Matrix metalloproteinase-9 · UniProt P14780
MMP9 (CLG4B) is a secreted, zinc-dependent endopeptidase that remodels the extracellular matrix and processes membrane-bound and secreted protein substrates to regulate tissue remodeling, cell survival, and invasion (PMID:19329763, PMID:39613801). It is produced as a latent proenzyme activated through the cysteine-switch mechanism: oxidation of the prodomain cysteine by HOCl or the MPO/H2O2/Cl- system and by chloramines converts proMMP9 to its active form, while excess oxidant inactivates it (PMID:36009335), and proteolytic activation is also achieved by ADAM15 (PMID:26323669) and by pathogen-derived proteases (PMID:22035412). Active MMP9 controls cell fate by liberating bioactive ligands from precursors: it cleaves membrane SCF to soluble SCF that signals through c-Kit to protect renal epithelium from apoptosis and to support nephron development (PMID:19329763, PMID:19713309), processes pro-BDNF to mature BDNF following stress-induced secretory autophagy (PMID:34330919), sheds Syndecan-4 to desensitize chondrocytes to IL-1β (PMID:33246160), and acts as a PAR1 agonist to drive mesenchymal transition in pancreatic cancer (PMID:32809114). MMP9 expression is transcriptionally tuned by NF-κB, which binds the MMP9 promoter and is antagonized by PPARγ (PMID:33636289), by TGF-β-driven Smad3/Cited2 co-activation (PMID:16619037), and by promoter CpG demethylation (PMID:23417766), while FOXP3 represses it in an LCK-phosphorylation-dependent manner (PMID:24155921). Beyond the extracellular space, nuclear MMP9 in melanoma proteolytically clips the histone H3 N-terminal tail in a p300/CBP-acetylation-dependent manner to activate growth-regulatory genes (PMID:34785776), and active MMP9 interacts with GPX4 and glutathione reductase to suppress antioxidant defenses and drive ferroptosis independently of its collagenase activity (PMID:39252956). MMP9 partners with LCN2 and LOXL2 in a ternary complex and with dysadherin to promote ECM degradation, invasion, and a protumorigenic microenvironment (PMID:37753805, PMID:39613801).
Established the genomic locus of MMP9, providing the foundation for genetic and regulatory studies of the gene.
Evidence Linkage analysis and somatic cell hybrid genotyping mapping CLG4B to 20q11.2-q13.1
Resolved how TGF-β drives MMP9 transcription by identifying Cited2 as a Smad3 co-activator recruited to the MMP9 promoter, linking cytokine signaling to invasion-associated MMP9 induction.
Evidence Co-IP, two-hybrid, GST pulldown, ChIP and luciferase reporter in MDA-MB-231 cells
Demonstrated that MMP9 is not merely an ECM protease but liberates a survival ligand, cleaving membrane SCF to soluble SCF that engages c-Kit to suppress apoptosis in kidney injury and development.
Evidence MMP9 knockout mice, in vitro SCF cleavage assay, recombinant SCF rescue across AKI and organotypic kidney models
Identified converging regulatory inputs controlling MMP9 levels — promoter CpG demethylation, FOXP3 repression gated by LCK phosphorylation, and integrin/SFK signaling — explaining how diverse stimuli set MMP9 expression.
Evidence Bisulfite sequencing and reporter assays, in vitro kinase assay with Y342F mutagenesis, integrin loss-of-function with SFK inhibition
PMID:17762853 PMID:23417766 PMID:24155921
Showed MMP9 acts as a downstream effector of metabolic and proteolytic stress in barrier disruption, including hyperhomocysteinemia-driven BBB permeability and functional interdependency with uPA in tissue remodeling.
Evidence Cbs+/-/Mmp9-/- double knockout with permeability tracers and VE-cadherin readout; MMP9/uPA double knockout phenotyping
Placed MMP9 within signaling cascades as a downstream effector of Ca2+ entry and superoxide, establishing that upstream stress signals converge on MMP9 to cause organ injury.
Evidence TRPV4 knockout and SB-3CT blockade in lung; endothelin-1 allelic series with MMP9 KO and SOD-mimetic rescue in heart
Expanded the MMP9 substrate repertoire to include receptor and proteoglycan signaling — PAR1 agonism driving mesenchymal transition and Syndecan-4 shedding modulating IL-1β responses — and revealed an intracellular role suppressing autophagy.
Evidence PAR1 cleavage and inhibition assays, Sdc4 siRNA/inhibitor shedding assays, CRISPR ablation plus inhibitors with autophagic flux measurement
PMID:32809114 PMID:33064567 PMID:33246160
Discovered a nuclear, chromatin-modifying function for MMP9 — proteolytic clipping of histone H3 N-terminal tails dependent on p300/CBP acetylation — recasting MMP9 as a direct epigenetic regulator of oncogenic gene expression.
Evidence Nuclear localization, H3NT proteolysis and genome-wide assays, p300/CBP co-dependence, RNAi and small-molecule inhibition with tumor growth phenotype in melanoma
Characterized MMP9 secretion via stress-induced secretory autophagy and refined understanding of its activation and domain-dependent surface interactions.
Evidence FKBP51 secretory autophagy assays with in vivo microdialysis and proBDNF cleavage; HOCl/MPO cysteine-switch reconstitution; C9-PEX bispecific inhibitor disrupting MMP9/CD44
PMID:33600567 PMID:34330919 PMID:36009335
Defined non-ECM intracellular functions and protein complexes of MMP9 — driving ferroptosis through GPX4 interaction and antioxidant suppression, and cooperating with dysadherin and the LCN2/LOXL2 complex to remodel the tumor microenvironment.
Evidence LC-MS/MS interactome with collagenase-inactive MMP9 construct and GPX4 assays; dysadherin KO with MMP9 OE rescue in humanized models; co-IP and fractionation of LCN2-LOXL2-MMP9 complex
PMID:37753805 PMID:39252956 PMID:39613801
| Year | Finding | Method | Journal | Conf | PMIDs |
|---|---|---|---|---|---|
| 1995 | CLG4B (MMP9) was chromosomally localized to chromosome 20q11.2-q13.1 using linkage analysis with a polymorphic dinucleotide repeat in its 5' flanking region and somatic cell hybrid analysis. | Linkage analysis, somatic cell hybrid panel genotyping | Annals of human genetics | High | 7762981 8978762 |
| 2009 | MMP9 cleaves membrane-bound stem cell factor (SCF) to release its soluble form (sSCF), which then signals through c-Kit to protect renal tubular epithelial cells (S3 proximal tubule and intercalated cells of collecting duct) from apoptosis during acute kidney injury. MMP9 knockout increased apoptosis and SCF rescue reversed the phenotype. | MMP9 knockout mice, in vivo/in vitro MMP9 cleavage assay for SCF, recombinant SCF rescue, mouse models of folic acid-induced and ischemia-reperfusion AKI | Journal of the American Society of Nephrology : JASN | High | 19329763 |
| 2009 | MMP9 deficiency during embryonic kidney development delays maturation, increases apoptosis in metanephric mesenchyme, reduces nephron number, and impairs ureteric bud branching. MMP9 releases soluble SCF from kidney cells, and recombinant SCF partially rescues the branching and apoptosis defects in MMP9-deficient kidneys. | MMP9 knockout mice, organotypic kidney cultures, SCF secretion assay, recombinant SCF rescue | Journal of the American Society of Nephrology : JASN | High | 19713309 |
| 2006 | Cited2 physically associates with Smad2 and Smad3 (co-IP, two-hybrid, GST pulldown), and is recruited with Smad3 to the MMP9 promoter upon TGF-β stimulation (ChIP), thereby acting as a transcriptional co-activator that enhances TGF-β-mediated MMP9 upregulation and tumor cell invasion. | Co-immunoprecipitation, mammalian two-hybrid, GST pulldown, ChIP, luciferase reporter assay, Cited2 knockdown in MDA-MB-231 cells | Oncogene | High | 16619037 |
| 2007 | Integrin alpha3beta1 cooperates with TGF-β to induce MMP-9 expression in immortalized keratinocytes through a Src family kinase (SFK)-dependent pathway; alpha3beta1 does not regulate TGF-β gene expression, bioavailability, or Smad signaling, indicating a parallel non-Smad pathway. | Integrin alpha3beta1 loss-of-function in immortalized keratinocytes, SFK inhibitor, mRNA/protein measurement | The Journal of investigative dermatology | Medium | 17762853 |
| 2011 | Concomitant deficiency of MMP9 and urokinase-type plasminogen activator (uPA) causes gestational failure, impairs bone growth, and delays cutaneous wound healing in mice beyond that seen with individual knockouts, demonstrating functional interdependency between MMP9 and uPA in tissue remodeling. | MMP9/uPA double knockout mice, wound healing assay, bone growth measurement | Developmental biology | Medium | 21802414 |
| 2013 | MMP9 gene promoter demethylation, specifically at the -36 bp CpG site, correlates with increased MMP9 mRNA expression in TNF-α-stimulated keratinocytes; dual-luciferase reporter assays showed this site controls transcriptional activity. | Bisulfite sequencing PCR, methylation-specific PCR, dual-luciferase reporter assay, TNF-α stimulation | Journal of molecular endocrinology | Medium | 23417766 |
| 2013 | FOXP3 transcription factor suppresses MMP9 expression; LCK phosphorylates FOXP3 at Tyr-342, and a Y342F mutation abrogates FOXP3's ability to suppress MMP9 expression and cell invasion, demonstrating that LCK-mediated phosphorylation of FOXP3 is required for MMP9 repression. | Co-immunoprecipitation, in vitro kinase assay, site-directed mutagenesis (Y342F), invasion assay | PloS one | Medium | 24155921 |
| 2013 | P. gingivalis supernatant (but not its LPS) activates MMP9 to its 82 kDa active form (vs 92 kDa proenzyme) in monocytes in a dose-dependent manner and increases monocyte migration; this migration is completely blocked by the MMP inhibitor GM6001, demonstrating MMP9 activation is mechanistically required for P. gingivalis-induced monocyte migration. | Gelatin zymography, western blot, quantitative PCR, transwell migration assay, pharmacological MMP inhibition | Journal of periodontal research | Medium | 22035412 |
| 2014 | ADAM15 proteolytically cleaves and activates pro-MMP9 in vitro and physically interacts with MMP9 in vivo; ADAM15 also upregulates MMP9 expression via the MEK-ERK pathway in lung cancer cells, and MMP9 knockdown attenuates ADAM15-driven invasion. | In vitro cleavage assay, co-immunoprecipitation, shRNA knockdown, MEK-ERK pathway inhibition, invasion assay | Oncology reports | Medium | 26323669 |
| 2015 | DNA methylation status of the MMP9 gene promoter (unmethylated profile) correlates with higher MMP9 mRNA expression in periapical inflammatory lesions compared with healthy mucosa. | Methylation-specific PCR, restriction enzyme digestion for methylation, qRT-PCR | Journal of endodontics | Low | 26549219 |
| 2015 | MMP-9 (along with MMP-2) is activated downstream of TRPV4-mediated Ca2+ entry in the lung; pharmacological blockade of MMP2/9 with SB-3CT protects against TRPV4 agonist-induced lung injury, and TRPV4-/- mice show no MMP activation, placing MMP9 downstream of TRPV4 Ca2+ signaling in lung barrier disruption. | TRPV4 knockout mice, TRPV4 agonist perfusion, western blot for active MMP isoforms, pharmacological MMP blockade, lung injury markers | American journal of physiology. Lung cellular and molecular physiology | Medium | 25150065 |
| 2015 | Endothelin-1 deficiency causes increased ventricular superoxide, overexpression of MMP9, and reduced ventricular stiffness/dilated cardiomyopathy in mice. A superoxide dismutase mimetic normalized superoxide, reduced MMP9 overexpression, and improved cardiac function; genetic absence of MMP9 also improved cardiac function but did not reduce superoxide, placing MMP9 downstream of superoxide in an endothelin-1 → superoxide → MMP9 → cardiac dysfunction cascade. | Endothelin-1 hypomorphic/hypermorphic mouse alleles, MMP9 knockout mice, superoxide dismutase mimetic treatment, cardiac function measurement | Proceedings of the National Academy of Sciences of the United States of America | High | 25848038 |
| 2020 | Macrophage-secreted MMP9 acts as a PAR1 agonist on pancreatic cancer cells, inducing mesenchymal transition; PAR1 cleavage assay identified MMP9 as the relevant macrophage-secreted PAR1 agonist, and MMP9 or PAR1 inhibition blocked macrophage-driven mesenchymal transition. | PAR1 cleavage assay, medium transfer experiments, MMP9/PAR1 pharmacological inhibition, siRNA knockdown of ZEB1 | Cellular oncology (Dordrecht, Netherlands) | Medium | 32809114 |
| 2020 | MMP-9 inhibition or genetic ablation upregulates autophagic flux in cardiomyocytes and fibroblasts via activation of the AMPKα/Beclin-1/Atg7 pathway and suppression of mTOR, indicating MMP9 suppresses autophagy in the failing heart. | CRISPR/Cas9 MMP9 genetic ablation, pharmacological MMP9 inhibitors (salvianolic acid B, MMP9 inhibitor-I), autophagic flux measurement with bafilomycin A1, western blot of LC3B-II and p62 | American journal of physiology. Heart and circulatory physiology | Medium | 33064567 |
| 2020 | MMP-9 mediates Syndecan-4 (Sdc4) shedding under osteoarthritis conditions; siRNA knockdown and inhibition of MMP-9 (but not MMP-2) decreased shed Sdc4 in vitro, and shed Sdc4 correlated with MMP-9 levels in synovial fluid, desensitizing chondrocytes to IL-1β signaling. | MMP-9 siRNA knockdown, MMP inhibitors, ELISA for shed Sdc4, western blot for IL-1β signaling (pERK/ERK) | Osteoarthritis and cartilage | Medium | 33246160 |
| 2021 | Glucocorticoid-mediated stress enhances secretory autophagy via FKBP51, resulting in MMP9 secretion; secreted MMP9 cleaves pro-BDNF to its mature form (mBDNF), as demonstrated by cellular assays and in vivo microdialysis. | Cellular secretory autophagy assays, in vivo microdialysis, proBDNF cleavage assay, FKBP51 stress response pathway | Nature communications | Medium | 34330919 |
| 2021 | Nuclear MMP-9 in melanoma cells proteolytically clips the histone H3 N-terminal tail (H3NT), selectively activating growth-regulatory genes. This H3NT proteolysis depends on p300/CBP-mediated acetylation of H3K18; RNAi knockdown and small-molecule inhibition of MMP-9 impede melanomagenic gene expression and melanoma tumor growth. | Nuclear MMP9 localization, H3NT proteolysis assay, genome-wide studies, p300/CBP co-dependence assay, RNAi knockdown, small-molecule inhibition | Oncogene | High | 34785776 |
| 2021 | PPARγ activation suppresses MMP9 expression post-intracerebral hemorrhage through downregulation of NF-κB; chromatin immunoprecipitation confirmed NF-κB binding to the MMP9 gene promoter, and co-localization of NF-κB with both PPARγ and MMP9 was demonstrated. | In vivo/in vitro PPARγ agonist (rosiglitazone) and antagonist (GW9662), NF-κB inhibitor (JSH-23), chromatin immunoprecipitation, protein co-immunoprecipitation | Neuroscience letters | Medium | 33636289 |
| 2021 | Simultaneous inhibition of the MMP9 catalytic domain and hemopexin (PEX) domain disrupts the MMP9/CD44 interaction on the cell surface; a multi-specific inhibitor (C9-PEX) reduces MMP9 catalytic activity, MMP9 cellular levels, MMP9 homodimerization, and downstream MAPK/ERK pathway activation in cancer cells. | Yeast surface display protein engineering, fluorescence-activity assays, cell surface binding assays, MAPK/ERK pathway analysis | The Biochemical journal | Medium | 33600567 |
| 2022 | HOCl and a MPO/H2O2/Cl- system activate proMMP9 to active MMP9 via the cysteine-switch mechanism (oxidation of the prodomain cysteine); low nM-μM concentrations of chloramines (from amino acids, serum albumin, ECM proteins, but not HEPES) also activate proMMP9, while high HOCl concentrations inactivate active MMP9. | Fluorescence-based activity assay, gel zymography, MPO enzymatic system, chloramine dose-response, methionine competition assay | Antioxidants (Basel, Switzerland) | High | 36009335 |
| 2023 | Metformin directly binds to MMP-9 (molecular docking and surface plasmon resonance) and attenuates MMP-9 enzymatic activity in overexpressing cells; metformin also drives MMP-9 degradation, reducing plaque MMP-9 and improving atherosclerotic plaque stability in a carotid artery model. | Molecular docking, surface plasmon resonance, zymography, MMP activity assay in HEK293A cells, carotid artery atherosclerosis model | Journal of cardiovascular development and disease | Medium | 36826550 |
| 2023 | LCN2, LOXL2, and MMP9 form a ternary protein complex; LCN2-LOXL2 and LCN2-MMP9 interactions occur both intracellularly and extracellularly, while LOXL2-MMP9 interactions occur only intracellularly. This complex enhances fibronectin/Matrigel degradation, filopodia formation, microfilament rearrangement via profilin 1 upregulation, and activates FAK/AKT/GSK3β signaling. | Protein-protein interaction assays (co-IP), subcellular fractionation, invasion/migration assays, in vivo tumor models | Molecular oncology | Medium | 37753805 |
| 2024 | Active MMP9 interacts with GPX4 and glutathione reductase (identified by LC-MS/MS of 83 MMP9-interacting proteins), reduces GPX4 expression and activity, suppresses key transcription factors (SP1, CREB1, NRF2, FOXO3, ATF4) and ferroptosis-suppressing proteins (GPX1, FSP1), and regulates iron metabolism proteins, thereby driving ferroptosis; an MMP9 construct lacking collagenase activity was used to isolate these non-ECM functions. | LC-MS/MS interactome analysis, MMP9 collagenase-inactive construct, GPX4 activity assay, IPA pathway analysis | iScience | Medium | 39252956 |
| 2024 | Dysadherin directly targets MMP9 (protein-protein interaction), and the dysadherin/MMP9 axis enhances CRC cell invasiveness, ECM proteolytic activity, cancer-associated fibroblast activation, and ECM remodeling. Dysadherin knockout reduces the immunosuppressive and proangiogenic tumor microenvironment; MMP9 overexpression reverses this, confirming MMP9 as the key downstream effector. | Co-IP/protein interaction assay, dysadherin knockout, MMP9 overexpression rescue, humanized mouse model, ECM proteolysis assay | Nature communications | High | 39613801 |
| 2014 | MMP9 ablation in hyperhomocysteinemia mice ameliorates cerebrovascular paracellular permeability and reduces fibrinogen-amyloid beta complex formation; Cbs+/-/Mmp9-/- double knockout mice show restored VE-cadherin expression and improved short-term memory compared to Cbs+/- single knockouts, placing MMP9 downstream of homocysteine in BBB disruption. | Dual-tracer cerebrovascular permeability assay, MMP9 knockout crossed with Cbs+/- mice, immunohistochemistry for VE-cadherin and Fg-Aβ complex, novel object recognition test | Journal of cerebral blood flow and metabolism | Medium | 24865997 |
| Year | Title | Journal | Citations | PMID |
|---|---|---|---|---|
| 2016 | MMP-9 in translation: from molecule to brain physiology, pathology, and therapy. | Journal of neurochemistry | 319 | 26525923 |
| 2013 | Mmp-9 inhibition: a therapeutic strategy in ischemic stroke. | Molecular neurobiology | 244 | 24026771 |
| 2022 | MMP9: A Tough Target for Targeted Therapy for Cancer. | Cancers | 184 | 35406619 |
| 2012 | Matrix metalloproteinases in diabetic retinopathy: potential role of MMP-9. | Expert opinion on investigational drugs | 141 | 22519597 |
| 2002 | MMP9 potentiates pulmonary metastasis formation. | Cancer cell | 139 | 12398887 |
| 2021 | Stress-primed secretory autophagy promotes extracellular BDNF maturation by enhancing MMP9 secretion. | Nature communications | 107 | 34330919 |
| 2016 | MMP-9 signaling in the left ventricle following myocardial infarction. | American journal of physiology. Heart and circulatory physiology | 99 | 27208160 |
| 2014 | Glioma-associated microglial MMP9 expression is upregulated by TLR2 signaling and sensitive to minocycline. | International journal of cancer | 99 | 24752463 |
| 2020 | Elevated MMP9 expression in breast cancer is a predictor of shorter patient survival. | Breast cancer research and treatment | 95 | 32445177 |
| 2006 | Cited2 modulates TGF-beta-mediated upregulation of MMP9. | Oncogene | 93 | 16619037 |
| 2009 | MMP9 and SCF protect from apoptosis in acute kidney injury. | Journal of the American Society of Nephrology : JASN | 87 | 19329763 |
| 2017 | High expression of MMP9 in glioma affects cell proliferation and is associated with patient survival rates. | Oncology letters | 80 | 28454256 |
| 1995 | Characterization of a dinucleotide repeat in the 92 kDa type IV collagenase gene (CLG4B), localization of CLG4B to chromosome 20 and the role of CLG4B in aortic aneurysmal disease. | Annals of human genetics | 66 | 7762981 |
| 2014 | SEMA4b inhibits MMP9 to prevent metastasis of non-small cell lung cancer. | Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine | 65 | 25095981 |
| 2014 | Role of MMP2 and MMP9 in TRPV4-induced lung injury. | American journal of physiology. Lung cellular and molecular physiology | 65 | 25150065 |
| 2021 | Infarct in the Heart: What's MMP-9 Got to Do with It? | Biomolecules | 64 | 33805901 |
| 2014 | DLK1 promotes lung cancer cell invasion through upregulation of MMP9 expression depending on Notch signaling. | PloS one | 63 | 24621612 |
| 2021 | Dendrobine attenuates osteoclast differentiation through modulating ROS/NFATc1/ MMP9 pathway and prevents inflammatory bone destruction. | Phytomedicine : international journal of phytotherapy and phytopharmacology | 59 | 34801352 |
| 2020 | Matrix metalloproteinase 9 (MMP9) limits reactive oxygen species (ROS) accumulation and DNA damage in colitis-associated cancer. | Cell death & disease | 58 | 32943603 |
| 2019 | A circular RNA derived from MMP9 facilitates oral squamous cell carcinoma metastasis through regulation of MMP9 mRNA stability. | Cell transplantation | 56 | 31510782 |
| 2020 | MMP9 inhibition increases autophagic flux in chronic heart failure. | American journal of physiology. Heart and circulatory physiology | 54 | 33064567 |
| 2020 | Role of MMP-9 in epithelial-mesenchymal transition of thyroid cancer. | World journal of surgical oncology | 53 | 32698816 |
| 2019 | MiRNA-218 regulates osteoclast differentiation and inflammation response in periodontitis rats through Mmp9. | Cellular microbiology | 53 | 30444938 |
| 2019 | Macrophage-secreted TSLP and MMP9 promote bleomycin-induced pulmonary fibrosis. | Toxicology and applied pharmacology | 53 | 30653976 |
| 2021 | Tanshinone I, a new EZH2 inhibitor restricts normal and malignant hematopoiesis through upregulation of MMP9 and ABCG2. | Theranostics | 51 | 34093860 |
| 2015 | Expression of MMP-9 and VEGF in meningiomas and their correlation with peritumoral brain edema. | BioMed research international | 51 | 25821815 |
| 2020 | Macrophage-secreted MMP9 induces mesenchymal transition in pancreatic cancer cells via PAR1 activation. | Cellular oncology (Dordrecht, Netherlands) | 50 | 32809114 |
| 2020 | MMP-9 mediated Syndecan-4 shedding correlates with osteoarthritis severity. | Osteoarthritis and cartilage | 50 | 33246160 |
| 2024 | The dysadherin/MMP9 axis modifies the extracellular matrix to accelerate colorectal cancer progression. | Nature communications | 48 | 39613801 |
| 2024 | MMP-9 responsive hydrogel promotes diabetic wound healing by suppressing ferroptosis of endothelial cells. | Bioactive materials | 47 | 39386223 |
| 2015 | Endothelin-1 critically influences cardiac function via superoxide-MMP9 cascade. | Proceedings of the National Academy of Sciences of the United States of America | 47 | 25848038 |
| 2011 | Cyclophilin A promotes human hepatocellular carcinoma cell metastasis via regulation of MMP3 and MMP9. | Molecular and cellular biochemistry | 47 | 21667159 |
| 2017 | Silencing of TGF-β1 in tumor cells impacts MMP-9 in tumor microenvironment. | Scientific reports | 46 | 28819116 |
| 2021 | COL3A1 and MMP9 Serve as Potential Diagnostic Biomarkers of Osteoarthritis and Are Associated With Immune Cell Infiltration. | Frontiers in genetics | 41 | 34512730 |
| 2014 | CRKL promotes lung cancer cell invasion through ERK-MMP9 pathway. | Molecular carcinogenesis | 40 | 24664993 |
| 2014 | Ablation of MMP9 gene ameliorates paracellular permeability and fibrinogen-amyloid beta complex formation during hyperhomocysteinemia. | Journal of cerebral blood flow and metabolism : official journal of the International Society of Cerebral Blood Flow and Metabolism | 39 | 24865997 |
| 2007 | Integrin alpha3beta1 potentiates TGFbeta-mediated induction of MMP-9 in immortalized keratinocytes. | The Journal of investigative dermatology | 39 | 17762853 |
| 2020 | Targeting the MAPK7/MMP9 axis for metastasis in primary bone cancer. | Oncogene | 38 | 32655131 |
| 2012 | MMP-9 and uPAR regulated glioma cell migration. | Cell adhesion & migration | 37 | 23076139 |
| 2018 | CCN6-mediated MMP-9 activation enhances metastatic potential of human chondrosarcoma. | Cell death & disease | 36 | 30237403 |
| 2017 | P2X7 receptor antagonism modulates IL-1β and MMP9 in human atherosclerotic vessels. | Scientific reports | 36 | 28687781 |
| 2016 | HOXC6 promotes gastric cancer cell invasion by upregulating the expression of MMP9. | Molecular medicine reports | 35 | 27573865 |
| 2020 | USP19 Enhances MMP2/MMP9-Mediated Tumorigenesis in Gastric Cancer. | OncoTargets and therapy | 34 | 32904569 |
| 2021 | Elevated expression of MMP8 and MMP9 contributes to diabetic osteoarthritis progression in a rat model. | Journal of orthopaedic surgery and research | 33 | 33468174 |
| 2020 | SETDB1 promotes gastric carcinogenesis and metastasis via upregulation of CCND1 and MMP9 expression. | The Journal of pathology | 33 | 33044755 |
| 2018 | Cigarette smoke-induced RANKL expression enhances MMP-9 production by alveolar macrophages. | International journal of chronic obstructive pulmonary disease | 33 | 30587964 |
| 2009 | MMP9 limits apoptosis and stimulates branching morphogenesis during kidney development. | Journal of the American Society of Nephrology : JASN | 33 | 19713309 |
| 2014 | Anti-inflammatory macrophages activate invasion in pancreatic adenocarcinoma by increasing the MMP9 and ADAM8 expression. | Medical oncology (Northwood, London, England) | 32 | 24526468 |
| 2021 | TPM3 mediates epithelial-mesenchymal transition in esophageal cancer via MMP2/MMP9. | Annals of translational medicine | 31 | 34532475 |
| 2017 | Blockade of MMP-2 and MMP-9 inhibits corneal lymphangiogenesis. | Graefe's archive for clinical and experimental ophthalmology = Albrecht von Graefes Archiv fur klinische und experimentelle Ophthalmologie | 31 | 28669039 |
| 2013 | CD147, MMP9 expression and clinical significance of basal-like breast cancer. | Medical oncology (Northwood, London, England) | 31 | 23292863 |
| 2021 | Muscone derivative ZM-32 inhibits breast tumor angiogenesis by suppressing HuR-mediated VEGF and MMP9 expression. | Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie | 29 | 33450490 |
| 2015 | ADAM15 targets MMP9 activity to promote lung cancer cell invasion. | Oncology reports | 29 | 26323669 |
| 2015 | DNA Methylation of MMP9 Is Associated with High Levels of MMP-9 Messenger RNA in Periapical Inflammatory Lesions. | Journal of endodontics | 29 | 26549219 |
| 2013 | Phosphorylation of FOXP3 by LCK downregulates MMP9 expression and represses cell invasion. | PloS one | 29 | 24155921 |
| 2019 | Induction of ALP and MMP9 activity facilitates invasive behavior in heterogeneous human BMSC and HNSCC 3D spheroids. | FASEB journal : official publication of the Federation of American Societies for Experimental Biology | 28 | 31366234 |
| 2016 | Largescale Transcriptomics Analysis Suggests Over-Expression of BGH3, MMP9 and PDIA3 in Oral Squamous Cell Carcinoma. | PloS one | 28 | 26745629 |
| 2011 | MMP9 expression in oesophageal adenocarcinoma is upregulated with visceral obesity and is associated with poor tumour differentiation. | Molecular carcinogenesis | 28 | 22121096 |
| 2011 | ERK5 regulates invasiveness of osteosarcoma by inducing MMP-9. | Journal of orthopaedic research : official publication of the Orthopaedic Research Society | 28 | 22213073 |
| 2015 | SiRNA directed against annexin II receptor inhibits angiogenesis via suppressing MMP2 and MMP9 expression. | Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology | 27 | 25633185 |
| 2013 | Role of site-specific DNA demethylation in TNFα-induced MMP9 expression in keratinocytes. | Journal of molecular endocrinology | 27 | 23417766 |
| 2020 | L-theanine suppresses the metastasis of prostate cancer by downregulating MMP9 and Snail. | The Journal of nutritional biochemistry | 25 | 33249185 |
| 2017 | Expression of NGF, GDNF and MMP-9 in prostate carcinoma. | Pathology, research and practice | 25 | 28237042 |
| 2018 | MMP2 and MMP9 are Associated with Apical Periodontitis Progression and Might be Modulated by TLR2 and MyD88. | Brazilian dental journal | 24 | 29267523 |
| 2015 | Cadmium exposure inhibits MMP2 and MMP9 activities in the prostate and testis. | Biochemical and biophysical research communications | 24 | 25600809 |
| 2015 | CD151 knockdown inhibits osteosarcoma metastasis through the GSK-3β/β-catenin/MMP9 pathway. | Oncology reports | 24 | 26707073 |
| 2023 | A protein complex of LCN2, LOXL2 and MMP9 facilitates tumour metastasis in oesophageal cancer. | Molecular oncology | 23 | 37753805 |
| 2017 | Sirtuin 6 contributes to migration and invasion of osteosarcoma cells via the ERK1/2/MMP9 pathway. | FEBS open bio | 23 | 28904859 |
| 2015 | Dual-Functions of miR-373 and miR-520c by Differently Regulating the Activities of MMP2 and MMP9. | Journal of cellular physiology | 22 | 25545756 |
| 2013 | Involvement of p38 MAPK in haemozoin-dependent MMP-9 enhancement in human monocytes. | Cell biochemistry and function | 22 | 23468369 |
| 2011 | Porphyromonas gingivalis promotes monocyte migration by activating MMP-9. | Journal of periodontal research | 22 | 22035412 |
| 2022 | Activation and Inhibition of Human Matrix Metalloproteinase-9 (MMP9) by HOCl, Myeloperoxidase and Chloramines. | Antioxidants (Basel, Switzerland) | 21 | 36009335 |
| 2021 | Expression and Potential Role of MMP-9 in Intrauterine Adhesion. | Mediators of inflammation | 21 | 33574731 |
| 2021 | Simultaneous targeting of CD44 and MMP9 catalytic and hemopexin domains as a therapeutic strategy. | The Biochemical journal | 21 | 33600567 |
| 2021 | PPARγ activation suppresses the expression of MMP9 by downregulating NF-κB post intracerebral hemorrhage. | Neuroscience letters | 21 | 33636289 |
| 2021 | Macrophage morphological plasticity and migration is Rac signalling and MMP9 dependant. | Scientific reports | 20 | 33980872 |
| 2020 | Expression of RSK4, CD44 and MMP-9 is upregulated and positively correlated in metastatic ccRCC. | Diagnostic pathology | 20 | 32209138 |
| 2016 | MMP2 and MMP9 associate with crescentic glomerulonephritis. | Clinical kidney journal | 20 | 28584626 |
| 2014 | Increased VEGFR2 and MMP9 protein levels are associated with epithelial dysplasia grading. | Pathology, research and practice | 20 | 25441661 |
| 2022 | Evaluation of the matrix metalloproteinase 9 (MMP9) inhibitor Andecaliximab as an Anti-invasive therapeutic in Head and neck squamous cell carcinoma. | Oral oncology | 19 | 35803110 |
| 2022 | Lactobacilli metabolites restore E-cadherin and suppress MMP9 in cervical cancer cells. | Current research in toxicology | 19 | 36176311 |
| 2021 | MMP-9 Deletion Attenuates Arteriovenous Fistula Neointima through Reduced Perioperative Vascular Inflammation. | International journal of molecular sciences | 19 | 34064140 |
| 2021 | MMP-9 drives the melanomagenic transcription program through histone H3 tail proteolysis. | Oncogene | 19 | 34785776 |
| 2020 | MMP9, CXCR1, TLR6, and MPO participant in the progression of coronary artery disease. | Journal of cellular physiology | 19 | 32052443 |
| 2016 | Epb41l3 suppresses esophageal squamous cell carcinoma invasion and inhibits MMP2 and MMP9 expression. | Cell biochemistry and function | 19 | 26916087 |
| 2011 | Expression of MMP-2 and MMP-9 in retinoblastoma and their significance. | International journal of ophthalmology | 19 | 22553708 |
| 1996 | Reassignment of the 92-kDa type IV collagenase gene (CLG4B) to human chromosome 20. | Cytogenetics and cell genetics | 19 | 8978762 |
| 2021 | Expression and clinical significance of MMP-9 and P53 in lung cancer. | European review for medical and pharmacological sciences | 18 | 33629306 |
| 2011 | Concomitant lack of MMP9 and uPA disturbs physiological tissue remodeling. | Developmental biology | 18 | 21802414 |
| 2021 | MMP-9 Signaling Pathways That Engage Rho GTPases in Brain Plasticity. | Cells | 17 | 33467671 |
| 2020 | Ceramide induces MMP-9 expression through JAK2/STAT3 pathway in airway epithelium. | Lipids in health and disease | 17 | 32829707 |
| 2023 | Fatty acid-binding protein-4 (FABP4) and matrix metalloproteinase-9 (MMP9) as predictive values for nonalcoholic steatohepatitis (NASH). | Lipids in health and disease | 16 | 36609276 |
| 2022 | Exosomes from EGFR-Mutated Adenocarcinoma Induce a Hybrid EMT and MMP9-Dependant Tumor Invasion. | Cancers | 16 | 35954442 |
| 2021 | Berberine Regulated miR150-5p to Inhibit P2X7 Receptor, EMMPRIN and MMP-9 Expression in oxLDL Induced Macrophages. | Frontiers in pharmacology | 16 | 33959010 |
| 2018 | MiR-5692a promotes the invasion and metastasis of hepatocellular carcinoma via MMP9. | European review for medical and pharmacological sciences | 16 | 30070322 |
| 2005 | Expression of RECK gene and MMP-9 in hilar cholangiocarcinoma and its clinical significance. | Journal of Huazhong University of Science and Technology. Medical sciences = Hua zhong ke ji da xue xue bao. Yi xue Ying De wen ban = Huazhong keji daxue xuebao. Yixue Yingdewen ban | 16 | 16463672 |
| 2023 | Metformin Directly Binds to MMP-9 to Improve Plaque Stability. | Journal of cardiovascular development and disease | 15 | 36826550 |
| 2014 | Expression and significance of MMP-9 and MDM2 in the oncogenesis of lung cancer in rats. | Asian Pacific journal of tropical medicine | 15 | 25063292 |
| 2024 | MMP9 drives ferroptosis by regulating GPX4 and iron signaling. | iScience | 14 | 39252956 |
| 2018 | Expression and correlation of MMP-9, VEGF, and p16 in infantile hemangioma. | European review for medical and pharmacological sciences | 14 | 30070315 |
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