. Microarray analysis, promoter deletion and mutation analysis, ChIP, immunohistochemistry, expression correlation across 38 CRC cell lines Proceedings of the National Academy of Sciences of the United States of America High 19188603
2008 CDX1 promotes intestinal differentiation by activating PPARγ gene expression through functional interaction with C/EBPα; butyrate increases CDX1-C/EBPα protein interaction, leading to enhanced PPARγ expression. Co-immunoprecipitation, CDX1/C/EBPα co-transfection with PPARγ reporter, butyrate treatment, Western blot FEBS letters Medium 19059241
2009 Cdx2 binds the Cdx1 promoter region in intestinal metaplasia (demonstrated by ChIP and EMSA) and upregulates Cdx1 transcription; siRNA-mediated Cdx2 knockdown reduces Cdx1 promoter activity; the Cdx1 promoter is unmethylated in Cdx2-transgenic mouse intestinal metaplasia. ChIP, EMSA, luciferase reporter assay, siRNA knockdown of Cdx2, bisulfite sequencing of Cdx1 promoter The FEBS journal High 19725873
2009 Cdx2 substituted for Cdx1 in a knock-in model perfectly complements the Cdx1-null skeletal phenotype and Hox gene expression, demonstrating functional redundancy between Cdx1 and Cdx2 for vertebral anteroposterior patterning. Cdx2 knock-in at Cdx1 locus, skeletal phenotyping, Hox gene expression analysis Developmental biology High 19328777
2010 Cdx1 directly represses Mafb expression in the neural tube posterior to the r6/r7 boundary, thereby refining hindbrain patterning; Cdx1 binds regulatory sequences of the Mafb gene as demonstrated by ChIP and in situ hybridization, establishing Mafb as a direct early target of Cdx1. ChIP, immunofluorescence, in situ hybridization, identification of Mafb regulatory sequences responsive to Cdx1 Development (Cambridge, England) High 21098558
2011 CDX1 regulates the ASBT (apical sodium-dependent bile acid transporter) gene; CDX1 binds to six of nine predicted CDX binding sites in the ASBT promoter (verified by EMSA and ChIP in living cells), strongly induces ASBT promoter activity in reporter assays, and CDX1 siRNA knockdown reduces ASBT mRNA in intestinal cells. ChIP, EMSA, luciferase reporter assay, siRNA knockdown, correlation analysis in Barrett's esophagus biopsies American journal of physiology. Gastrointestinal and liver physiology High 22016432
2012 Oxidative stress (H2O2) silences CDX1 in colorectal cancer cells through epigenetic mechanisms: ROS upregulates DNMT1 and HDAC1 expression and activity, enhances DNMT1-HDAC1 association, and increases CDX1 promoter methylation; 5-aza-2'-deoxycytidine reverses these effects. H2O2 treatment, MSP, RT-PCR, Western blot, DNMT1/HDAC1 activity assays, N-acetylcysteine rescue, 5-aza-dC reversal Gene Medium 23618814
2012 CDX1 directly activates SALL4 and KLF5 expression in gastric epithelial cells; CDX1-induced SALL4 and KLF5 convert gastric epithelial cells into intestinal stem-like progenitor cells, which then transdifferentiate into intestinal epithelial cells; inhibiting SALL4 or KLF5 suppresses CDX1-induced intestinal differentiation markers. Gene expression profiling (microarray), siRNA knockdown of SALL4 and KLF5, stable CDX1 expression in gastric epithelial cells, immunohistochemistry in human and mouse IM specimens Proceedings of the National Academy of Sciences of the United States of America High 23112162
2012 NF-κB binding to the CDX1 promoter is methylation-dependent; a methylated CDX1 promoter is associated with closed chromatin structure and reduced NF-κB binding; competitive EMSA and ChIP quantified this interaction, and along the gastritis-metaplasia-carcinoma sequence, CDX1 promoter methylation pattern inversely correlates with NF-κB signaling activity. Competitive EMSA, ChIP, bisulfite sequencing, TNF-α protein expression analysis across gastric tissue stages The American journal of pathology Medium 22749770
2012 Cdx1 induces multiple microRNAs (miR-9, miR-16, miR-22) in colorectal tumor cells that directly bind the CDX2 mRNA 3'UTR and destabilize it; simultaneous mutation of miR-9 and miR-16 binding sites in CDX2 3'UTR was sufficient to block Cdx2 suppression by Cdx1. Microarray miRNA profiling, 3'UTR luciferase reporter with CDX2 binding-site mutations, stable Cdx1 transfection in SW480 cells The Biochemical journal Medium 22849325
2012 Cdx1 directly binds to conserved CDX-binding sites within the HoxC8 early enhancer during Xenopus embryogenesis (demonstrated by ChIP), and Cdx1 overexpression or knockdown causes precocious or delayed HoxC8 expression respectively; differential binding affinity to multiple CDX sites may contribute to the temporal control of Hox activation. Xenopus Cdx1 overexpression and morpholino knockdown, mouse HoxC8 early enhancer reporter in Xenopus, ChIP, mutagenesis of Cdx binding sites FASEB journal : official publication of the Federation of American Societies for Experimental Biology High 22426122
2013 CDX1 exhibits context-dependent transcriptional specificity: in intestinal cells, CDX2 is significantly less potent than CDX1 at activating the Cdx1 promoter autoregulatory response, and in vivo, CDX2 cannot substitute for CDX1 in the autoregulatory loop (unlike in paraxial mesoderm/vertebral patterning), demonstrated by a gene swap approach. Cell-based promoter reporter assay comparing Cdx1 vs Cdx2 potency, in vivo gene swap knock-in model PloS one Medium 23382958
2014 Combined somatic loss of Cdx2 and the Cdx1 null allele in APC(Min/+) mice significantly increases the incidence of tumors in the distal colon relative to APC(Min/+)-Cdx2 mutants alone, demonstrating that Cdx1 functions as a tumor suppressor specifically in the distal colon. Conditional somatic Cdx2 deletion combined with Cdx1 null allele in APC(Min/+) background, tumor quantification by location The Journal of biological chemistry High 25320087
2014 CDX1 restricts invasion of HTR-8/SVneo trophoblast cells by inhibiting MMP-9 expression and increasing TIMP-1 expression, independently of the PI3K/AKT signaling pathway; however, CDX1 itself is regulated by PI3K/AKT signaling. Stable CDX1 transfection, invasion assay (QCM ECMatrix kit), RT-PCR and Western blot for MMP-9/TIMP-1, PI3K inhibitor (perifosine) treatment Placenta Medium 24836459
2014 Transient CDX1 expression in embryonic epicardium promotes epithelial-to-mesenchymal transition (EMT) and migration/differentiation of epicardium-derived cells into α-SMA+ vascular smooth muscle; sustained high-level CDX1 or CDX1 deficiency both attenuate this process, and CDX1 induction alters transcript levels of genes involved in neuronal development, angiogenesis, and cell adhesion. Doxycycline-inducible CDX1 mouse model, primary epicardium culture, ex vivo heart culture, RNA-seq, α-SMA immunostaining, Cdx1 knockout comparison PloS one Medium 25068460
2015 CDX1 directly activates miR-215 transcription (confirmed by ChIP-PCR and promoter luciferase assay); miR-215 mediates repression of cell cycle and stemness genes downstream of CDX1, including the stemness gene BMI1, linking CDX1 to colorectal cancer stem cell differentiation. Small RNA sequencing, ChIP-PCR, promoter luciferase assay, miR-215 overexpression/knockdown, FACS-enriched cancer stem cell comparison, clonogenicity assays Proceedings of the National Academy of Sciences of the United States of America High 25775580
2022 EHF physically interacts with CDX1 via the EHF PNT domain; EHF and CDX1 co-operatively drive transcription of the colonic differentiation marker VIL1; compound genetic deletion of Ehf and Cdx1 in the mouse colon disrupts normal colonic differentiation and enhances colorectal tumour progression. Co-immunoprecipitation identifying EHF-CDX1 interaction, VIL1 promoter reporter co-transfection, compound Ehf/Cdx1 conditional knockout mouse, chromatin remodeling assays, transcriptional profiling Cell death and differentiation High 35606410

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2000 Cdx1 and cdx2 expression during intestinal development. Gastroenterology 506 11040183
1995 Disruption of the murine homeobox gene Cdx1 affects axial skeletal identities by altering the mesodermal expression domains of Hox genes. Cell 291 7585967
2002 Cdx1 and Cdx2 have overlapping functions in anteroposterior patterning and posterior axis elongation. Development (Cambridge, England) 238 11959827
2003 Expression of intestine-specific transcription factors, CDX1 and CDX2, in intestinal metaplasia and gastric carcinomas. The Journal of pathology 236 12474224
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