Affinage

MSX2

Homeobox protein MSX-2 · UniProt P35548

Length
267 aa
Mass
28.9 kDa
Annotated
2026-06-10
100 papers in source corpus 41 papers cited in narrative 41 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MSX2 is a homeodomain transcriptional repressor that acts as a dosage-sensitive node integrating BMP, Wnt, Notch, FGF, and TNF signals to control osteogenic, adipogenic, chondrogenic, and broader cell-fate decisions (PMID:12925529, PMID:15841209, PMID:19407244). It represses transcription largely independently of direct DNA binding through a core suppressor domain (residues 97–208, with residues 132–148 essential) that engages the basal machinery and other regulators: it binds both subunits of TFIIF (RAP74/RAP30) (PMID:9265625), blocks promoter occupancy of partner factors such as the OCFRE-binding protein and Runx2 (PMID:9368026, PMID:15131132), and cooperates with TLE1 to recruit HDAC1 to suppress Runx2-driven osteoblast differentiation (PMID:15060165). MSX2 maintains osteoprogenitors in a proliferative, undifferentiated state—occupying the osteocalcin promoter until it is replaced by Dlx/Runx2 as a differentiation switch (PMID:15456894)—and competes with Dlx5 and inhibits adipogenic factors PPARγ and C/EBPα through protein–protein interaction, biasing mesenchymal progenitors toward bone over fat (PMID:12925529, PMID:15383550, PMID:15175325). Its bone-anabolic output is amplified by paracrine canonical Wnt signaling: MSX2 represses Dkk1 and induces Wnt7a/Wnt7b to drive β-catenin/TCF activity (PMID:15841209, PMID:18487199). Beyond the skeleton, MSX2 represses pluripotency and lineage programs by directly binding promoters—repressing SOX2 while activating Nodal in mesendoderm commitment (PMID:26427715)—and prevents cBAF-dependent chromatin remodeling at syncytiotrophoblast differentiation genes (PMID:34507999). MSX2 protein abundance is tightly controlled by FBXW2-mediated ubiquitin–proteasomal degradation, enhanced by hypoxia-induced VRK2, and by CLU/AKT/DNM1L-driven mitophagic clearance that excludes it from the nucleus to derepress SOX2 (PMID:31548378, PMID:36779631). Gain-of-function P148H enhances homeodomain DNA-binding affinity to cause Boston-type craniosynostosis while paradoxically increasing proteasomal susceptibility, and homeodomain loss-of-function mutations cause parietal foramina by haploinsufficiency, establishing strict gene-dosage requirements for calvarial morphogenesis (PMID:8106171, PMID:10742103, PMID:18786927).

Mechanistic history

Synthesis pass · year-by-year structured walk · 13 steps
  1. 1993 High

    Established MSX2 as a disease gene by linking a homeodomain point mutation to a calvarial malformation, framing the question of how a transcription factor controls suture morphogenesis.

    Evidence Human genetic mapping and segregation of the P148H missense allele, with mouse calvarial in situ localization

    PMID:8106171

    Open questions at the time
    • Did not establish the molecular mechanism of repression
    • Left open how enhanced DNA binding translates to premature suture fusion
  2. 1996 Medium

    Defined MSX2's biochemical activity as a transcriptional repressor that acts independently of its consensus DNA-binding site, with repressor potency mapping outside the homeodomain.

    Evidence DNA-binding site selection, EMSA, and luciferase reporter assays with deletion mutants in cell lines

    PMID:7726844 PMID:8861098

    Open questions at the time
    • Did not identify the protein partners mediating DNA-binding-independent repression
    • Endogenous target genes not defined
  3. 1997 High

    Mapped the core suppressor domain and identified its first mechanistic basis—engagement of the basal transcription factor TFIIF—explaining DNA-binding-independent repression.

    Evidence Systematic deletion mutagenesis with osteoblast reporter assays, Far-Western binding, and RAP74 rescue, plus blocking of OCFREB binding to the FGF-response element

    PMID:9265625 PMID:9368026

    Open questions at the time
    • Did not show TFIIF interaction at endogenous loci by ChIP
    • Did not establish which signaling inputs deploy the suppressor domain
  4. 1999 High

    Identified nuclear protein partners (MINT, Miz1) and positioned MSX2 as a brake on osteoblast differentiation that keeps progenitors proliferative.

    Evidence Far-Western expression cloning, yeast two-hybrid, EMSA, fractionation, and reciprocal gain/loss-of-function in calvarial osteoblasts

    PMID:10328922 PMID:10451362 PMID:9256341

    Open questions at the time
    • Interaction stoichiometry and complex architecture unresolved
    • Whether MINT and Miz1 act on the same target promoters not tested
  5. 2000 High

    Resolved the dosage logic of MSX2 in the skull by contrasting gain-of-function craniosynostosis with loss-of-function parietal foramina.

    Evidence Human mutation identification with in vitro DNA-binding assays and Msx2-null mouse phenotyping including BrdU and epistasis with Msx1

    PMID:10742103 PMID:10742104

    Open questions at the time
    • Direct transcriptional targets governing osteoprogenitor proliferation not enumerated
    • Mechanism of Msx1/Msx2 functional redundancy at the molecular level unclear
  6. 2003 High

    Distinguished two mechanistic modes of MSX2 action—DNA-binding-dependent pro-osteogenic versus protein-interaction-dependent anti-adipogenic—and placed it downstream of BMP2/Wnt via Smad4/Lef1 promoter activation.

    Evidence Structure-function mutagenesis (P148H, T147A) in mesenchymal cells, C/EBPα interaction studies, and Msx2 promoter dissection with ChIP in Smad4-null cells

    PMID:12925529 PMID:14551209

    Open questions at the time
    • How a single protein switches between the two modes in vivo not defined
    • Direct osteogenic DNA targets requiring binding not identified
  7. 2004 High

    Defined MSX2 as the repressive arm of a temporal molecular switch at osteoblast promoters, acting via competition with Dlx5/Runx2 and recruitment of TLE1/HDAC1.

    Evidence Stage-resolved ChIP at osteocalcin, EMSA competition at the ALP promoter, Co-IP and HDAC recruitment assays, and ternary complexes with necdin/MAGE-D1 and Runx2/MINT

    PMID:15060165 PMID:15131132 PMID:15175325 PMID:15272023 PMID:15383550 PMID:15456894

    Open questions at the time
    • Quantitative thresholds for the Dlx5:Msx2 switch in vivo not established
    • Generality of HDAC1/TLE1 recruitment across MSX2 targets untested
  8. 2005 High

    Showed MSX2 acts non-cell-autonomously through secreted Wnt ligands and Dkk1 repression to bias mesenchymal fate toward bone.

    Evidence Conditioned media transfer, TOPGAL reporter mice, nuclear β-catenin imaging, and Dkk1 rescue

    PMID:15841209

    Open questions at the time
    • Whether Wnt ligand induction is direct transcriptional activation by MSX2 not shown here
    • Receiving cell populations in vivo not fully defined
  9. 2008 High

    Expanded MSX2's regulatory inputs (Notch/RBP-Jk, PXR, BMP2/Smad in Runx2-null context) and outputs (Osterix, Ihh, smooth-muscle and cardiac genes), and revealed proteasomal control of the protein.

    Evidence Promoter mutagenesis with RBP-Jk-null cells, ChIP for PXR/p300, Runx2-null epistasis for Osterix, T-box partner pulldowns, and pulse-chase/ubiquitylation with Praja1 E3 ligase

    PMID:17030628 PMID:17516553 PMID:17875939 PMID:18285513 PMID:18682398 PMID:18703512 PMID:18786927 PMID:19407244

    Open questions at the time
    • How distinct upstream signals select between activation and repression outputs unresolved
    • Physiological relevance of Praja1-mediated turnover in vivo not established
  10. 2011 High

    Defined a TNF→TNFR1→Nox/ROS axis driving MSX2 to inhibit osteogenesis and demonstrated Notch–BMP synergy at the MSX2 promoter via a Smad1–N1ICD complex.

    Evidence TNFR1/TNFR2 and p47phox knockouts, Nox inhibitors, H2O2 promoter assays, and Co-IP/ChIP at the Msx2 promoter

    PMID:20440096 PMID:21471203 PMID:22685265

    Open questions at the time
    • Downstream osteogenic targets repressed by ROS-induced MSX2 not enumerated
    • Cell-type specificity of the ROS input not generalized
  11. 2015 High

    Generalized MSX2 to stem-cell fate control as a direct repressor of SOX2 and activator of Nodal, establishing mutual MSX2–SOX2 antagonism at both transcript and protein levels.

    Evidence ChIP for direct promoter binding, gain/loss-of-function in hPSCs, and protein half-life measurements

    PMID:26427715

    Open questions at the time
    • Co-factors at the SOX2 and Nodal promoters not identified
    • Mechanism of SOX2-driven MSX2 protein destabilization unresolved
  12. 2021 High

    Connected MSX2 repression to chromatin remodeling control, showing it restrains cBAF/H3K27ac at differentiation loci, and identified cBAF as a physical interactor.

    Evidence ChIP-seq for MSX2/H3K27ac/cBAF, IP-MS for interactors, and gain/loss-of-function in human trophoblast stem cells

    PMID:34507999

    Open questions at the time
    • Whether MSX2 directly antagonizes cBAF recruitment or via intermediaries not resolved
    • Generality of cBAF antagonism beyond trophoblast loci untested
  13. 2023 Medium

    Established a second post-translational control layer: mitophagic degradation of MSX2 via CLU/AKT/DNM1L excludes it from the nucleus to derepress SOX2 and sustain cancer stemness.

    Evidence CLU gain/loss-of-function, nuclear/cytoplasmic fractionation, mitophagy assays, AKT inhibition and DNM1L phosphorylation assays in oral cancer stem cells

    PMID:36779631

    Open questions at the time
    • How nuclear MSX2 is routed to mitochondria for mitophagy mechanistically unclear
    • Single-lab finding awaiting independent confirmation

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how MSX2 dynamically selects between its repressor and activator outputs and between DNA-binding-dependent and -independent modes in response to distinct upstream signals at the level of a unified molecular mechanism.
  • No structural model of MSX2 bound to its co-repressors or to cBAF
  • Genome-wide direct target catalogs across lineages incomplete
  • Integration of transcriptional inputs with FBXW2/VRK2 and mitophagy degradation control not unified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 6 GO:0003677 DNA binding 4 GO:0140097 catalytic activity, acting on DNA 2
Localization
GO:0005634 nucleus 3 GO:0005829 cytosol 1
Pathway
R-HSA-162582 Signal Transduction 5 R-HSA-74160 Gene expression (Transcription) 5 R-HSA-1266738 Developmental Biology 4 R-HSA-392499 Metabolism of proteins 2 R-HSA-4839726 Chromatin organization 2
Partners
C/EBPALPHAFBXW2HDAC1MINT (SPEN)MYOCARDINRAP74 (GTF2F1)RUNX2TLE1
Complex memberships
cBAF (SWI/SNF)

Evidence

Reading pass · 41 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1993 A Pro148His substitution in the MSX2 homeodomain (position 7 of the N-terminal arm) causes autosomal dominant craniosynostosis (Boston type) by enhancing DNA binding affinity of the homeodomain, acting as a gain-of-function mutation. Human genetic mapping, segregation analysis, identification of missense mutation exclusively in affected individuals; mouse Msx2 transcript localization to calvarial sutures by in situ hybridization Cell High 8106171
1995 Both wild-type and P148H mutant Msx2 specifically bind a high-affinity homeodomain DNA target and repress reporter gene transcription in a dose-dependent but DNA binding site-independent manner, establishing MSX2 as a transcriptional repressor. Co-transfection reporter assays in cell lines, EMSA/DNA-binding assays with recombinant proteins Biochemical and biophysical research communications Medium 7726844
1996 MSX-1 and MSX-2 share a common consensus DNA binding site but MSX-2 has higher apparent DNA-binding affinity; both function as transcriptional repressors independently of their consensus DNA binding sites; the distinction in repressor potency maps to sequences N-terminal to the homeodomain. DNA binding site selection, EMSA, luciferase reporter transcription assays with deletion mutants Mechanisms of development Medium 8861098
1997 The core suppressor domain of Msx2 maps to residues 97–208, with residues 132–148 (upstream of and overlapping the homeodomain N-terminal extension) being essential; transcriptional suppression of the osteocalcin promoter does not require direct DNA binding by Msx2. Msx2 binds both subunits of TFIIF (RAP74 and RAP30) through its core suppressor domain; overexpression of RAP74 partially reverses Msx2-mediated suppression. Systematic N- and C-terminal and internal deletion mutagenesis with luciferase reporter assays in MC3T3-E1 osteoblasts; Far-Western blotting with recombinant proteins; co-transfection rescue experiments Biochemistry High 9265625
1997 Msx2 suppresses FGF2/cAMP-stimulated but not calcitriol-stimulated osteocalcin promoter activity; mechanistically, Msx2 inhibits binding of the OCFRE-binding protein (OCFREB) to the FGF2-response element (OCFRE) via its core suppressor residues 132–148, without itself binding the OCFRE. Luciferase reporter assays in MC3T3-E1 osteoblasts with FGF2/forskolin or calcitriol stimulation; EMSA with recombinant GST-Msx2 and nuclear extracts; protein purification of OCFREB; deletion-mutant analysis The Journal of biological chemistry High 9368026
1997 Miz1, a zinc-finger protein identified by yeast two-hybrid screen, directly interacts with Msx2 in vitro, enhances its DNA binding affinity for the osteocalcin promoter, and augments the effect of the P148H craniosynostosis mutation on DNA binding. Yeast two-hybrid screen, in vitro protein interaction assays, EMSA with Msx2 and Miz1, Northern blot and in situ hybridization for expression overlap Mechanisms of development Medium 9256341
1997 Constitutive ectopic Msx2 expression induces apoptosis in aggregated P19 cells, and BMP4 induces cell death via an Msx2-dependent pathway; BMP4 treatment of P19 cells induces Msx2 transcription, placing Msx2 downstream of BMP4 in programmed cell death. Stable transfection of P19 cells with Msx2 expression construct; BMP4 treatment; apoptosis quantification; Msx2 mRNA induction assay Developmental biology Medium 9205134
1999 MINT (Msx2-interacting nuclear target protein) was identified as a direct binding partner of Msx2; the interaction requires Msx2 residues necessary for transcriptional suppression; MINT's N-terminal RRM domain selectively binds the G/T-rich OCFRE in the osteocalcin promoter; MINT co-segregates with Msx2 in chromatin/nuclear matrix fractions. Far-Western expression cloning with radiolabeled GST-Msx2 fusion; Farwestern analysis; EMSA with MINT RRM domain; cellular fractionation and co-sedimentation with topoisomerase II; transient transfection reporter assays Biochemistry High 10451362
1999 Overexpression of Msx2 in primary chick calvarial osteoblasts prevents osteoblast differentiation and matrix mineralization; antisense Msx2 decreases proliferation and accelerates differentiation; Msx2 promoter activity is downregulated in differentiating osteoblastic nodules, indicating Msx2 maintains osteoprogenitors in a proliferative, undifferentiated state. Retroviral-mediated sense and antisense Msx2 overexpression in primary calvarial osteoblasts; mineralization and differentiation marker assays; Msx2 promoter-reporter analysis Developmental biology Medium 10328922
1999 Msx2 gene dosage controls the number of proliferative osteogenic cells in calvarial sutures; tissue-specific Msx2 overexpression in suture mesenchymal cells enhances parietal bone growth and increases BrdU-labeling of osteoblastic cells at the osteogenic front, consistent with a transient retardation of osteogenic cell differentiation. Transgenic mouse overexpression with Msx2-specific suture promoter; BrdU proliferation assay; histological analysis of cranial sutures Developmental biology Medium 9917362
2000 Heterozygous loss-of-function mutations in the MSX2 homeodomain (RK159-160del and R172H) cause >85% reduction in DNA binding and produce parietal foramina via haploinsufficiency, demonstrating that MSX2 dosage is critical for calvarial ossification; in contrast, the gain-of-function P148H mutation that enhances DNA binding causes craniosynostosis. Human genetic mutation identification; in vitro DNA binding assays with mutant Msx2 proteins; mouse phenotype analysis Nature genetics High 10742103
2000 Msx2-deficient mice have defective proliferation of osteoprogenitors at the calvarial osteogenic front; Msx2 is required for both chondrogenesis and osteogenesis in axial and appendicular skeleton, operating downstream of Pth/Pthrp receptor signaling; Msx1/Msx2 genetic dosage interactions modify PFM phenotype, indicating functional overlap. Msx2 null mutant mouse phenotyping; BrdU labeling; in situ hybridization for marker genes; Msx1/Msx2 compound mutant analysis Nature genetics High 10742104
2002 Pax3 represses Msx2 expression via a direct effect on a conserved Pax3-binding site in the Msx2 promoter; in Splotch (Pax3-null) mutant mice, upregulation of Msx2 causes the deficiency in cardiac neural crest development, establishing Msx2 as an immediate downstream effector of Pax3. Genetic epistasis (Splotch mutant × Msx2 mutant compound mice); molecular analysis of Pax3 binding to Msx2 promoter; in situ hybridization Development (Cambridge, England) High 11807043
2003 Msx2 promotes osteogenic and suppresses adipogenic differentiation of mesenchymal progenitors; osteogenic actions require intrinsic DNA binding (gain-of-function P148H enhances mineralization; DNA-binding-deficient T147A is inactive for osteogenesis); suppression of adipogenesis does not require DNA binding but occurs via protein-protein interactions with C/EBPalpha controlling PPARgamma transcription. Viral transduction of Msx2 variants (wild-type, P148H, T147A) in C3H10T1/2 and aortic myofibroblast cells; alkaline phosphatase assay; mineralized nodule quantification; adipogenesis assays; C/EBPalpha interaction studies The Journal of biological chemistry High 12925529
2003 Smad4 and Lef1 cooperatively activate the Msx2 promoter in response to BMP2; Wnt/beta-catenin signaling activates Msx2 via Lef1 binding and synergizes with BMP2; Wnt-dependent Msx2 activation requires Smad4 (not Smad1) even in the absence of BMP autocrine loops, demonstrated by chromatin immunoprecipitation showing Smad4 in the Lef1 transcriptional complex. Msx2 promoter-reporter assays; mutagenesis of Smad-binding elements and Lef1/TCF sites; Smad4-deficient ES cells; chromatin immunoprecipitation; co-transfection with Smad1, Smad4, and Lef1 The Journal of biological chemistry High 14551209
2004 Msx2 occupies the osteocalcin gene promoter in proliferating (undifferentiated) osteoblasts and represses it; after proliferation, Msx2 is replaced by Dlx3, Dlx5, and Runx2, forming a molecular switch for osteocalcin transcriptional activation during osteoblast differentiation. Chromatin immunoprecipitation (ChIP) across stages of osteoblast differentiation; RNA interference knockdown of Dlx3; overexpression experiments; RNA polymerase II ChIP Molecular and cellular biology High 15456894
2004 Msx2 suppresses BMP2-induced alkaline phosphatase (ALP) expression by competing with Dlx5 for the same cis-acting element in the ALP promoter; high Msx2 levels counteract Dlx5-stimulated ALP transcription until the Dlx5:Msx2 ratio exceeds a threshold. ALP promoter dissection with EMSA and site-directed mutagenesis; Msx2 overexpression in C2C12 and Runx2(-/-) cells; ALP mRNA and enzyme activity assays The Journal of biological chemistry High 15383550
2004 Msx2 colocalizes with Runx2/Osf2 and suppresses Runx2 transcriptional activity cooperatively with TLE1, recruiting HDAC1 activity to inhibit osteoblast differentiation in ligament fibroblasts; stable Msx2 knockdown in PDL-L2 cells induces osteoblastic differentiation and matrix mineralization. Co-immunoprecipitation; co-localization studies; RNA interference; stable overexpression in MC3T3-E1 cells; in situ hybridization; RT-PCR Molecular and cellular biology High 15060165
2004 MINT interacts functionally with Runx2 to enhance OCFRE-driven osteocalcin transcription; Msx2 abrogates Runx2-MINT activation by selectively inhibiting Runx2 binding to OC chromatin (demonstrated by ChIP); MINT adopts a reticular nuclear matrix distribution co-localizing with phospho-RNA polymerase II. Luciferase reporter assays in MC3T3E1 and CV1 cells; confocal immunofluorescence microscopy; chromatin immunoprecipitation; MINT RNA interference The Journal of biological chemistry High 15131132
2004 Necdin associates with Msx2 via MAGE-D1; a ternary complex of necdin, MAGE-D1, and Msx2 forms in vitro and is detected as an endogenous complex in differentiating embryonal carcinoma cells; co-expression of necdin and MAGE-D1 relieves Msx2-dependent transcriptional repression and rescues Msx2-inhibited myogenic differentiation in C2C12 cells. In vitro binding assays; co-immunoprecipitation; stable transfection of C2C12 with Msx2; rescue co-expression experiments; differentiation marker assays The Journal of biological chemistry High 15272023
2004 Msx2 inhibits transcriptional activity of PPARgamma, C/EBPbeta, and C/EBPdelta, blocking adipocyte differentiation induced by overexpression of each; and promotes osteoblast differentiation independently of Runx2 (active in Runx2-null cells). Overexpression and reporter assays in C3H10T1/2, C2C12, 3T3-F442A, and Runx2(-/-) mesenchymal cells; alkaline phosphatase and adipogenesis assays The Journal of biological chemistry Medium 15175325
2005 Msx2-expressing cells secrete paracrine Wnt signals (upregulating Wnt3a and Wnt7a, downregulating Dkk1) that promote osteogenic and suppress adipogenic differentiation; Msx2 induces nuclear beta-catenin accumulation and TCF/LEF transcriptional activity; Dkk1 treatment reverses these effects; in vivo, TOPGAL reporter mice confirm augmented Wnt signaling in Msx2-transgenic aorta. Conditioned media transfer experiments; TCF/LEF reporter (TOPGAL) transgenic mice; immunofluorescence for nuclear beta-catenin; alkaline phosphatase assay; Dkk1 rescue; qRT-PCR for Wnt ligands The Journal of clinical investigation High 15841209
2006 Msx1 and Msx2 form a ternary complex with SRF and myocardin, inhibiting SRF/myocardin binding to the CArG-box motif and suppressing smooth muscle cell marker gene (SM22alpha, caldesmon) transcription; this interaction is induced downstream of BMP2/4/6 signaling. Co-immunoprecipitation; gel-shift (EMSA); chromatin immunoprecipitation; promoter-reporter assays; BMP treatment of VSMCs Molecular and cellular biology High 17030628
2007 Msx2 is a direct transcriptional target of Notch/RBP-Jk signaling; Notch1 intracellular domain (N1-ICD) induces Msx2 gene expression via an RBP-Jk-binding element within the Msx2 promoter; RBP-Jk-deficient cells fail to induce Msx2 in response to N1-ICD; Msx2 mediates N1-ICD-induced ALP activity and vascular smooth muscle cell mineralization. Msx2 promoter deletion and site-directed mutagenesis; RBP-Jk-deficient fibroblasts; siRNA knockdown of Msx2 and RBP-Jk; ALP activity assay; immunohistochemistry of human calcifying plaques Arteriosclerosis, thrombosis, and vascular biology High 19407244
2007 Vitamin K2 (MK4) activates Msx2 gene transcription through PXR binding to a PXRE in the Msx2 promoter; ChIP shows PXR and p300 coactivator recruitment to this element; MK4-bound PXR cooperates with estrogen-bound ERalpha on the Msx2 promoter; knockdown of PXR or Msx2 attenuates MK4-induced osteoblast differentiation. 2D-SDS-PAGE proteomics; Msx2 promoter reporter mapping; ChIP for PXR and p300; siRNA knockdown; co-transfection of PXR/RXRalpha/ERalpha Molecular and cellular biology High 17875939
2008 BMP2 regulates Osterix via two parallel pathways: a Runx2-dependent pathway and a Runx2-independent pathway through Msx2; Msx2 (induced by BMP2 in Runx2-null cells via Smad1/Smad4) induces Osterix expression; Msx2 knockdown inhibits BMP2-induced Osterix in Runx2-null cells. Runx2-deficient mesenchymal cells; Msx2 overexpression and siRNA knockdown; BMP2 treatment; Osterix and ALP expression assays; Smad overexpression/inhibitory Smad experiments The Journal of biological chemistry High 18703512
2008 Msx2 exerts bone anabolic effects by reducing Dkk1 expression and enhancing canonical Wnt (Wnt7a, Wnt7b) signaling; Msx2 inhibits Dkk1 promoter activity and reduces RNA polymerase II association with Dkk1 chromatin; RNAi knockdown of Wnt7a, Wnt7b, and LRP6 significantly reduces Msx2-induced alkaline phosphatase; confirmed in Msx2-transgenic mice using TOPGAL reporter. CMV-Msx2 transgenic mice; microCT; histomorphometry; TOPGAL Wnt reporter; Msx2 siRNA; ChIP for RNA Pol II at Dkk1; Wnt7a/7b/LRP6 knockdown with rescue assay The Journal of biological chemistry High 18487199
2008 Msx1 and Msx2 bind cardiac T-box proteins Tbx2, Tbx3, and Tbx5 via their homeodomain and T-box domains; Msx proteins together with Tbx2/Tbx3 suppress Connexin43 (Cx43) promoter activity; Msx1 binds the Cx43 promoter at a conserved site adjacent to a T-box site (by ChIP), and Msx activity on the Cx43 promoter depends on the presence of Tbx3. Yeast two-hybrid screen; in vitro pull-down; reporter assays in rat heart-derived cells; chromatin immunoprecipitation Cardiovascular research High 18285513
2008 Msx2 promotes chondrocyte maturation in part by upregulating Ihh (Indian hedgehog) expression; cyclopamine (hedgehog pathway inhibitor) blocks Msx2-induced chondrogenesis; Msx2's chondrogenic action requires BMP2/Smad signaling (Smad1/4 enhance, Smad6 blocks). Overexpression of constitutively active Msx2 in primary chondrocytes and metatarsal explants; cyclopamine treatment; Smad overexpression/inhibition; ALP and collagen X expression assays; Msx2 siRNA knockdown The Journal of biological chemistry Medium 18682398
2008 The Boston craniosynostosis P148H mutation renders MSX2 more susceptible to ubiquitin-dependent proteasomal degradation; Praja1 E3 ubiquitin ligase mediates MSX2 degradation; P148H shows greater ubiquitylation and shorter protein half-life than wild-type; P148H functions as a dominant-negative by increasing ubiquitylation of wild-type MSX2. Pulse-chase protein half-life experiments; ubiquitylation assays; Praja1 co-expression; dominant-negative co-expression; osteoblast proliferation and cyclin D1 assays The Journal of biological chemistry High 18786927
2008 BMP4-induced EMT in pancreatic cancer cells requires MSX2; BMP4 induces MSX2 via ERK, p38 MAPK, and Smad pathways; siRNA-mediated MSX2 knockdown abolishes BMP4-induced E-cadherin repression, vimentin induction, and enhanced cell migration. BMP4 treatment of Panc-1 cells; MSX2 siRNA knockdown; pathway inhibitors (ERK, p38, Smad); migration assays; Western blot for EMT markers Journal of cellular physiology Medium 17516553
2010 Msx2 mediates TNF-alpha inhibition of BMP2-induced osteoblast differentiation; TNF-alpha induces Msx2 via NF-kappaB pathway (not JNK); Msx2 siRNA rescues ALP expression suppressed by TNF-alpha, placing Msx2 as a downstream effector of TNF-alpha/NF-kappaB in inhibiting osteogenesis. TNF-alpha treatment of C2C12 and Runx2(-/-) calvarial cells; pathway-specific inhibitors for NF-kappaB and JNK; Msx2 siRNA knockdown; ALP expression assays Experimental & molecular medicine Medium 20440096
2011 TNF acts through TNFR1 (not TNFR2) to upregulate Msx2 via reactive oxygen species (ROS) generated by NADPH oxidase (Nox); hydrogen peroxide directly upregulates Msx2 mRNA and promoter activity; Nox inhibition, p47phox genetic deficiency, and rotenone reduce TNF-induced Msx2; TNFR1-null aortic myofibroblasts express ~5% of wild-type Msx2 and are non-inducible by TNF. TNFR1-/- and TNFR2-/- cells; p47phox-/- cells; Nox inhibitors; H2O2 treatment; Msx2 promoter reporter; antisense oligonucleotides in SM22-TNF transgenic mice Endocrinology High 22685265
2011 Notch signaling (N1-ICD/RBP-Jk) enhances BMP2-responsiveness of the Msx2 promoter; Smad1 interacts with N1-ICD to form a complex within the Msx2 promoter; RBP-Jk binding element is required for this synergistic BMP2 + Notch induction of Msx2 gene expression and subsequent ALP activity/mineralization in smooth muscle cells. Msx2 promoter deletion/mutation analysis; RBP-Jk-deficient cells; siRNA for RBP-Jk; co-immunoprecipitation of Smad1 and N1-ICD; ChIP at Msx2 promoter The Journal of biological chemistry High 21471203
2012 MSX2 directly regulates ABCG2 transcription in functional cooperation with SP1 via SP1-binding elements within the ABCG2 promoter; MSX2 overexpression or siRNA knockdown proportionally changes ABCG2 expression, and MSX2 expression correlates with chemoresistance. ABCG2 promoter reporter assay with MSX2 and SP1 co-expression; siRNA knockdown; overexpression; correlation of MSX2 and ABCG2 mRNA levels across cell lines Journal of cellular physiology Medium 21465479
2012 FOXC1 directly occupies a conserved element in the MSX2 promoter (by ChIP) and transcriptionally activates both human and mouse MSX2 promoters; FOXC1 siRNA reduces endogenous MSX2 expression; heterologous Foxc1 expression in C2C12 cells elevates ALP activity and Runx2 and Msx2 levels, placing FOXC1 upstream of MSX2 in early osteoblast differentiation. ChIP for FOXC1 at MSX2 promoter; luciferase reporter assays; siRNA; Foxc1 overexpression in C2C12 PloS one Medium 23145080
2013 Msx2 and Wnt7b signaling maintain aortic endothelial cell (EC) phenotype and oppose endothelial-mesenchymal transition (EndoMT); EC-specific deletion of Wnt7b upregulates osteogenic genes including Msx2 and nuclear phospho-Smad1/5; Msx2 in ECs has the opposite effect to mesenchymal cells, preserving EC identity. Cdh5-Cre;Wnt7b(fl/fl);LDLR(-/-) conditional knockout mice; adenoviral transduction of aortic ECs; immunofluorescence; Western blot; calcium/collagen quantification Arteriosclerosis, thrombosis, and vascular biology Medium 23685555
2015 MSX2 is a direct BMP pathway target in human pluripotent stem cells (hPSCs); MSX2 destabilizes pluripotency by directly binding the SOX2 promoter and repressing SOX2 transcription; simultaneously MSX2 directly activates Nodal promoter to commit cells to mesendoderm; SOX2 can promote MSX2 protein degradation, establishing mutual antagonism; Wnt signals via LEF1 synergistically activate MSX2 during mesendoderm induction. MSX2 overexpression and shRNA depletion in hPSCs; ChIP for MSX2 binding to SOX2 and Nodal promoters; promoter reporter assays; protein half-life measurement; MSX2/SOX2 co-expression Cell research High 26427715
2019 MSX2 is a substrate of FBXW2 E3 ubiquitin ligase; FBXW2 binds MSX2, promotes its ubiquitylation and proteasomal degradation, and shortens its protein half-life; hypoxia induces VRK2 kinase to facilitate MSX2-FBXW2 binding and enhance FBXW2-mediated MSX2 degradation; MSX2 accumulation (upon FBXW2 inactivation) represses SOX2 transcription. Co-immunoprecipitation; ubiquitylation assays; FBXW2 overexpression and siRNA knockdown with pulse-chase; VRK2 kinase assays; in vitro and in vivo breast cancer models Proceedings of the National Academy of Sciences of the United States of America High 31548378
2021 MSX2 represses the syncytiotrophoblast transcriptional program in human trophoblast stem cells; MSX2 directly binds and regulates differentiation genes; MSX2 interacts with the SWI/SNF canonical BAF (cBAF) subcomplex and co-occupies target genes together with H3K27ac; MSX2 depletion increases H3K27ac and cBAF occupancy at differentiation genes, indicating MSX2 prevents chromatin remodeling at syncytiotrophoblast loci. MSX2 shRNA depletion and overexpression in human trophoblast stem cells; ChIP-seq for MSX2, H3K27ac, and cBAF components; immunoprecipitation-mass spectrometry for MSX2 interactors; transcriptomics Proceedings of the National Academy of Sciences of the United States of America High 34507999
2023 CLU-mediated mitophagy promotes degradation of MSX2 in mitochondria/cytoplasm, preventing its nuclear translocation; when MSX2 is degraded by mitophagy, SOX2 expression is derepressed, maintaining cancer stemness; CLU activates AKT, which phosphorylates DNM1L/Drp1 at Ser616 to initiate mitochondrial fission preceding mitophagy of MSX2. CLU gain/loss-of-function in oral CSCs; MSX2 nuclear vs. cytoplasmic fractionation; mitophagy assays; AKT inhibition; DNM1L phosphorylation assays; SOX2 reporter/expression assays Autophagy Medium 36779631

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2000 Msx2 deficiency in mice causes pleiotropic defects in bone growth and ectodermal organ formation. Nature genetics 598 10742104
1993 A mutation in the homeodomain of the human MSX2 gene in a family affected with autosomal dominant craniosynostosis. Cell 563 8106171
2008 BMP2 regulates Osterix through Msx2 and Runx2 during osteoblast differentiation. The Journal of biological chemistry 427 18703512
2005 Msx2 promotes cardiovascular calcification by activating paracrine Wnt signals. The Journal of clinical investigation 365 15841209
2003 MSX2 promotes osteogenesis and suppresses adipogenic differentiation of multipotent mesenchymal progenitors. The Journal of biological chemistry 282 12925529
2007 Aortic Msx2-Wnt calcification cascade is regulated by TNF-alpha-dependent signals in diabetic Ldlr-/- mice. Arteriosclerosis, thrombosis, and vascular biology 268 17932314
1992 Expression patterns of the homeobox gene, Hox-8, in the mouse embryo suggest a role in specifying tooth initiation and shape. Development (Cambridge, England) 268 1358591
2004 Dlx3 transcriptional regulation of osteoblast differentiation: temporal recruitment of Msx2, Dlx3, and Dlx5 homeodomain proteins to chromatin of the osteocalcin gene. Molecular and cellular biology 230 15456894
2000 Functional haploinsufficiency of the human homeobox gene MSX2 causes defects in skull ossification. Nature genetics 229 10742103
2003 Smad4 and beta-catenin co-activators functionally interact with lymphoid-enhancing factor to regulate graded expression of Msx2. The Journal of biological chemistry 173 14551209
2003 Osteoblast-related transcription factors Runx2 (Cbfa1/AML3) and MSX2 mediate the expression of bone sialoprotein in human metastatic breast cancer cells. Cancer research 165 12750290
2004 Bone morphogenetic protein-2-induced alkaline phosphatase expression is stimulated by Dlx5 and repressed by Msx2. The Journal of biological chemistry 163 15383550
1999 Msx2 gene dosage influences the number of proliferative osteogenic cells in growth centers of the developing murine skull: a possible mechanism for MSX2-mediated craniosynostosis in humans. Developmental biology 157 9917362
1996 Regulation of Msx-1, Msx-2, Bmp-2 and Bmp-4 during foetal and postnatal mammary gland development. Development (Cambridge, England) 155 8787747
2005 Combined deficiencies of Msx1 and Msx2 cause impaired patterning and survival of the cranial neural crest. Development (Cambridge, England) 154 16221730
2003 Msx2 and Twist cooperatively control the development of the neural crest-derived skeletogenic mesenchyme of the murine skull vault. Development (Cambridge, England) 153 14597577
2004 Reciprocal roles of MSX2 in regulation of osteoblast and adipocyte differentiation. The Journal of biological chemistry 152 15175325
1993 Expression of homeobox genes Msx-1 (Hox-7) and Msx-2 (Hox-8) during cardiac development in the chick. Developmental dynamics : an official publication of the American Association of Anatomists 143 8106021
1997 Msx2 is a transcriptional regulator in the BMP4-mediated programmed cell death pathway. Developmental biology 131 9205134
2003 'Cyclic alopecia' in Msx2 mutants: defects in hair cycling and hair shaft differentiation. Development (Cambridge, England) 123 12466204
2007 Vascular Bmp Msx2 Wnt signaling and oxidative stress in arterial calcification. Annals of the New York Academy of Sciences 119 18056036
1996 Comparison of MSX-1 and MSX-2 suggests a molecular basis for functional redundancy. Mechanisms of development 117 8861098
2005 Analysis of Msx1; Msx2 double mutants reveals multiple roles for Msx genes in limb development. Development (Cambridge, England) 114 15930102
2007 Concerted action of Msx1 and Msx2 in regulating cranial neural crest cell differentiation during frontal bone development. Mechanisms of development 110 17693062
1999 Ectopic Msx2 overexpression inhibits and Msx2 antisense stimulates calvarial osteoblast differentiation. Developmental biology 100 10328922
1998 Ectopic expression of Msx-2 in posterior limb bud mesoderm impairs limb morphogenesis while inducing BMP-4 expression, inhibiting cell proliferation, and promoting apoptosis. Developmental biology 100 9578615
1998 Expression of Msx-2 during development, regeneration, and wound healing in axolotl limbs. The Journal of experimental zoology 100 9846383
2013 Dkk1 and MSX2-Wnt7b signaling reciprocally regulate the endothelial-mesenchymal transition in aortic endothelial cells. Arteriosclerosis, thrombosis, and vascular biology 97 23685555
1997 Perinatal lethality and multiple craniofacial malformations in MSX2 transgenic mice. Human molecular genetics 96 9147639
2000 Positionally-dependent chondrogenesis induced by BMP4 is co-regulated by Sox9 and Msx2. Developmental dynamics : an official publication of the American Association of Anatomists 91 10767084
2019 The FBXW2-MSX2-SOX2 axis regulates stem cell property and drug resistance of cancer cells. Proceedings of the National Academy of Sciences of the United States of America 89 31548378
2004 Homeobox protein MSX2 acts as a molecular defense mechanism for preventing ossification in ligament fibroblasts. Molecular and cellular biology 88 15060165
1997 Miz1, a novel zinc finger transcription factor that interacts with Msx2 and enhances its affinity for DNA. Mechanisms of development 87 9256341
2007 Bone morphogenetic protein 4 induces epithelial-mesenchymal transition through MSX2 induction on pancreatic cancer cell line. Journal of cellular physiology 84 17516553
1999 The RRM domain of MINT, a novel Msx2 binding protein, recognizes and regulates the rat osteocalcin promoter. Biochemistry 83 10451362
2008 Msx2 exerts bone anabolism via canonical Wnt signaling. The Journal of biological chemistry 81 18487199
1995 Gene defect in hypodontia: exclusion of MSX1 and MSX2 as candidate genes. Human genetics 81 7649547
2002 Msx2 is an immediate downstream effector of Pax3 in the development of the murine cardiac neural crest. Development (Cambridge, England) 79 11807043
2007 Msx1 and Msx2 regulate survival of secondary heart field precursors and post-migratory proliferation of cardiac neural crest in the outflow tract. Developmental biology 77 17601530
2006 Bone morphogenetic protein-induced MSX1 and MSX2 inhibit myocardin-dependent smooth muscle gene transcription. Molecular and cellular biology 75 17030628
2004 Msx2 and necdin combined activities are required for smooth muscle differentiation in mesoangioblast stem cells. Circulation research 75 15155529
2001 Msx2 is a repressor of chondrogenic differentiation in migratory cranial neural crest cells. Developmental dynamics : an official publication of the American Association of Anatomists 75 11668602
2014 BMP4 promotes EMT and mesodermal commitment in human embryonic stem cells via SLUG and MSX2. Stem cells (Dayton, Ohio) 72 24549638
2007 Msx2 -/- transgenic mice develop compound amelogenesis imperfecta, dentinogenesis imperfecta and periodental osteopetrosis. Bone 72 17878071
2008 Msx1 and Msx2 are required for endothelial-mesenchymal transformation of the atrioventricular cushions and patterning of the atrioventricular myocardium. BMC developmental biology 70 18667074
2007 Vitamin K induces osteoblast differentiation through pregnane X receptor-mediated transcriptional control of the Msx2 gene. Molecular and cellular biology 70 17875939
2008 Msx1 and Msx2 are functional interacting partners of T-box factors in the regulation of Connexin43. Cardiovascular research 69 18285513
1999 PDX-1 and Msx-2 expression in the regenerating and developing pancreas. The Journal of endocrinology 69 10588826
2011 Notch signaling pathway enhances bone morphogenetic protein 2 (BMP2) responsiveness of Msx2 gene to induce osteogenic differentiation and mineralization of vascular smooth muscle cells. The Journal of biological chemistry 68 21471203
1997 Structure-function analysis of Msx2-mediated transcriptional suppression. Biochemistry 68 9265625
2004 Necdin interacts with the Msx2 homeodomain protein via MAGE-D1 to promote myogenic differentiation of C2C12 cells. The Journal of biological chemistry 67 15272023
1994 Regulation of the Msx2 homeobox gene during mouse embryogenesis: a transgene with 439 bp of 5' flanking sequence is expressed exclusively in the apical ectodermal ridge of the developing limb. Mechanisms of development 67 7893602
2014 Targeted reduction of vascular Msx1 and Msx2 mitigates arteriosclerotic calcification and aortic stiffness in LDLR-deficient mice fed diabetogenic diets. Diabetes 66 25056439
2008 Up-regulation of MSX2 enhances the malignant phenotype and is associated with twist 1 expression in human pancreatic cancer cells. The American journal of pathology 65 18349132
2015 MSX2 mediates entry of human pluripotent stem cells into mesendoderm by simultaneously suppressing SOX2 and activating NODAL signaling. Cell research 64 26427715
2009 Notch signaling induces osteogenic differentiation and mineralization of vascular smooth muscle cells: role of Msx2 gene induction via Notch-RBP-Jk signaling. Arteriosclerosis, thrombosis, and vascular biology 62 19407244
2001 Antagonistic regulation of Dlx2 expression by PITX2 and Msx2: implications for tooth development. Gene expression 62 11763998
2018 MSX2 Initiates and Accelerates Mesenchymal Stem/Stromal Cell Specification of hPSCs by Regulating TWIST1 and PRAME. Stem cell reports 61 30033084
2008 Genetic interplays between Msx2 and Foxn1 are required for Notch1 expression and hair shaft differentiation. Developmental biology 61 19103190
2011 Msx1 and Msx2 promote meiosis initiation. Development (Cambridge, England) 58 22071108
2004 Msx2 controls ameloblast terminal differentiation. Developmental dynamics : an official publication of the American Association of Anatomists 54 15499554
1996 Regulated expression of homeobox genes Msx-1 and Msx-2 in mouse mammary gland development suggests a role in hormone action and epithelial-stromal interactions. Developmental biology 52 8660900
2010 Msx2 mediates the inhibitory action of TNF-alpha on osteoblast differentiation. Experimental & molecular medicine 51 20440096
2012 TNFR1-activated reactive oxidative species signals up-regulate osteogenic Msx2 programs in aortic myofibroblasts. Endocrinology 50 22685265
2004 MINT, the Msx2 interacting nuclear matrix target, enhances Runx2-dependent activation of the osteocalcin fibroblast growth factor response element. The Journal of biological chemistry 50 15131132
2003 Fibroblast growth factors lead to increased Msx2 expression and fusion in calvarial sutures. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 48 12674336
2021 MSX2 safeguards syncytiotrophoblast fate of human trophoblast stem cells. Proceedings of the National Academy of Sciences of the United States of America 46 34507999
2011 Transcriptional repression in the Notch pathway: thermodynamic characterization of CSL-MINT (Msx2-interacting nuclear target protein) complexes. The Journal of biological chemistry 46 21372128
2003 Microphthalmia resulting from MSX2-induced apoptosis in the optic vesicle. Investigative ophthalmology & visual science 46 12766037
2004 Msx-1 and Msx-2 in mammary gland development. Journal of mammary gland biology and neoplasia 45 15300013
1999 Expression of osteocalcin and its transcriptional regulators core-binding factor alpha 1 and MSX2 in osteoid-forming tumours. Journal of orthopaedic research : official publication of the Orthopaedic Research Society 43 10569470
1997 Stimulus-selective inhibition of rat osteocalcin promoter induction and protein-DNA interactions by the homeodomain repressor Msx2. The Journal of biological chemistry 43 9368026
2011 Msx1 and Msx2 in limb mesenchyme modulate digit number and identity. Developmental dynamics : an official publication of the American Association of Anatomists 40 21465616
2004 Alx4 and Msx2 play phenotypically similar and additive roles in skull vault differentiation. Journal of anatomy 37 15198690
1997 Gestational exposure to ethanol suppresses msx2 expression in developing mouse embryos. Proceedings of the National Academy of Sciences of the United States of America 37 9207129
2000 Craniofacial disorders caused by mutations in homeobox genes MSX1 and MSX2. Journal of craniofacial genetics and developmental biology 36 10879654
2012 The homeobox gene MSX2 determines chemosensitivity of pancreatic cancer cells via the regulation of transporter gene ABCG2. Journal of cellular physiology 35 21465479
2010 Enamel protein regulation and dental and periodontal physiopathology in MSX2 mutant mice. The American journal of pathology 35 20934968
2009 Msx2 induces epithelial-mesenchymal transition in mouse mammary epithelial cells through upregulation of Cripto-1. Journal of cellular physiology 35 19170109
2000 The homeobox genes MSX2 and MOX2 are candidates for regulating epithelial-mesenchymal cell interactions in the human placenta. Placenta 34 10831122
2008 MSX2 stimulates chondrocyte maturation by controlling Ihh expression. The Journal of biological chemistry 33 18682398
2005 Ventral abdominal wall dysmorphogenesis of Msx1/Msx2 double-mutant mice. The anatomical record. Part A, Discoveries in molecular, cellular, and evolutionary biology 33 15803476
2012 Initiation of early osteoblast differentiation events through the direct transcriptional regulation of Msx2 by FOXC1. PloS one 32 23145080
2006 MSX2 promotes vaginal epithelial differentiation and wolffian duct regression and dampens the vaginal response to diethylstilbestrol. Molecular endocrinology (Baltimore, Md.) 32 16513791
2023 CLU (clusterin) promotes mitophagic degradation of MSX2 through an AKT-DNM1L/Drp1 axis to maintain SOX2-mediated stemness in oral cancer stem cells. Autophagy 30 36779631
2015 MSX2 in ameloblast cell fate and activity. Frontiers in physiology 28 25601840
2013 BMP-mediated functional cooperation between Dlx5;Dlx6 and Msx1;Msx2 during mammalian limb development. PloS one 28 23382810
2007 Syndromic craniosynostosis due to complex chromosome 5 rearrangement and MSX2 gene triplication. American journal of medical genetics. Part A 28 18000908
2005 Structural analysis of the group II intron splicing factor CRS2 yields insights into its protein and RNA interaction surfaces. Journal of molecular biology 28 15567410
2006 Functional interactions between Dlx2 and lymphoid enhancer factor regulate Msx2. Nucleic acids research 26 17068080
2005 Formation of the CRS2-CAF2 group II intron splicing complex is mediated by a 22-amino acid motif in the COOH-terminal region of CAF2. The Journal of biological chemistry 26 16379013
1995 DNA binding and transcriptional properties of wild-type and mutant forms of the homeodomain protein Msx2. Biochemical and biophysical research communications 26 7726844
2016 Msx1 and Msx2 function together in the regulation of primordial germ cell migration in the mouse. Developmental biology 25 27435625
2011 MSX2 is an oncogenic downstream target of activated WNT signaling in ovarian endometrioid adenocarcinoma. Oncogene 25 21499300
2008 The Boston-type craniosynostosis mutation MSX2 (P148H) results in enhanced susceptibility of MSX2 to ubiquitin-dependent degradation. The Journal of biological chemistry 25 18786927
2002 Msx2 and p21(CIP1/WAF1) mediate the proapoptotic effects of bone morphogenetic protein-4 on ventricular zone progenitor cells. Journal of neuroscience research 25 12205674
2006 Msx1 and Msx2 have shared essential functions in neural crest but may be dispensable in epidermis and axis formation in Xenopus. The International journal of developmental biology 24 16586351
2005 Agonistic and antagonistic action of AP2, Msx2, Pax6, Prox1 AND Six3 in the regulation of Sox2 expression. Ophthalmic research 24 16118513
2002 Tgf-beta1, Tgf-beta2, Tgf-beta3 and Msx2 expression is elevated during frontonasal suture morphogenesis and during active postnatal facial growth. Orthodontics & craniofacial research 24 12416537
2014 Msx1 and Msx2 act as essential activators of Atoh1 expression in the murine spinal cord. Development (Cambridge, England) 23 24715462

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