Affinage

HOXB9

Homeobox protein Hox-B9 · UniProt P17482

Length
250 aa
Mass
28.1 kDa
Annotated
2026-04-28
100 papers in source corpus 30 papers cited in narrative 30 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

HOXB9 is a homeodomain transcription factor that patterns the axial skeleton and mammary gland during development and, in adult tissues, regulates cell proliferation, angiogenesis, epithelial–mesenchymal transitions, and metastatic competence. It binds promoters of diverse target genes—including angiogenic factors (VEGF, bFGF, IL-8), cell-cycle regulators (CDK6, RBL2), oncogenes (KRAS, EZH2), and SLC7A11—often as a heterodimer with cofactors such as PBX2 and BTG1/BTG2 (PMID:19576624, PMID:20080567, PMID:37272544, PMID:10617598, PMID:36001969). Its transcriptional activity and stability are tuned by post-translational modifications: PCAF-mediated acetylation at K27 triggers nuclear-to-cytoplasmic translocation and attenuates oncogenic target gene expression, while AMPKα-mediated phosphorylation at T133 promotes Praja2-dependent ubiquitination and proteasomal degradation (PMID:27613418, PMID:29654889, PMID:36001969). Loss-of-function studies in mice demonstrate that Hoxb9 is required for thoracic rib and sternum specification and, together with paralogous Hox genes, for pregnancy-induced mammary epithelial expansion (PMID:9013929, PMID:9892669).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1993 Medium

    Cross-species ectopic expression established that mouse HoxB9 retains homeotic selector function, converting Drosophila head structures toward posterior identities, thereby placing HOXB9 within a functionally conserved HOX framework.

    Evidence Heat-shock–driven expression of mouse HoxB9 in Drosophila, scored for homeotic transformations

    PMID:8105876

    Open questions at the time
    • Transformation phenotype partially overlaps Antp and Abd-B classes, so precise paralog-equivalent identity in flies is ambiguous
    • No DNA-binding target analysis in this system
  2. 1997 High

    Gene targeting in mice revealed that Hoxb9 is required for normal rib and sternum morphogenesis, and genetic epistasis with Hoxa9 showed synergistic skeletal defects, establishing paralog cooperation in thoracic patterning.

    Evidence Targeted gene disruption in mice and Hoxa9/Hoxb9 double-mutant intercross with skeletal phenotyping

    PMID:9013929

    Open questions at the time
    • Downstream skeletal target genes not identified
    • Mechanisms of paralog cooperation (shared or distinct targets) unknown
  3. 1999 High

    Triple-paralog knockout (Hoxa9/Hoxb9/Hoxd9) demonstrated a direct requirement for these Hox genes in pregnancy-induced mammary gland expansion, extending HOXB9 function beyond embryonic skeletal patterning to adult epithelial organogenesis.

    Evidence Compound knockout mice; mammary gland hypoplasia post-pregnancy phenotype

    PMID:9892669

    Open questions at the time
    • Individual contribution of Hoxb9 vs. paralog redundancy in mammary gland not fully resolved
    • Mammary target genes not identified
  4. 2000 High

    Identification of BTG1 and BTG2 as HOXB9-interacting cofactors that enhance its DNA binding and transcriptional activity provided the first molecular insight into how HOXB9 assembles productive transcription complexes, with the N-terminal domain required for activation.

    Evidence Yeast two-hybrid, co-immunoprecipitation, DNA-binding assay, and reporter assays

    PMID:10617598

    Open questions at the time
    • Endogenous genomic targets of the HOXB9–BTG complex not mapped
    • Structural basis of the interaction unknown
  5. 2009 High

    Two landmark studies established HOXB9 as an activator of angiogenic factors, ErbB ligands, and TGF-β in breast cancer EMT and as a WNT/TCF-responsive driver of organ-specific metastasis in lung adenocarcinoma, repositioning HOXB9 as a cancer-relevant transcription factor.

    Evidence Gain/loss-of-function in breast and lung cancer cell lines; in vivo xenograft, metastasis, and angiogenesis models

    PMID:19576624 PMID:20080567

    Open questions at the time
    • Direct versus indirect transcriptional control of individual angiogenic targets not distinguished at this stage
    • Relative importance of WNT input versus other upstream signals in different tumor types unresolved
  6. 2012 Medium

    ChIP and homeodomain-deletion analysis confirmed that HOXB9 directly occupies promoters of angiogenic and growth factor genes through its homeodomain, linking sequence-specific DNA binding to its tumorigenic output.

    Evidence ChIP assay and homeodomain deletion mutant in breast cancer cells with soft-agar colony assay

    PMID:22863320

    Open questions at the time
    • Genome-wide binding profile not determined
    • Cofactor requirements at individual promoters not dissected
  7. 2014 Medium

    Context-dependent roles of HOXB9 in epithelial plasticity were revealed: it promotes EMT via TGF-β1 in hepatocellular carcinoma but induces MET by repressing Snail/Twist/ZEB1 in colon adenocarcinoma, indicating tissue-specific transcriptional programs.

    Evidence Knockdown/overexpression, EMT marker analysis, RNA-seq, and in vivo metastasis models in HCC and colon cancer cells

    PMID:25025961 PMID:25081022

    Open questions at the time
    • Molecular basis for opposing EMT/MET outcomes in different tissues not resolved
    • Tissue-specific cofactors that redirect HOXB9 activity not identified
  8. 2016 High

    Discovery that PCAF acetylates HOXB9 at K27 (reversed by SIRT1) introduced the first post-translational switch controlling HOXB9: acetylation suppresses JMJD6 transcription and reduces migration, revealing a regulatory layer beyond expression control.

    Evidence In vitro acetylation assay, site-directed mutagenesis (K27), ChIP, migration and xenograft assays

    PMID:27613418

    Open questions at the time
    • Full spectrum of acetylation-sensitive target genes unknown
    • Whether other acetyltransferases modify K27 not tested
  9. 2018 High

    Acetylation at K27 was shown to trigger nucleocytoplasmic translocation of HOXB9, functionally uncoupling it from nuclear targets such as EZH2 and explaining how acetylation suppresses oncogenic transcription.

    Evidence Subcellular fractionation, immunofluorescence, luciferase reporter, and K27 mutagenesis in colon cancer cells

    PMID:29654889

    Open questions at the time
    • Export mechanism (NES, carrier) mediating cytoplasmic translocation not characterized
    • Potential cytoplasmic functions of acetylated HOXB9 unknown
  10. 2022 High

    AMPKα phosphorylation of HOXB9 at T133 was identified as a second post-translational switch, triggering Praja2-mediated ubiquitination and degradation; blocking this axis stabilizes HOXB9 and upregulates its direct target KRAS, linking metabolic sensing to HOXB9 protein turnover.

    Evidence Phosphorylation assay, T133A mutagenesis, ubiquitination assay, ChIP at KRAS promoter, xenograft models

    PMID:36001969

    Open questions at the time
    • Other E3 ligases that may target HOXB9 not surveyed
    • Integration between AMPK-mediated degradation and PCAF-mediated acetylation signals not examined
  11. 2023 Medium

    HOXB9 was shown to heterodimerize with PBX2 to co-occupy and activate the CDK6 promoter, directly coupling HOXB9 to G1/S cell-cycle progression and explaining its proliferation-promoting activity in gastric cancer.

    Evidence Co-immunoprecipitation, ChIP, reporter assay, CDK6 inhibitor rescue, and cell-cycle analysis in gastric cancer cells

    PMID:37272544

    Open questions at the time
    • Whether PBX2 is required at all HOXB9 target promoters unknown
    • Structural details of HOXB9-PBX2 heterodimer not available
  12. 2024 Medium

    Identification of SLC7A11 as a direct HOXB9 transcriptional target linked HOXB9 to ferroptosis regulation; CAP-enhanced PCAF-mediated acetylation destabilizes HOXB9 and suppresses SLC7A11, revealing convergence of the acetylation and ubiquitination regulatory axes on redox defense.

    Evidence ChIP, luciferase, Co-IP, ubiquitination assay, ferroptosis assays, and xenograft in lung cancer

    PMID:39127016

    Open questions at the time
    • Whether HOXB9-driven SLC7A11 regulation occurs outside the CAP treatment context not established
    • Relative contribution of HOXB9 versus other transcription factors to SLC7A11 expression unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the genome-wide direct target repertoire of HOXB9 (no ChIP-seq available), the structural basis for cofactor selectivity (PBX2, BTG1/2), the mechanism of acetylation-driven nuclear export, and how tissue context dictates opposing EMT vs. MET outputs.
  • No genome-wide binding data (ChIP-seq/CUT&RUN) for HOXB9
  • No structural model of HOXB9 homeodomain with DNA or cofactors
  • Crosstalk between K27 acetylation and T133 phosphorylation not investigated
  • Molecular determinants of tissue-specific EMT versus MET outcomes uncharacterized

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 9 GO:0003677 DNA binding 4
Localization
GO:0005634 nucleus 2 GO:0005829 cytosol 1
Pathway
R-HSA-74160 Gene expression (Transcription) 8 R-HSA-1266738 Developmental Biology 3 R-HSA-162582 Signal Transduction 3 R-HSA-1640170 Cell Cycle 2

Evidence

Reading pass · 30 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 Paralogous Hox genes Hoxa9, Hoxb9, and Hoxd9 are required for expansion and/or differentiation of mammary epithelial ductal system in response to pregnancy; triple knockout mice cannot raise pups due to mammary gland hypoplasia after pregnancy, demonstrating a direct functional role in adult mammary development. Targeted gene disruption (loss-of-function) in mice with specific phenotypic readout (mammary gland hypoplasia post-pregnancy) Proceedings of the National Academy of Sciences of the United States of America High 9892669
1997 Targeted disruption of hoxb-9 causes defects in first and second rib development and sternal abnormalities; compound hoxa-9/hoxb-9 double knockouts show synergistic, more severe thoracic skeletal defects, demonstrating genetic interaction between paralogous Hox genes in thoracic skeletal specification. Targeted gene disruption (loss-of-function) and genetic epistasis via double-mutant intercross Developmental biology High 9013929
2009 HOXB9, a WNT/TCF target gene, mediates chemotactic invasion and colony outgrowth of lung adenocarcinoma cells and enhances competence to colonize bone and brain; reduction of HOXB9 attenuates brain and bone metastasis in mice independently of primary lung tumor growth. Gain-of-function/loss-of-function (shRNA knockdown, overexpression) in metastatic lung adenocarcinoma cell lines and mouse metastasis models Cell High 19576624
2009 HOXB9 is a transcription factor that induces expression of multiple angiogenic factors (VEGF, bFGF, IL-8, ANGPTL-2) and ErbB ligands (amphiregulin, epiregulin, neuregulins) as well as TGF-β, leading to increased cell motility and acquisition of mesenchymal phenotypes; in vivo, HOXB9 promotes formation of well-vascularized tumors that metastasize to the lung. Overexpression in breast cancer cells, in vitro motility assays, in vivo xenograft and metastasis models, gene expression analysis Proceedings of the National Academy of Sciences of the United States of America High 20080567
2000 BTG1 and BTG2 physically interact with HOXB9 homeodomain protein (identified by yeast two-hybrid screening), enhance HOXB9-mediated transcription in transfected cells, form a complex on HOXB9-responsive DNA targets, and facilitate HOXB9 binding to DNA; the transcriptional activity depends on the N-terminal activation domain of HOXB9. Yeast two-hybrid screening, co-immunoprecipitation, DNA-binding assay, transfection reporter assay The Journal of biological chemistry High 10617598
2016 PCAF acetyltransferase interacts with and acetylates HOXB9 at lysine 27 (K27) both in vivo and in vitro; this acetylation can be reversed by deacetylase SIRT1. AcK27-HOXB9 suppresses transcription of target gene JMJD6 by occupying its promoter, and decreases cancer cell migration and tumor growth compared to wild-type HOXB9. Co-immunoprecipitation, in vitro acetylation assay, site-directed mutagenesis (K27), ChIP assay, functional cell migration and xenograft assays Nucleic acids research High 27613418
2018 AcK27-HOXB9 translocates from the nucleus to the cytoplasm, suppressing transcription of oncogenic EZH2, whereas non-acetylated HOXB9 remains nuclear and promotes EZH2 expression and colon cancer progression; subcellular fractionation and immunofluorescence confirmed nucleocytoplasmic translocation. Subcellular fractionation, immunofluorescence, luciferase reporter assay, site-directed mutagenesis Cancer letters High 29654889
2022 AMPKα phosphorylates HOXB9 at threonine 133 (T133), promoting E3 ligase Praja2-mediated HOXB9 ubiquitination and degradation; blocking HOXB9 phosphorylation (AMPKα1/2 depletion or HOXB9-T133A mutant) stabilizes HOXB9, upregulates KRAS (identified as a direct HOXB9 transcriptional target), and promotes tumor growth. Phosphorylation assay, site-directed mutagenesis (T133A), ubiquitination assay, ChIP, cell and xenograft models Cell reports High 36001969
2012 HOXB9 binds directly to promoters of tumor growth and angiogenic factors (via its homeodomain) and regulates their expression; the homeodomain is required for transcriptional regulation of these factors and for 3D colony formation in soft agar, indicating a functional role in tumorigenesis. ChIP assay, homeodomain deletion mutant analysis, soft agar colony formation assay The FEBS journal Medium 22863320
2011 HOXB9 is transcriptionally regulated by estrogen (E2) via estrogen-response elements (EREs) in its promoter; estrogen receptors ERα and ERβ and histone methylases MLL1 and MLL3 bind to HOXB9 EREs in an E2-dependent manner and are required for E2-mediated HOXB9 transcriptional activation. Luciferase reporter assay, chromatin immunoprecipitation (ChIP), ERE mutational analysis Biochemistry Medium 21428455
2016 BPA (bisphenol-A) induces HOXB9 expression via EREs in the HOXB9 promoter, requiring estrogen receptors and chromatin modification factors (MLL histone methylase, CBP/P300 acetylases) that bind to HOXB9 EREs in the presence of BPA, leading to histone methylation and acetylation and gene activation. Luciferase reporter assay, ChIP, ERE mutational analysis, in vivo animal (OVX rat) model Gene Medium 27182052
2008 The Hoxb9 promoter forms secondary DNA structures that regulate promoter activity; FBXL10 (KDM2B) was isolated as a protein that specifically binds these secondary-structured promoter DNA sequences and affects Hoxb9 promoter activity. DNA secondary structure analysis, promoter-activity reporter assay, protein isolation/pulldown from nuclear extracts Nucleic acids research Medium 18276649
2016 miR-192 suppresses HOXB9 (and EGR1) expression, thereby globally downregulating angiogenic pathways in cancer cells; HOXB9 is identified as a central downstream effector of miR-192's anti-angiogenic function. miRNA overexpression/inhibition in vitro and in vivo, gene expression analysis, in vivo DOPC nanoliposome delivery Nature communications Medium 27041221
2014 HOXB9 promotes epithelial-to-mesenchymal transition (EMT) in hepatocellular carcinoma cells through the TGF-β1 pathway; knockdown decreases migration and invasion while overexpression increases these phenotypes. siRNA knockdown, overexpression, cell migration/invasion assays, EMT marker analysis Clinical and experimental medicine Medium 25081022
2014 HOXB9 promotes mesenchymal-to-epithelial transition (MET) in colon adenocarcinoma cells by downregulating EMT-promoting transcription factors (Snail, Twist, FOXC2, ZEB1) and upregulating epithelial proteins (E-cadherin, claudins, occludin, ZO-1); overexpression inhibits colon cancer cell growth, migration, invasion, and metastasis in vivo. Overexpression/knockdown, RNA sequencing, in vitro cell assays, in vivo xenograft/metastasis model British journal of cancer Medium 25025961
2015 HOXB9 hexapeptide motif acts as a 'brake' on its own MET induction and tumor suppression activity in gastric carcinoma; a HOXB9 mutant lacking the hexapeptide motif showed more potent MET induction and tumor suppression than wild-type HOXB9. Domain deletion/mutant analysis, cell proliferation/migration/invasion assays, structural analysis Oncotarget Medium 26536658
2018 E2F1 directly binds the HOXB9 promoter region (-404 to -392) and transcriptionally activates HOXB9 in breast cancer cells; confirmed by EMSA, ChIP, and mutational analysis of the binding site. Dual luciferase reporter assay, EMSA, ChIP, mutational analysis, Q-PCR in multiple cell lines PloS one Medium 25136922
2019 GRP78 regulates HOXB9 expression through the Wnt signaling pathway by chaperoning LRP6; GRP78 promotes LRP6 maturation, and knockdown of GRP78 leads to LRP6 misfolding and ERAD degradation, reducing mature LRP6 and suppressing Wnt/HOXB9 signaling, thereby decreasing HCC invasion and metastasis. GRP78 knockdown, Wnt pathway inhibition, LRP6 maturation assay, rescue experiments with HOXB9 overexpression Experimental cell research Medium 31310747
2022 HOXB9 blocks cell cycle progression at G0/G1 in pancreatic cancer cells by transcriptionally upregulating RBL2 and inhibiting c-Myc; DNMT1 suppresses HOXB9 expression in pancreatic cancer by promoting methylation of the HOXB9 promoter; the DNMT1/HOXB9/RBL2/c-Myc axis regulates pancreatic cancer cell proliferation. Cell cycle PCR array, flow cytometry, ChIP-qPCR, luciferase assay, promoter methylation analysis, xenograft models Cancer letters Medium 35182659
2023 HOXB9 interacts with PBX2 to form a heterodimer that transcriptionally upregulates CDK6; E2F1 activates HOXB9 expression upstream; depletion of HOXB9 causes G1-phase cell cycle arrest by downregulating CDK6 and a subset of cell cycle genes in gastric cancer cells. Co-immunoprecipitation, ChIP, reporter assay, siRNA knockdown, flow cytometry (cell cycle), CDK6 inhibitor rescue The Journal of pathology Medium 37272544
2018 HOXB9 directly binds the promoter of microRNA-765 and facilitates its transcription, which in turn targets FOXA2; this HOXB9-miR-765-FOXA2 axis promotes cancer stem cell self-renewal and ER stress resistance in melanoma. ChIP assay, luciferase reporter, miRNA target validation, cancer stem cell assays The Journal of investigative dermatology Medium 29408459
2016 HOXB9 is expressed in mammalian oocytes and early embryos; its protein distribution is dynamic, mainly nuclear, with strong nuclear staining in trophoblastic cells, while epiblast cells progressively downregulate HOXB9 after blastocyst stage; presence in apical vacuoles of mouse visceral endoderm cells suggests roles in early embryonic development. Immunofluorescence localization in mouse and bovine oocytes and early embryos (multiple developmental stages) PloS one Low 27798681
2015 GalNAc-T14 increases sensitivity of the WNT response and stability of β-catenin protein, leading to induced HOXB9 expression and acquisition of invasive phenotype in lung cancer; pharmacological inhibition of β-catenin suppressed HOXB9 expression and invasion, placing HOXB9 downstream of the GalNAc-T14/WNT/β-catenin axis. Microarray, biochemical analysis, pharmacological inhibition (β-catenin inhibitor), invasion assays Oncotarget Low 26544896
2018 HOXB9 promotes E2F3 expression by directly binding to the E2F3 promoter in endometrial cancer cells; E2F3 knockdown abolished HOXB9-enhanced cell migration. ChIP assay, reporter assay, siRNA knockdown, migration assay Cell death & disease Low 29724991
2024 CAP (cold atmospheric plasma) promotes HOXB9 interaction with PCAF acetyltransferase, increasing HOXB9 acetylation; acetylated HOXB9 has altered ubiquitination and reduced protein stability. HOXB9 directly targets and regulates SLC7A11 transcription (confirmed by luciferase and ChIP assays); CAP suppresses HOXB9/SLC7A11 axis to enhance ferroptosis in lung cancer cells. ChIP, luciferase assay, Co-IP, ubiquitination assay, ferroptosis assays, xenograft model Redox biology Medium 39127016
2020 HOXB9-dependent expression of MMP9 in NSCLC cells leads to degradation of intercellular adhesion proteins in BBB endothelial cells and promotes tumor cell transmigration; knockdown and overexpression experiments using an in vitro BBB model confirmed this mechanism. In vitro BBB model, knockdown/overexpression, MMP9 expression analysis, endothelial junctional protein analysis Aging Low 33411683
2024 HOXB9 directly binds to the promoter of SPP1 and transcriptionally upregulates SPP1 expression, promoting osteosarcoma cell survival and growth under glucose starvation. ChIP assay, luciferase reporter, functional cell survival and growth assays under glucose starvation Biochemical pharmacology Low 38621423
1993 Mouse HoxB9 (Hox-2.5), when expressed in Drosophila under heat shock promoter control, induces homeotic transformations of head toward more posterior identities in larvae and adults, sharing similarities with Abdominal-B phenotypes but also with Antennapedia/HoxB6 transformations, indicating partial functional conservation with Drosophila HOM genes. Ectopic expression in Drosophila (heat shock promoter), phenotypic analysis of homeotic transformations Mechanisms of development Medium 8105876
2007 HOXB9 expression in Hodgkin lymphoma is regulated by E2F3A (activator) and BMI1 (repressor); constitutively active ERK5 signaling mediates HOXB9 expression by repressing BMI1. Knockdown and overexpression assays indicated HOXB9 influences both proliferation and apoptosis in HL cells. RT-PCR, microarray profiling, siRNA knockdown, overexpression, ERK5 pathway analysis Blood Low 17148583
2000 A targeted 90-kb deletion from Hoxb1 to Hoxb9 produces an ordered series of one-segment anterior homeotic transformations along the cervical and thoracic vertebral column; the resulting phenotype is approximately the sum of individual single-gene mutant phenotypes, with no synergistic or novel phenotypes, except loss of ventral curvature at cervicothoracic boundary. Targeted chromosomal deletion in mice, skeletal analysis, neurofilament staining Developmental biology High 10885747

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1989 Segmental expression of Hox-2 homoeobox-containing genes in the developing mouse hindbrain. Nature 515 2571936
2009 WNT/TCF signaling through LEF1 and HOXB9 mediates lung adenocarcinoma metastasis. Cell 502 19576624
1990 Sequential activation of HOX2 homeobox genes by retinoic acid in human embryonal carcinoma cells. Nature 474 1975088
1992 Exogenous retinoic acid rapidly induces anterior ectopic expression of murine Hox-2 genes in vivo. Development (Cambridge, England) 442 1363087
1993 The zinc finger gene Krox20 regulates HoxB2 (Hox2.8) during hindbrain segmentation. Cell 291 8093858
1990 Isolation of the mouse Hox-2.9 gene; analysis of embryonic expression suggests that positional information along the anterior-posterior axis is specified by mesoderm. Development (Cambridge, England) 287 1983472
1991 Introduction of a subtle mutation into the Hox-2.6 locus in embryonic stem cells. Nature 282 1672446
1992 Cell adhesion molecules as targets for Hox genes: neural cell adhesion molecule promoter activity is modulated by cotransfection with Hox-2.5 and -2.4. Proceedings of the National Academy of Sciences of the United States of America 221 1347944
1991 Effects of retinoic acid excess on expression of Hox-2.9 and Krox-20 and on morphological segmentation in the hindbrain of mouse embryos. The EMBO journal 219 1915273
1990 Mouse Hox-2.2 specifies thoracic segmental identity in Drosophila embryos and larvae. Cell 163 1979525
1999 Paralogous mouse Hox genes, Hoxa9, Hoxb9, and Hoxd9, function together to control development of the mammary gland in response to pregnancy. Proceedings of the National Academy of Sciences of the United States of America 156 9892669
1992 Neuroectodermal autonomy of Hox-2.9 expression revealed by rhombomere transpositions. Nature 151 1545869
1988 Characterization of a murine homeo box gene, Hox-2.6, related to the Drosophila Deformed gene. Genes & development 143 2463210
2009 HOXB9, a gene overexpressed in breast cancer, promotes tumorigenicity and lung metastasis. Proceedings of the National Academy of Sciences of the United States of America 139 20080567
1989 Expression of Hox-2.4 homeobox gene directed by proviral insertion in a myeloid leukemia. Nucleic acids research 130 2564662
2000 The leukemia-associated protein Btg1 and the p53-regulated protein Btg2 interact with the homeoprotein Hoxb9 and enhance its transcriptional activation. The Journal of biological chemistry 122 10617598
1992 Analysis of the murine Hox-2.7 gene: conserved alternative transcripts with differential distributions in the nervous system and the potential for shared regulatory regions. The EMBO journal 118 1582411
1990 Hox-2.3 upstream sequences mediate lacZ expression in intermediate mesoderm derivatives of transgenic mice. Development (Cambridge, England) 105 1977573
2016 A miR-192-EGR1-HOXB9 regulatory network controls the angiogenic switch in cancer. Nature communications 104 27041221
1997 Targeted mutations in hoxa-9 and hoxb-9 reveal synergistic interactions. Developmental biology 101 9013929
1991 The murine Hox-2 genes display dynamic dorsoventral patterns of expression during central nervous system development. Development (Cambridge, England) 100 1685115
1991 The expression of murine Hox-2 genes is dependent on the differentiation pathway and displays a collinear sensitivity to retinoic acid in F9 cells and Xenopus embryos. Nucleic acids research 99 1682879
1993 Conditional immortalization of mouse myelomonocytic, megakaryocytic and mast cell progenitors by the Hox-2.4 homeobox gene. The EMBO journal 97 8104786
2017 MiR-215-5p is a tumor suppressor in colorectal cancer targeting EGFR ligand epiregulin and its transcriptional inducer HOXB9. Oncogenesis 81 29199273
1987 Sequence analysis of the murine Hox-2.2, -2.3, and -2.4 homeo boxes: evolutionary and structural comparisons. Genomics 72 2891608
1991 The oncogenic potential of an activated Hox-2.4 homeobox gene in mouse fibroblasts. Molecular and cellular biology 68 1670897
2016 PCAF-mediated acetylation of transcriptional factor HOXB9 suppresses lung adenocarcinoma progression by targeting oncogenic protein JMJD6. Nucleic acids research 66 27613418
1989 Differential expression of human HOX-2 genes along the anterior-posterior axis in embryonic central nervous system. Differentiation; research in biological diversity 64 2570724
1989 The developmental expression pattern of a new murine homeo box gene: Hox-2.5. Developmental biology 61 2567250
1990 Expression of the murine homeobox-containing gene Hox-2.3 suggests multiple time-dependent and tissue-specific roles during development. Development (Cambridge, England) 60 1983116
2000 A large targeted deletion of Hoxb1-Hoxb9 produces a series of single-segment anterior homeotic transformations. Developmental biology 58 10885747
2018 HOXB9 promotes endometrial cancer progression by targeting E2F3. Cell death & disease 57 29724991
2007 Comprehensive analysis of homeobox genes in Hodgkin lymphoma cell lines identifies dysregulated expression of HOXB9 mediated via ERK5 signaling and BMI1. Blood 57 17148583
2015 Kindlin-2 induced by TGF-β signaling promotes pancreatic ductal adenocarcinoma progression through downregulation of transcriptional factor HOXB9. Cancer letters 54 25724625
2015 The role of HOXB9 and miR-196a in head and neck squamous cell carcinoma. PloS one 54 25860510
1992 Ectopic expression of Hox-2.3 induces craniofacial and skeletal malformations in transgenic mice. Mechanisms of development 54 1362649
1993 Colinearity in the Xenopus laevis Hox-2 complex. Mechanisms of development 53 8095151
2012 Homeodomain-containing protein HOXB9 regulates expression of growth and angiogenic factors, facilitates tumor growth in vitro and is overexpressed in breast cancer tissue. The FEBS journal 51 22863320
1993 Cooperative DNA binding of the human HoxB5 (Hox-2.1) protein is under redox regulation in vitro. Molecular and cellular biology 51 8101633
1988 Structure and expression of Hox-2.2, a murine homeobox-containing gene. Proceedings of the National Academy of Sciences of the United States of America 51 2899893
1993 Proximal cis-acting elements cooperate to set Hoxb-7 (Hox-2.3) expression boundaries in transgenic mice. Development (Cambridge, England) 50 8104144
2016 Endocrine disrupting chemical, bisphenol-A, induces breast cancer associated gene HOXB9 expression in vitro and in vivo. Gene 47 27182052
1990 The zebrafish homeobox gene hox-2.2: transcription unit, potential regulatory regions and in situ localization of transcripts. The EMBO journal 47 1968004
1990 A yeast artificial chromosome containing the mouse homeobox cluster Hox-2. Proceedings of the National Academy of Sciences of the United States of America 45 1972280
2012 HOXB9 expression promoting tumor cell proliferation and angiogenesis is associated with clinical outcomes in breast cancer patients. Annals of surgical oncology 43 22396001
1992 Xenopus Hox-2 genes are expressed sequentially after the onset of gastrulation and are differentially inducible by retinoic acid. Development (Cambridge, England) 43 1363722
1992 Dominant mutation of the murine Hox-2.2 gene results in developmental abnormalities. The Journal of experimental zoology 41 1358998
1992 Inhibition of specific pathways of myeloid cell differentiation by an activated Hox-2.4 homeobox gene. Cell growth & differentiation : the molecular biology journal of the American Association for Cancer Research 41 1359901
2022 HOXB9 blocks cell cycle progression to inhibit pancreatic cancer cell proliferation through the DNMT1/RBL2/c-Myc axis. Cancer letters 38 35182659
2018 Regulation of Cancer Stem Cell Self-Renewal by HOXB9 Antagonizes Endoplasmic Reticulum Stress-Induced Melanoma Cell Apoptosis via the miR-765-FOXA2 Axis. The Journal of investigative dermatology 38 29408459
1997 Lineage-specific regulation of hematopoiesis by HOX-B8 (HOX-2.4): inhibition of granulocytic differentiation and potentiation of monocytic differentiation. Blood 38 9292516
1992 Expression of selected human HOX-2 genes in B/T acute lymphoid leukemia and interleukin-2/interleukin-1 beta-stimulated natural killer lymphocytes. Blood 35 1351762
2014 Elevated HOXB9 expression promotes differentiation and predicts a favourable outcome in colon adenocarcinoma patients. British journal of cancer 34 25025961
2014 HOXB9 promotes epithelial-to-mesenchymal transition via transforming growth factor-β1 pathway in hepatocellular carcinoma cells. Clinical and experimental medicine 33 25081022
1991 Expression of the HOX-2.3 homeobox gene in human lymphocytes and lymphoid tissues. Blood 33 1676919
2019 GRP78 activates the Wnt/HOXB9 pathway to promote invasion and metastasis of hepatocellular carcinoma by chaperoning LRP6. Experimental cell research 32 31310747
1986 The murine Hox-2 cluster of homeo box containing genes maps distal on chromosome 11 near the tail-short (Ts) locus. Cytogenetics and cell genetics 32 2875852
1992 DNA sequence polymorphism within hominoid species exceeds the number of phylogenetically informative characters for a HOX2 locus. Molecular biology and evolution 30 1352841
2009 Do HOXB9 and COL1A1 genes play a role in congenital dislocation of the hip? Study in a Caucasian population. Osteoarthritis and cartilage 29 19341834
2015 GalNAc-T14 promotes metastasis through Wnt dependent HOXB9 expression in lung adenocarcinoma. Oncotarget 28 26544896
2011 Histone methylases MLL1 and MLL3 coordinate with estrogen receptors in estrogen-mediated HOXB9 expression. Biochemistry 28 21428455
1993 Functional analysis of the mouse homeobox gene HoxB9 in Drosophila development. Mechanisms of development 27 8105876
2015 HOXB9 induction of mesenchymal-to-epithelial transition in gastric carcinoma is negatively regulated by its hexapeptide motif. Oncotarget 25 26536658
1988 A zebrafish homologue of the murine Hox-2.1 gene. FEBS letters 25 2895022
1987 The mouse Hox2.3 homeobox-containing gene: regulation in differentiating pluripotent stem cells and expression pattern in embryos. Differentiation; research in biological diversity 25 2892751
1988 In situ hybridization patterns of zebrafish homeobox genes homologous to Hox-2.1 and En-2 of mouse. Biochemical and biophysical research communications 24 2904810
2018 HOXB9 acetylation at K27 is responsible for its suppression of colon cancer progression. Cancer letters 22 29654889
2018 HOXB9 inhibits proliferation in gastric carcinoma cells via suppression of phosphorylated-Akt and NF-κB-dependent Snail expression. Digestive and liver disease : official journal of the Italian Society of Gastroenterology and the Italian Association for the Study of the Liver 22 30314948
1991 Characterization of the murine Hox-2.3 promoter: involvement of the transcription factor USF (MLTF). Mechanisms of development 22 1677811
2017 HoxB9 promotes the migration and invasion via TGF-β1/Smad2/Slug signaling pathway in oral squamous cell carcinoma. American journal of translational research 21 28386341
2024 Cold atmospheric plasma enhances SLC7A11-mediated ferroptosis in non-small cell lung cancer by regulating PCAF mediated HOXB9 acetylation. Redox biology 19 39127016
2021 Genome-Wide Epigenetic Landscape of Lung Adenocarcinoma Links HOXB9 DNA Methylation to Intrinsic EGFR-TKI Resistance and Heterogeneous Responses. JCO precision oncology 19 34036228
2014 An E2F1-HOXB9 transcriptional circuit is associated with breast cancer progression. PloS one 18 25136922
1988 The Hox-2 homeo box gene complex on mouse chromosome 11 is closely linked to Re. Genetics 18 2896141
1991 A deletion and a rearrangement distinguish between the intracisternal A-particle of Hox-2.4 and that of interleukin-3 in the same leukemic cells. Virology 17 1673811
1991 Possible role of DNA topoisomerase II on transcription of the homeobox gene Hox-2.1 in F9 embryonal carcinoma cells. Nucleic acids research 17 1683482
2022 HOXB9 Overexpression Promotes Colorectal Cancer Progression and Is Associated with Worse Survival in Liver Resection Patients for Colorectal Liver Metastases. International journal of molecular sciences 16 35216396
2019 Experimental and clinicopathological analysis of HOXB9 in gastric cancer. Oncology letters 14 30867739
2003 [Transmission disequilibrium test for congenital dislocation of the hip and HOXB9 gene or COL1AI gene]. Zhonghua yi xue yi chuan xue za zhi = Zhonghua yixue yichuanxue zazhi = Chinese journal of medical genetics 14 12778441
2023 The E2F1-HOXB9/PBX2-CDK6 axis drives gastric tumorigenesis and serves as a therapeutic target in gastric cancer. The Journal of pathology 13 37272544
2016 Dynamic Pattern of HOXB9 Protein Localization during Oocyte Maturation and Early Embryonic Development in Mammals. PloS one 13 27798681
1993 Induction of HOX-2 genes in P19 embryocarcinoma cells is dependent on retinoic acid receptor alpha. Experimental cell research 13 8097725
2022 The AMPK-HOXB9-KRAS axis regulates lung adenocarcinoma growth in response to cellular energy alterations. Cell reports 12 36001969
2021 miR-557 suppressed the malignant behaviours of osteosarcoma cells by reducing HOXB9 and deactivating the EMT process. Artificial cells, nanomedicine, and biotechnology 12 33666541
2021 A Systematic Review on HOX Genes as Potential Biomarkers in Colorectal Cancer: An Emerging Role of HOXB9. International journal of molecular sciences 12 34948228
2020 HOXB9 enhances the ability of lung cancer cells to penetrate the blood-brain barrier. Aging 12 33411683
2008 Secondary DNA structure formation for Hoxb9 promoter and identification of its specific binding protein. Nucleic acids research 12 18276649
1992 The murine Hox-2.4 promoter contains a functional octamer motif. Nucleic acids research 11 1349742
2022 HOXB9 mediates resistance to chemotherapy and patient outcomes through the TGFβ pathway in pancreatic cancer. Oncotarget 10 35634239
2016 HOXB9 acts as a negative regulator of activated human T cells in response to amino acid deficiency. Immunology and cell biology 10 26926958
2005 Ets identified as a trans-regulatory factor of amphioxus Hox2 by transgenic analysis using ascidian embryos. Developmental biology 10 16051213
1988 Restriction fragment length polymorphism and chromosome mapping of a mouse homeo box gene, Hox-2.1. Biochemical genetics 10 2907403
2023 HOXB9 Overexpression Confers Chemoresistance to Ovarian Cancer Cells by Inducing ERCC-1, MRP-2, and XIAP. International journal of molecular sciences 9 36674764
2023 HOXB9 a miR-122-5p regulated gene, suppressed the anticancer effects of brusatol by upregulating SCD1 expression in melanoma. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 9 37031492
2019 Acetylated HOXB9 at lysine 27 is of differential diagnostic value in patients with pancreatic ductal adenocarcinoma. Frontiers of medicine 7 31372881
2024 HOXB9 promotes osteosarcoma cell survival and malignancy under glucose starvation via upregulating SPP1 expression. Biochemical pharmacology 6 38621423
2022 Activation of CTHRC1 by HOXB9 Promotes Angiogenesis through Fatty Acid Metabolism in Lung Adenocarcinoma. Revista de investigacion clinica; organo del Hospital de Enfermedades de la Nutricion 6 37205792
2020 Silencing of HOXB9 suppresses cellular proliferation, angiogenesis, migration and invasion of prostate cancer cells. Journal of biosciences 6 32098919
1994 The 5'-sequence of the murine Hox-b3 (Hox-2.7) gene and its intron contain multiple transcription-regulatory elements. The International journal of biochemistry 6 7890121
1992 Assignment of the HOX2 and HOX3 gene clusters to the bovine chromosome regions 19q17-qter and 5q14-23. Animal genetics 5 1359814