Affinage

EMX1

Homeobox protein EMX1 · UniProt Q04741

Length
290 aa
Mass
31.3 kDa
Annotated
2026-06-09
45 papers in source corpus 21 papers cited in narrative 21 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

EMX1 is a homeodomain transcription factor that specifies dorsal telencephalic (archipallial) identity and guides the wiring of the developing cerebral cortex, acting in the nuclei of cortical glutamatergic/pyramidal neurons across all layers and, distinctively, also localizing to olfactory sensory neuron axons (PMID:8843394, PMID:11709490). During forebrain development it functions in both proliferating ventricular-zone progenitors and postmitotic neurons (PMID:8743751), and its lineage gives rise to radial glia, Cajal-Retzius cells, glutamatergic neurons, astrocytes, and oligodendrocytes but not most GABAergic neurons (PMID:12151506). EMX1 acts largely redundantly with its paralog EMX2: single mutants show subtle defects, whereas Emx1;Emx2 double mutants display severe cortical hypoplasia, loss of hippocampus and Cajal-Retzius/subplate neurons, failure of cortical efferent axons to exit the cortex with non-cell-autonomous thalamocortical pathfinding defects, transformation of archipallium into roof plate, and loss of the cortical hem and choroid plexus, establishing a cooperative requirement for archipallial identity over roof plate/choroid plexus fate (PMID:9006071, PMID:12091317, PMID:12561075, PMID:15147765, PMID:16793035). Independently of EMX2, EMX1 is required for corpus callosum formation: it directly activates Nrp1 transcription in anterior cingulate axons, and NRP1 overexpression rescues the midline-crossing defect of Emx1 knockouts (PMID:8812118, PMID:26534986). EMX1 also supports adult hippocampal neurogenesis through a VEGF–Cofilin1 phosphorylation pathway in neural stem cells (PMID:17490651, PMID:27799894), and contributes to neocortical area patterning (PMID:26901526). EMX1 physically interacts with WDR11, a partnership relevant to olfactory neuron development (PMID:20887964). In non-neural and tumor contexts, EMX1 acts as a transcriptional repressor and tumor suppressor, repressing stem-cell regulatory genes and the Wnt/β-catenin pathway—directly binding the WASF2 promoter to suppress it (PMID:34016958, PMID:34364391, PMID:35849030)—while in hepatocellular carcinoma the full-length isoform binds the EGFR promoter to activate EGFR-ERK signaling and drive invasion (PMID:38007497).

Mechanistic history

Synthesis pass · year-by-year structured walk · 17 steps
  1. 1996 Medium

    Establishing whether EMX1 has a non-redundant developmental role, knockout mice revealed a specific requirement in interhemispheric connectivity, defining the corpus callosum as a key EMX1-dependent structure.

    Evidence N-terminal deletion knockout mouse with histological analysis

    PMID:8812118

    Open questions at the time
    • Cortical histology appeared normal, leaving the molecular mechanism of the callosal defect unexplained
    • Single lab; later shown to be background-dependent
  2. 1996 Medium

    To place EMX1 functionally, protein and transcript localization showed nuclear (transcription-factor-consistent) expression across all cortical layers in proliferating and postmitotic cells, plus an unusual axonal pool in olfactory sensory neurons.

    Evidence Immunohistochemistry and in situ hybridization across developmental stages

    PMID:8743751 PMID:8843394

    Open questions at the time
    • Function of the axonal/olfactory EMX1 pool not defined
    • No direct transcriptional targets identified at this stage
  3. 1997 High

    Addressing why single mutants are mild, double-mutant epistasis demonstrated functional redundancy and cooperation between EMX1 and EMX2 in dorsal forebrain patterning.

    Evidence Emx1, Emx2, and Emx1;Emx2 double knockout comparative phenotyping

    PMID:9006071

    Open questions at the time
    • Shared versus distinct target genes of the two paralogs not resolved
    • Mechanism of compensation unknown
  4. 2001 High

    To define the cell-type scope of EMX1, multiple orthogonal methods established that expression is restricted to glutamatergic pyramidal neurons, and fate mapping delineated the full Emx1 lineage as pallial excitatory/glial but not GABAergic.

    Evidence ISH, EM-level IHC, glutamate double-labeling, chimeric clonal analysis, and IRES-Cre genetic fate mapping

    PMID:11709490 PMID:12151506

    Open questions at the time
    • Does not establish cell-autonomous transcriptional targets within these cell types
  5. 2002 High

    Comprehensive double-mutant analyses defined the cooperative roles of EMX1/EMX2 in cortical size, lamination, Cajal-Retzius/subplate neuron production, tangential migration, and axon scaffolding, showing several effects are non-cell-autonomous.

    Evidence Emx1/Emx2 double knockout with BrdU labeling, cell transplantation, axon tracing, and marker analysis

    PMID:12091317 PMID:12561075

    Open questions at the time
    • Downstream effector genes mediating the environmental/non-cell-autonomous signal not identified
    • Individual contribution of EMX1 versus EMX2 not separable
  6. 2002 Medium

    Testing whether EMX1 patterns cortical areas, single and double mutants showed no arealization change at perinatal stages, initially excluding EMX1 from area specification.

    Evidence Knockout with seven area markers, axon tracing, and epistasis analysis

    PMID:12196586

    Open questions at the time
    • Examined only at perinatal stages; later contradicted by mature-age analysis (#14)
  7. 2004 Medium

    Probing the fate decision underlying archipallial development, double mutants showed transformation of archipallium into roof plate with loss of cortical hem and choroid plexus, establishing EMX1/EMX2 as enforcers of archipallial identity.

    Evidence Emx1;Emx2 double knockout with marker IHC/ISH

    PMID:15147765

    Open questions at the time
    • Direct target genes mediating fate suppression unknown
    • Single lab
  8. 2006 Medium

    A gain-of-function test in chick midline showed that Emx1 misexpression instructs neuroepithelial over choroid plexus identity, demonstrating EMX1 actively suppresses choroid plexus fate.

    Evidence In ovo misexpression with fate-marker in situ hybridization

    PMID:16793035

    Open questions at the time
    • Direct transcriptional targets among Gli3/Lhx2/Otx2/BMP7 not established
    • Avian system; single lab
  9. 2007 Medium

    Asking whether EMX1 acts beyond embryogenesis, knockouts revealed a selective requirement for adult dentate gyrus neurogenesis with behavioral consequences.

    Evidence Knockout mouse with stereology, BrdU/doublecortin labeling, and behavioral testing

    PMID:17490651

    Open questions at the time
    • Molecular pathway linking EMX1 to neurogenesis not defined here
    • Single lab
  10. 2010 Medium

    To identify EMX1 protein partners, a Co-IP screen tied to a disease translocation showed EMX1 binds WDR11, with disease mutations disrupting the interaction, linking EMX1 to olfactory development and puberty.

    Evidence Co-immunoprecipitation, translocation breakpoint mapping, and missense-mutation binding analysis

    PMID:20887964

    Open questions at the time
    • Functional consequence of the EMX1-WDR11 complex on transcription not shown
    • No reciprocal/structural validation
  11. 2015 High

    Resolving the mechanism of the callosal phenotype, EMX1 was shown to directly activate Nrp1 in cingulate axons, with NRP1 overexpression rescuing the midline-crossing defect.

    Evidence Knockout mouse, axon tracing, in vitro transcriptional activation, and NRP1 rescue

    PMID:26534986

    Open questions at the time
    • Whether EMX1 binds the Nrp1 promoter directly in vivo not shown
    • Other callosal target genes unexplored
  12. 2016 Medium

    Revisiting arealization at mature ages with two independent lines reversed the earlier negative result, establishing EMX1 as a regulator of neocortical area boundaries.

    Evidence Two independent knockout lines with area marker analysis at mature ages

    PMID:26901526

    Open questions at the time
    • Direct contradiction with perinatal analysis (#6) unresolved at the mechanistic level
    • Target genes mediating area shifts unidentified
  13. 2016 Medium

    To explain the neurogenesis defect mechanistically, proteomics in Emx1 KO neural stem cells implicated impaired VEGF-induced Cofilin1 phosphorylation and reduced FLK expression in defective self-renewal and migration.

    Evidence Neurosphere and migration assays, 2D-PAGE/MS proteomics, and p-Cofilin1 western blot in KO NSCs

    PMID:27799894

    Open questions at the time
    • Whether EMX1 transcriptionally controls VEGF/FLK directly not established
    • Single lab
  14. 2018 Medium

    An ortholog study extended EMX1 function beyond the CNS, showing zebrafish emx1 directs distal nephron segment fate downstream of mecom/tbx2b and upstream of irx and sim1a genes.

    Evidence Zebrafish loss-of-function, genetic epistasis, segment-marker ISH, and retinoic acid treatment

    PMID:30575756

    Open questions at the time
    • Direct versus indirect regulation of segment markers unresolved
    • Relevance to mammalian EMX1 function unknown
  15. 2021 Medium

    In a tumor context, gain- and loss-of-function established EMX1 (with EMX2) as a tumor suppressor that represses stem-cell genes and the Wnt/β-catenin pathway, with Wnt activation epistatically bypassing EMX repression.

    Evidence Ectopic expression and KO models, in vitro/in vivo sarcoma assays, Wnt activation rescue, qRT-PCR

    PMID:34016958 PMID:34364391

    Open questions at the time
    • Direct EMX1 target genes within the stemness/Wnt program not all mapped
    • Single lab
  16. 2022 Medium

    Pinpointing a direct repression mechanism, EMX1 was shown to bind the WASF2 promoter and suppress Wnt/β-catenin signaling through WASF2 inhibition in glioma cells.

    Evidence ChIP-qPCR, luciferase reporter, gain/loss-of-function, and WASF2-restoration rescue

    PMID:35849030

    Open questions at the time
    • Generalizability of WASF2 axis beyond spinal cord glioma unknown
    • Single lab
  17. 2023 Medium

    Demonstrating context- and isoform-dependent activity, full-length EMX1 was shown to bind the EGFR promoter and activate EGFR-ERK signaling to drive hepatocellular carcinoma invasion, unlike the EMX1-X1 splice isoform.

    Evidence Gain-of-function and siRNA knockdown, migration/invasion and metastasis assays, promoter binding, and bisulfite sequencing

    PMID:38007497

    Open questions at the time
    • Mechanism switching EMX1 between repressor and activator roles unresolved
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • How EMX1's DNA-binding and target selectivity convert into opposite outcomes—transcriptional activation (NRP1, EGFR) versus repression (WASF2, stem-cell/Wnt genes)—across neural development and tumor contexts remains unresolved.
  • No structural model of EMX1-DNA binding in the corpus
  • Cofactors determining activation versus repression unidentified
  • Genome-wide direct target map absent

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 4 GO:0003677 DNA binding 2
Localization
GO:0005634 nucleus 2
Pathway
R-HSA-1266738 Developmental Biology 4 R-HSA-1643685 Disease 4 R-HSA-162582 Signal Transduction 3 R-HSA-74160 Gene expression (Transcription) 3
Partners
EMX2WDR11

Evidence

Reading pass · 21 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 Emx1 knockout mice (deletion of N-terminal coding region) lack most or all of their corpus callosum, the principal fiber tract connecting the cerebral hemispheres, demonstrating a role for EMX1 in corpus callosum formation. Histology and molecular properties of the cerebral cortex appeared normal in mutants. Gene targeting/knockout mouse, histology Developmental biology Medium 8812118
1996 EMX1 protein localizes to nuclei of proliferating, differentiating, and mature cortical neurons across all layers in the developing cerebral cortex, and is also present in axons of olfactory sensory neurons along their entire length including terminals in olfactory bulb glomeruli. Immunohistochemistry, subcellular localization Mechanisms of development Medium 8843394
1996 Emx1 is expressed in both proliferating (ventricular zone) and differentiated (postmitotic) neurons throughout all cortical layers during development, in contrast to Emx2 which is restricted to proliferating cells; this indicates EMX1 functions in both proliferating and differentiating cortical cells. In situ hybridization across developmental stages The European journal of neuroscience Medium 8743751
1997 Loss-of-function mutation of Emx1 in mice produces subtle or undetectable defects in early forebrain patterning where Emx1 expression overlaps Emx2, suggesting Emx2 can compensate for Emx1 during early dorsal forebrain patterning. In double mutants (Emx1;Emx2), forebrain defects are far more severe, demonstrating functional redundancy and cooperation between the two paralogs. Homologous recombination knockout, comparative phenotypic analysis of single and double mutants Development (Cambridge, England) High 9006071
2001 Emx1 expression is restricted to pyramidal (glutamatergic) neurons of the cerebral cortex, identified by co-localization with glutamate, pyramidal neuron morphology (soma shape, ultrastructure), laminar distribution (all layers except I), and radial clone analysis in chimeric mice. In situ hybridization, immunocytochemistry (light and electron microscopy), double-labeling with glutamate, chimeric mouse clone analysis Cerebral cortex (New York, N.Y. : 1991) High 11709490
2002 Using Emx1(IRES)cre fate mapping, the Emx1-expressing lineage gives rise to radial glia, Cajal-Retzius cells, glutamatergic neurons, astrocytes, and oligodendrocytes of pallial structures, but not to most pallial GABAergic neurons, which arise outside the Emx1-expressing lineage. Cre-mediated genetic fate mapping (IRES-Cre knock-in × reporter strains) The Journal of neuroscience : the official journal of the Society for Neuroscience High 12151506
2002 Emx1 single-mutant mice show no detectable changes in neocortical arealization as assessed by seven area marker genes and thalamocortical projections, indicating EMX1 does not regulate neocortical area specification. Emx1;Emx2 double mutants show similar area shifts as Emx2 single mutants, confirming absence of EMX1 contribution to arealization. Gene targeting KO, in situ hybridization of area markers, axon tracing, epistasis analysis The Journal of neuroscience : the official journal of the Society for Neuroscience Medium 12196586
2002 In Emx1/Emx2 double mutant neocortex, tangential migration of cells from the ganglionic eminence into the neocortex was greatly inhibited; transplanted wild-type ganglionic eminence cells also failed to migrate into double mutant telencephalon, indicating a non-cell-autonomous requirement for Emx1/Emx2 in the cortical environment for tangential cell migration. Cajal-Retzius cells and subplate neurons were absent throughout development in double mutants. Emx1/Emx2 double knockout, BrdU labeling, cell transplantation assay, immunohistochemistry Development (Cambridge, England) High 12091317
2003 Emx1;Emx2 double mutant cortex displays greatly reduced size, absent hippocampus and dentate gyrus, defective olfactory bulb growth, abnormal cortical lamination, reduced Cajal-Retzius and subplate neurons, failure of cortical efferent axons to exit the cortex, and thalamocortical axon pathfinding defects. The TCA pathfinding defect is non-cell autonomous, supporting that cortical efferents act as scaffolds to guide TCAs into cortex. Double knockout mouse analysis, axon tracing, marker gene expression, immunohistochemistry The Journal of comparative neurology High 12561075
2004 Emx1 and Emx2 cooperate in establishing the boundary between the roof plate and archipallium; in double mutants the archipallium is transformed into roof plate, the cortical hem signaling center fails to form, and the choroid plexus does not develop, indicating Emx1/Emx2 are required for archipallial identity against roof plate fate. Emx1;Emx2 double knockout, immunohistochemistry, in situ hybridization Mechanisms of development Medium 15147765
2006 Ectopic expression of Emx1 or Emx2 in the chick telencephalic midline region (prior to HH13) instructed neuroepithelial identity instead of choroid plexus fate, with expansion of Gli3 and Lhx2 (neuroepithelial markers) and downregulation of Otx2 and BMP7 (choroid plexus markers). This demonstrates that Emx1 suppresses choroid plexus fate and promotes neuroepithelial identity. In ovo misexpression (chick electroporation/lentivirus), in situ hybridization for fate markers Developmental biology Medium 16793035
2007 Emx1 null mice exhibit a smaller dentate gyrus, reduced number of proliferating progenitor cells and immature neurons in the adult dentate gyrus (but normal subventricular zone neurogenesis), indicating EMX1 is required for adult hippocampal neurogenesis. Emx1 deletion also impaired forelimb skill reaching and attenuated training-induced hippocampal neurogenesis. Knockout mouse, unbiased stereology, BrdU/doublecortin labeling, behavioral testing Experimental neurology Medium 17490651
2010 WDR11 physically interacts with EMX1 (homeodomain transcription factor), and disease-associated WDR11 missense mutations in WD domains reduce or abolish this interaction, implicating the WDR11-EMX1 interaction in development of olfactory neurons and normal puberty onset. Co-immunoprecipitation/protein-protein interaction assay, identification of chromosomal translocation breakpoint, missense mutation functional analysis American journal of human genetics Medium 20887964
2015 EMX1 is required for midline crossing of anterior cingulate cortex axons in the corpus callosum; in Emx1 knockout mice (C57Bl/6 background) cingulate axons show reduced NRP1 expression and form aberrant bundles. EMX1 directly activates Nrp1 transcription in vitro, and overexpression of NRP1 in Emx1 KO mice rescues the midline-crossing defect. Knockout mouse, axon tracing, in vitro transcriptional activation assay, rescue experiment (NRP1 overexpression in KO) Development (Cambridge, England) High 26534986
2016 Deletion of Emx1 results in expanded frontal/motor areas and reduced primary visual area (posterio-medial shift of area boundaries), demonstrating EMX1 is required for neocortical area patterning. This was consistent across two independent Emx1 deletion lines examined at mature (postnatal) ages. Two independent knockout mouse lines, area marker gene expression, mature-age analysis PloS one Medium 26901526
2016 Emx1 KO neural stem cells (NSCs) show reduced frequency and self-renewal capacity in the embryonic brain, and reduced migratory capacity in response to serum or VEGF. Proteomics revealed reduced phospho-Cofilin1 (p-Cofilin1) in KO NSCs; VEGF-induced Cofilin1 phosphorylation and FLK expression were impaired in KO NSCs, suggesting a defect in Cofilin1 phosphorylation downstream of VEGF contributes to reduced neurogenesis. Neurosphere assay, Boyden chamber migration assay, 2D-PAGE/tandem mass spectrometry proteomics, western blot for p-Cofilin1, Emx1 KO mouse Frontiers in molecular neuroscience Medium 27799894
2018 In zebrafish, emx1 loss of function alters the domains of distal nephron segments without affecting cell turnover, indicating emx1 directs distal segment fate during nephrogenesis. Retinoic acid negatively regulates emx1 expression. Genetic epistasis places emx1 downstream of mecom and tbx2b, and emx1 regulates expression of irx3b, irx1a (distal segmentation) and sim1a (corpuscle of Stannius formation). Loss-of-function (morpholino/mutant), genetic epistasis, in situ hybridization for segment markers, retinoic acid treatment Scientific reports Medium 30575756
2021 EMX1 and EMX2 act as tumor suppressors in sarcomas by repressing stem cell regulatory genes (OCT4, SOX2, KLF4, MYC, NANOG, NES, PROM1). EMX1/EMX2 overexpression reduces tumorigenic properties and cancer stem cell phenotype in vitro and in vivo; Emx KO murine models show more aggressive, infiltrative sarcomas with higher stem cell gene expression. Ectopic overexpression and KO mouse models, in vitro tumorigenesis assays, in vivo xenograft/carcinogen models, qRT-PCR Cell death & disease Medium 34016958
2021 EMX1 and EMX2 negatively regulate the canonical Wnt/β-catenin pathway in sarcoma cells; constitutive Wnt pathway activation bypasses EMX1/EMX2-dependent stemness repression and restores aggressive/stem phenotype even when EMX genes are overexpressed, placing EMX1 upstream of Wnt signaling in this context. Ectopic expression, Wnt pathway activation experiments, qRT-PCR, in vitro and in vivo sarcoma models, bioinformatic validation Journal of experimental & clinical cancer research : CR Medium 34364391
2022 EMX1 binds to the WASF2 promoter to suppress its transcription in spinal cord glioma cells; silencing WASF2 restores EMX1's tumor-suppressive effect, and EMX1 suppresses Wnt/β-catenin signaling via WASF2 inhibition. ChIP-qPCR, luciferase reporter assay, gain/loss-of-function in vitro and in vivo (xenograft), rescue with WASF2 restoration Brain and behavior Medium 35849030
2023 EMX1 full-length isoform (EMX1-FL) binds to the EGFR promoter, promotes EGFR transcription, and activates EGFR-ERK signaling to trigger hepatocellular carcinoma cell migration and invasion. The alternative splicing isoform EMX1-X1 does not induce migration/invasion. Gene body hypermethylation positively correlates with EMX1 mRNA expression in HCC. Gain-of-function and siRNA knockdown, in vitro migration/invasion assays, in vivo metastasis models, promoter binding assays, bisulfite sequencing Cell death & disease Medium 38007497

Source papers

Stage 0 corpus · 45 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Cortical excitatory neurons and glia, but not GABAergic neurons, are produced in the Emx1-expressing lineage. The Journal of neuroscience : the official journal of the Society for Neuroscience 1053 12151506
2000 Pallial and subpallial derivatives in the embryonic chick and mouse telencephalon, traced by the expression of the genes Dlx-2, Emx-1, Nkx-2.1, Pax-6, and Tbr-1. The Journal of comparative neurology 760 10906711
1997 Emx1 and Emx2 functions in development of dorsal telencephalon. Development (Cambridge, England) 312 9006071
1998 Expression of the Emx-1 and Dlx-1 homeobox genes define three molecularly distinct domains in the telencephalon of mouse, chick, turtle and frog embryos: implications for the evolution of telencephalic subdivisions in amniotes. Development (Cambridge, England) 274 9570774
2004 Expression of Dbx1, Neurogenin 2, Semaphorin 5A, Cadherin 8, and Emx1 distinguish ventral and lateral pallial histogenetic divisions in the developing mouse claustroamygdaloid complex. The Journal of comparative neurology 191 15174069
2002 Distinct actions of Emx1, Emx2, and Pax6 in regulating the specification of areas in the developing neocortex. The Journal of neuroscience : the official journal of the Society for Neuroscience 187 12196586
1996 Emx1 and Emx2 show different patterns of expression during proliferation and differentiation of the developing cerebral cortex in the mouse. The European journal of neuroscience 176 8743751
2010 WDR11, a WD protein that interacts with transcription factor EMX1, is mutated in idiopathic hypogonadotropic hypogonadism and Kallmann syndrome. American journal of human genetics 131 20887964
2003 Emx1 and Emx2 cooperate to regulate cortical size, lamination, neuronal differentiation, development of cortical efferents, and thalamocortical pathfinding. The Journal of comparative neurology 126 12561075
2004 Expression of the genes Emx1, Tbr1, and Eomes (Tbr2) in the telencephalon of Xenopus laevis confirms the existence of a ventral pallial division in all tetrapods. The Journal of comparative neurology 118 15174073
2002 Absence of Cajal-Retzius cells and subplate neurons associated with defects of tangential cell migration from ganglionic eminence in Emx1/2 double mutant cerebral cortex. Development (Cambridge, England) 108 12091317
2001 Emx1 is a marker for pyramidal neurons of the cerebral cortex. Cerebral cortex (New York, N.Y. : 1991) 99 11709490
1996 Mutation of the Emx-1 homeobox gene disrupts the corpus callosum. Developmental biology 90 8812118
2000 Specificity and efficiency of Cre-mediated recombination in Emx1-Cre knock-in mice. Biochemical and biophysical research communications 86 10873661
1996 EMX1 homeoprotein is expressed in cell nuclei of the developing cerebral cortex and in the axons of the olfactory sensory neurons. Mechanisms of development 83 8843394
2003 Emx1, emx2 and pax6 in specification, regionalization and arealization of the cerebral cortex. Cerebral cortex (New York, N.Y. : 1991) 81 12764040
2009 Emx1-lineage progenitors differentially contribute to neural diversity in the striatum and amygdala. The Journal of neuroscience : the official journal of the Society for Neuroscience 56 20016109
2004 Emx1 and Emx2 cooperate in initial phase of archipallium development. Mechanisms of development 56 15147765
2000 Emx1-specific expression of foreign genes using "knock-in" approach. Biochemical and biophysical research communications 41 10772936
2007 Reduced hippocampal neurogenesis and skill reaching performance in adult Emx1 mutant mice. Experimental neurology 34 17490651
2002 Reduced anxiety-- and depression-like behaviors in Emx1 homozygous mutant mice. Brain research 29 12020859
2006 The transcription factors Emx1 and Emx2 suppress choroid plexus development and promote neuroepithelial cell fate. Developmental biology 28 16793035
2002 Developmental expression of zebrafish emx1 during early embryogenesis. Gene expression patterns : GEP 27 12617801
2018 Homeogene emx1 is required for nephron distal segment development in zebrafish. Scientific reports 24 30575756
2011 Emx1-expressing neural stem cells in the subventricular zone give rise to new interneurons in the ischemic injured striatum. The European journal of neuroscience 24 21219481
2000 Normal corpus callosum in Emx1 mutant mice with C57BL/6 background. Biochemical and biophysical research communications 23 11027526
2013 Gli3 is required in Emx1+ progenitors for the development of the corpus callosum. Developmental biology 21 23396189
2015 EMX1 regulates NRP1-mediated wiring of the mouse anterior cingulate cortex. Development (Cambridge, England) 16 26534986
2016 Deciphering the Role of Emx1 in Neurogenesis: A Neuroproteomics Approach. Frontiers in molecular neuroscience 15 27799894
2021 Regulation of sarcomagenesis by the empty spiracles homeobox genes EMX1 and EMX2. Cell death & disease 14 34016958
2023 DNA methylation-activated full-length EMX1 facilitates metastasis through EMX1-EGFR-ERK axis in hepatocellular carcinoma. Cell death & disease 13 38007497
2021 Empty spiracles homeobox genes EMX1 and EMX2 regulate WNT pathway activation in sarcomagenesis. Journal of experimental & clinical cancer research : CR 12 34364391
2012 Reliable in vivo identification of both GABAergic and glutamatergic neurons using Emx1-Cre driven fluorescent reporter expression. Cell calcium 12 22658827
2011 Hypomyelination phenotype caused by impaired differentiation of oligodendrocytes in Emx1-cre mediated Cdk5 conditional knockout mice. Neurochemical research 12 21210220
2016 Emx1 Is Required for Neocortical Area Patterning. PloS one 10 26901526
2013 Restoration of SMN to Emx-1 expressing cortical neurons is not sufficient to provide benefit to a severe mouse model of Spinal Muscular Atrophy. Transgenic research 8 23512182
2022 Emx1-Cre Is Expressed in Peripheral Autonomic Ganglia That Regulate Central Cardiorespiratory Functions. eNeuro 6 36192157
2024 Role of PAX6, TRPA1, BCL11B, MCOLN2, CUX1, EMX1 in colorectal cancer and osteosarcoma. Medicine 4 38306561
2024 Extra-cerebral recombination activity of Emx1-Cre and nestin-Cre in the kidney. Frontiers in cell and developmental biology 4 39479517
2022 "SIT" with Emx1-NuTRAP Mice: Simultaneous INTACT and TRAP for Paired Transcriptomic and Epigenetic Sequencing. Current protocols 3 36287035
2016 Training of the impaired forelimb after traumatic brain injury enhances hippocampal neurogenesis in the Emx1 null mice lacking a corpus callosum. Behavioural brain research 3 27614007
2022 EMX1 functions as a tumor inhibitor in spinal cord glioma through transcriptional suppression of WASF2 and inactivation of the Wnt/β-catenin axis. Brain and behavior 1 35849030
2014 Rostral-caudal distribution of Emx1-lineage stem/transit amplifying cells and lineage progression in embryonic cortex depend on Hedgehog signaling. Developmental neurobiology 1 24771701
2025 Transferring Mouse Emx1 and Emx2 Lentiviruses into the Chicken Embryonic Brain and Their Implication to the Organization and Evolution of the Amniote Pallium. Developmental neuroscience 0 39864431
2024 Restoring social deficits in IRSp53-deleted mice: chemogenetic inhibition of ventral dentate gyrus Emx1-expressing cells. Translational psychiatry 0 39375329

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