Affinage

MYL9

Myosin regulatory light polypeptide 9 · UniProt P24844

Length
172 aa
Mass
19.8 kDa
Annotated
2026-06-10
87 papers in source corpus 26 papers cited in narrative 26 extracted findings
Cross-family judge vs UniProt: tie faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MYL9 (MLC2) is the regulatory light chain of myosin II whose reversible phosphorylation operates as the central activating switch for actomyosin contractility across smooth, cardiac, skeletal, and non-muscle cells, and across processes from muscle contraction to cytokinesis and cell migration (PMID:8351204, PMID:35802750). The activating modification is phosphorylation at the conserved serine (Ser19/Ser15) and an adjacent priming tyrosine: in fast-twitch skeletal fibres MLCK phosphorylates MLC2 in a Ca2+-dependent manner to raise the Ca2+ sensitivity of the contractile apparatus (PMID:8351204), while PKM2-mediated Y118 phosphorylation primes ROCK2 binding and subsequent S15 phosphorylation during cytokinesis (PMID:25412762). Multiple upstream Rho-kinase cascades converge on this switch to control distinct contractile outputs—VE-cadherin/ROCK signalling generates junctional contractility that restrains endothelial sprouting (PMID:19345098), RhoA-driven phosphorylation drives polar body extrusion in oocyte meiosis (PMID:26701676), and a 12(S)-HETE→RHO→ROCK→MYPT→MLC2 cascade drives lymph-endothelial retraction (PMID:27362730). Genetic ablation of Myl9 in mice causes neonatal lethality with smooth-muscle organ failure, establishing its non-redundant requirement in intestinal, bladder, and bronchial smooth muscle (PMID:35802750), a role mirrored by the redundant requirement of its C. elegans ortholog mlc-2 in pharyngeal and body-wall muscle (PMID:9799259). MYL9 expression is set transcriptionally by a network including RUNX1 (PMID:20876458), MAL/SRF and MRTF-A/SMYD3 (PMID:19724058, PMID:24189459), Junb (PMID:20551518), and cardiac GATA/SRF enhancers (PMID:14711876), and its protein output is tuned post-translationally by mutually exclusive Ser19 phosphorylation versus O-GlcNAcylation (PMID:24477671) and by competing N-terminal acetylation versus methylation that partition MYL9 between cytoplasmic/contractile and nuclear/DNA-binding roles (PMID:30242065). In cancer, MYL9 acts as an effector node for traction-force generation, migration, invasion, and EMT control through physical partners and regulators including IQGAP1, MYO19, MR-1, the ITPRIP-DAPK1 complex, and PRPF19-mediated K63 ubiquitination (PMID:37926835, PMID:39437553, PMID:18948272, PMID:34111521, PMID:37031206, PMID:40938994).

Mechanistic history

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

    Established the core biochemical function: that MLCK-mediated MLC2 phosphorylation is the modification that tunes contractile force, answering how light-chain phosphorylation translates into mechanical output.

    Evidence 32P phosphorylation assay and force-Ca2+ measurement in skinned fast-twitch rat skeletal fibres

    PMID:8351204

    Open questions at the time
    • Did not identify the specific phosphorylated residue at sequence level
    • Restricted to fast-twitch skeletal muscle
  2. 1998 High

    Showed the regulatory light chain is genetically required for muscle function and is functionally interchangeable with a paralog, establishing its non-redundant tissue role in an intact organism.

    Evidence Genetic deletion, double-mutant epistasis, and transgenic rescue in C. elegans pharyngeal and body-wall muscle

    PMID:9799259

    Open questions at the time
    • Ortholog system, not human MYL9
    • Did not address phosphoregulation
  3. 2004 High

    Defined the cis-regulatory logic for cardiac-restricted MLC2 expression, answering how the gene is spatially controlled in the heart.

    Evidence Transgenic frog/mouse reporter assays with deletion and site-specific promoter mutagenesis (GATA/SRF/YY1 elements)

    PMID:14711876

    Open questions at the time
    • Concerns Xenopus XMLC2 enhancer; human enhancer conservation not directly tested
    • Does not address non-cardiac expression control
  4. 2006 High

    Placed MLC2-dependent contractility at a spatial point of action—endosome-localized Rho-ROCK-MLC2 signals driving adhesion disassembly—linking contractility to migration mechanics.

    Evidence Live-cell imaging, subcellular localization, ROCK inhibition, Endo180 knockdown

    PMID:17043135

    Open questions at the time
    • Direct MLC2 phosphorylation at endosome sites inferred from pathway, not site-mapped
    • Endo180-MLC2 physical link not established
  5. 2009 High

    Identified upstream transcriptional and signalling inputs—MAL/SRF setting MYL9 expression in megakaryocytes and VE-cadherin/ROCK driving junctional MLC2 contractility—connecting MYL9 to platelet formation and angiogenic restraint.

    Evidence ChIP, luciferase, shRNA rescue (megakaryocytes); organotypic and zebrafish angiogenesis with ROCK/actomyosin inhibition

    PMID:19345098 PMID:19724058

    Open questions at the time
    • MAL/SRF and VE-cadherin pathways studied in separate systems
    • Quantitative contribution of MYL9 vs other SRF targets not isolated in angiogenesis
  6. 2010 High

    Defined RUNX1 and Junb as direct transcriptional regulators of MYL9, linking its expression to cell spreading and vascular smooth-muscle contractility.

    Evidence ChIP/EMSA/reporter with siRNA (RUNX1); conditional knockout with Myl9 re-expression rescue (Junb)

    PMID:20551518 PMID:20876458

    Open questions at the time
    • How these TFs integrate with SRF/MRTF inputs at the MYL9 promoter is unresolved
    • Post-translational consequences downstream not addressed
  7. 2013 Medium

    Resolved a cooperative transcriptional mechanism in which MRTF-A and SMYD3 transactivate MYL9 to drive cancer cell migration, extending SRF-pathway control into tumour cell motility.

    Evidence Reporter/promoter-mutation, Co-IP, siRNA, and migration assays in MCF-7 cells

    PMID:24084383 PMID:24189459

    Open questions at the time
    • Both studies from related work in one cell type
    • SMYD3 methylation substrate at the locus not directly mapped
  8. 2014 High

    Uncovered priming-then-activation phospho-control (PKM2 Y118 priming ROCK2-dependent S15) during cytokinesis and a mutually exclusive Ser19-phosphorylation/O-GlcNAcylation switch in skeletal muscle, refining how the activating modification is gated.

    Evidence In vitro kinase assays, mutagenesis, Co-IP in tumour cells; 2D-gel, phospho/O-GlcNAc immunoblotting and enzyme co-localization in rat muscle

    PMID:24477671 PMID:25412762

    Open questions at the time
    • Y118 priming demonstrated in tumour cytokinesis; generality to other contexts untested
    • Stoichiometry of competing O-GlcNAc vs phosphorylation in vivo not quantified
  9. 2018 High

    Identified N-terminal acetylation versus methylation as competing modifications that partition MYL9 between cytoplasmic contractile and nuclear DNA-binding pools, revealing a non-canonical nuclear role.

    Evidence In vitro NatA/NRMT1 enzymatic assays, modification-exclusive mutants, phosphorylation and DNA-binding readouts

    PMID:30242065

    Open questions at the time
    • Nuclear DNA targets and transcriptional consequences not identified
    • In vivo balance of the two N-terminal modifications unmeasured
  10. 2021 Medium

    Mapped MYL9 into discrete cancer protein complexes—IQGAP1 (CAF cytokine secretion) and an ITPRIP-DAPK1 ternary complex shielding MYL9 from DAPK1 phosphorylation—showing context-specific protein partners modulate its phospho-state and signalling output.

    Evidence Co-IP, ChIP, ELISA, co-culture (IQGAP1/ZEB1); reciprocal Co-IP and differential knockdown (ITPRIP-DAPK1-MYL9)

    PMID:34111521 PMID:37926835

    Open questions at the time
    • Single-lab complex characterizations without structural validation
    • Direct binding interfaces not defined
  11. 2023 Medium

    Established protein-stability and additional binding control of MYL9—PRPF19 K63-ubiquitination stabilizing it to drive Src-YAP1 metastasis, and direct MYO19 binding suppressing EMT—broadening regulation beyond phosphorylation.

    Evidence Co-IP, ubiquitination assays, GST pull-down, and migration/invasion assays in CRC and NSCLC cells

    PMID:37031206 PMID:39437553

    Open questions at the time
    • Ubiquitination site on MYL9 not mapped
    • MYO19 and PRPF19 effects characterized in different cancers without cross-validation
  12. 2025 Medium

    Positioned MYL9 as a downstream effector of mechanotransduction in cancer, where DDX3X-driven derepression elevates MYL9 to remodel F-actin and increase traction forces for metastasis.

    Evidence CRISPR-Cas9 screen, mRNA stability assays, ChIP, traction-force microscopy, orthotopic xenograft

    PMID:40938994

    Open questions at the time
    • Whether traction-force increase requires MYL9 phosphorylation not isolated
    • Single-cancer-type validation

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the multiple input layers—transcriptional networks, competing N-terminal acetylation/methylation, O-GlcNAc/phospho switching, and ubiquitination—are integrated to set the cytoplasmic-versus-nuclear and contractile-versus-migratory balance of MYL9 in a given cell remains unresolved.
  • No unified quantitative model of competing PTMs
  • Nuclear DNA-binding function lacks identified targets
  • No human disease-causing mutation reported in the corpus

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 3 GO:0005198 structural molecule activity 2 GO:0003677 DNA binding 1
Localization
GO:0005856 cytoskeleton 3 GO:0005634 nucleus 1 GO:0005829 cytosol 1
Pathway
R-HSA-1643685 Disease 4 R-HSA-162582 Signal Transduction 3 R-HSA-1640170 Cell Cycle 2 R-HSA-397014 Muscle contraction 2
Partners
DAPK1IQGAP1ITPRIPMR-1MYO19PKM2PRPF19ROCK2
Complex memberships
ITPRIP-DAPK1-MYL9 ternary complexmyosin II

Evidence

Reading pass · 26 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2014 PKM2 phosphorylates MLC2/MYL9 at Y118, which primes binding of ROCK2 to MLC2 and subsequent ROCK2-dependent MLC2 S15 phosphorylation; Aurora B phosphorylates PKM2 at T45 to enable its localization to the contractile ring and interaction with MLC2 during cytokinesis. In vitro kinase assay, Co-IP, site-directed mutagenesis, loss-of-function in tumour cells Nature communications High 25412762
2010 MYL9 is a direct transcriptional target of RUNX1; RUNX1 binds four RUNX1 sites in the MYL9 promoter region (-729/-542 bp) as demonstrated by ChIP, EMSA, and reporter assays; RUNX1 siRNA reduces MYL9 expression and cell spreading on collagen and fibrinogen. Chromatin immunoprecipitation (ChIP), EMSA, reporter gene assay, siRNA knockdown, cell-spreading assay Blood High 20876458
2009 The MAL/SRF transcriptional complex directly regulates MYL9 expression; MAL knockdown in megakaryocyte progenitors reduces MYL9 expression, impairs proplatelet formation and stress fiber/filopodia/lamellipodia formation; shRNA-mediated MYL9 knockdown alone reproduces the proplatelet formation defect. Luciferase reporter assay, chromatin immunoprecipitation, shRNA knockdown, gene expression profiling Blood High 19724058
2010 Conditional ablation of the transcription factor Junb in mice reduces Myl9 expression; re-expression of either Junb or Myl9 in Junb-deficient cells restores stress fiber formation, cellular motility, and contractile capacity of vascular smooth muscle cells, establishing Myl9 as a direct functional target of Junb in actomyosin-based contractility. Conditional knockout mouse, re-expression rescue experiments, arterial contraction assays, stress fiber imaging The Journal of clinical investigation High 20551518
2009 VE-cadherin signals through Rho-kinase to phosphorylate MLC2/MYL9, generating actomyosin contractility at cell junctions that suppresses VEGF-driven, Rac1-dependent sprouting; inhibition of VE-cadherin, Rho-kinase, or actomyosin contractility each leads to increased sprouting. Organotypic angiogenesis assay, zebrafish embryo experiments, pharmacological inhibition of ROCK and actomyosin, VE-cadherin loss-of-function Current biology : CB High 19345098
2006 Endosomes containing Endo180 (CD280) generate localised Rho-ROCK-MLC2-based contractile signals; these endosomes are spatially positioned at sites of adhesion turnover to drive focal adhesion and cell-cell junction disassembly during cell migration. Live-cell imaging, subcellular fractionation/localisation, pharmacological inhibition of ROCK, siRNA knockdown of Endo180 The Journal of cell biology High 17043135
2018 N-terminal acetyltransferase NatA acetylates and N-terminal methyltransferase NRMT1 methylates the alpha-amino group of MYL9; Nα-acetylation promotes cytoplasmic/actomyosin roles of MYL9 (increased pSer19 phosphorylation) while Nα-methylation promotes nuclear roles (increased DNA binding and reduced cytoskeletal interactions). In vitro enzymatic assay with purified NatA and NRMT1, mutant MYL9 constructs exclusive for each modification, cell-based phosphorylation assays, DNA-binding assays The Biochemical journal High 30242065
2013 SMYD3 histone methyltransferase cooperates with MRTF-A to transactivate the MYL9 promoter; SMYD3 HMT activity is required, and the proximal MRTF-A binding element of the MYL9 promoter mediates the cooperative transactivation; overexpression promotes MCF-7 cell migration while SMYD3/MRTF-A siRNA suppresses MYL9 expression and migration. Luciferase reporter assay, mutation analysis of MYL9 promoter, siRNA knockdown, co-immunoprecipitation, migration assay Cancer letters Medium 24189459
2013 MRTF-A transactivates MYL9 and its overexpression promotes migration of MCF-7 breast cancer cells; 17β-estradiol upregulates MRTF-A and thereby MYL9 expression; MRTF-A siRNA suppresses MYL9 transcription and reduces cell migration. Reporter gene assay, siRNA knockdown, migration assay, Western blot Acta biochimica et biophysica Sinica Medium 24084383
2023 PRPF19 E3 ubiquitin ligase stabilises MYL9 via K63-linked ubiquitination; MYL9 is the downstream substrate mediating PRPF19-driven migration and invasion of colorectal cancer cells, and the PRPF19/MYL9 axis activates the Src-YAP1 cascade to promote metastasis. Co-immunoprecipitation, ubiquitination assay, gain- and loss-of-function migration/invasion assays, substrate validation Cell death & disease Medium 37031206
2023 MYL9 binds MYO19 via direct protein-protein interaction (confirmed by Co-IP and GST pull-down); MYL9/MYO19 complex suppresses epithelial-mesenchymal transition and migration in NSCLC cells. Co-immunoprecipitation, GST pull-down assay, scratch wound healing assay, Western blot for EMT markers Physiological genomics Medium 39437553
2022 MYL9 deficiency in mice causes neonatal lethality with distended bladder, shortened small intestine, and alveolar overdistension; LacZ reporter knockin shows MYL9 expression is restricted to muscularis propria of intestine and bladder and bronchial smooth muscle, indicating MYL9 is required for smooth muscle cell function in these organs. Germline knockout mouse, LacZ reporter knock-in for expression mapping, gross and histological phenotyping PloS one High 35802750
2015 RhoA-mediated phosphorylation of MLC2 regulates actin dynamics required for cytokinesis (polar body extrusion) during mouse oocyte meiosis; MLC2 knockdown causes failure of polar body extrusion and arrest at telophase with reduced cortical actin. siRNA knockdown in mouse oocytes, immunofluorescence staining of p-MLC2 and actin, RhoA inhibition Cell cycle (Georgetown, Tex.) Medium 26701676
2021 MYL9 in cancer-associated fibroblasts binds IQGAP1 and through ERK1/2 signalling regulates secretion of CCL2 and TGF-β1, which in turn activate the PI3K-AKT pathway in CRC tumour cells to promote progression; ZEB1 binds the MYL9 promoter in CAFs to enhance MYL9 expression. Co-immunoprecipitation, ChIP, dual luciferase assay, ELISA, siRNA knockdown, co-culture assay Journal of experimental & clinical cancer research : CR Medium 37926835
2021 ITPRIP recruits MYL9 to the kinase domain of DAPK1, forming an ITPRIP-DAPK1-MYL9 ternary complex; this interaction impedes DAPK1-mediated phosphorylation of MYL9 and promotes glioma progression; MYL9 knockdown does not disrupt the ITPRIP-DAPK1 association. Lentivirus co-infection, co-immunoprecipitation, knockdown of ITPRIP/DAPK1/MYL9, in vitro and in vivo tumour growth assays Cellular signalling Medium 34111521
2008 MYL9 interacts with MR-1 (myofibrillogenesis regulator 1); MR-1 knockdown decreases MYL9 phosphorylation and disrupts stress fiber formation, reducing hepatoma cell proliferation, migration, and adhesion via the MLC2/FAK/Akt pathway. siRNA knockdown, Western blot for phospho-MLC2/FAK/Akt, migration/adhesion assay, stress-fiber imaging The Journal of biological chemistry Medium 18948272
2014 Phosphorylation (pSer19) and O-GlcNAcylation of slow MYL9 (sMLC2) are mutually exclusive post-translational modifications in skeletal muscle; a multienzymatic complex including kinase, phosphatase, OGT, and OGA is colocalized at the Z-disc and modulates these modifications; hindlimb unloading (atrophy) decreases O-GlcNAcylation and increases phosphorylation of sMLC2, reversed upon reloading. 2D-gel electrophoresis, phospho-specific dye staining, immunoblotting, co-localization immunofluorescence, rat hindlimb unloading model Pflugers Archiv : European journal of physiology Medium 24477671
1993 Endogenous myosin light chain kinase phosphorylates MLC2/MYL9 at approximately 50% of sites after Ca2+ activation in fast-twitch rat skeletal muscle fibres; MLC2 phosphorylation markedly increases the Ca2+ sensitivity of the contractile apparatus and follows Ca2+ rise with a 1–2 s delay. Mechanically skinned single fibre preparation, 32P-radiolabelled phosphorylation assay, force-Ca2+ relationship measurement Pflugers Archiv : European journal of physiology High 8351204
2004 An 82-bp cardiac-specific enhancer in the Xenopus XMLC2 (MLC2) proximal promoter, composed of two GATA sites and a composite YY1/CArG-like site, is necessary and sufficient for heart-specific expression; the low-affinity SRF site is essential and requires at least one adjacent GATA site; YY1 acts primarily as a repressor of ectopic expression. Transgenic frog embryo reporter assay, deletion and site-specific mutation analysis of promoter elements, transgene expression in mice Development (Cambridge, England) High 14711876
1998 In C. elegans, mlc-2 (regulatory myosin light chain) is required for pharyngeal muscle function; mlc-1 and mlc-2 are redundant in body-wall muscle; transgenic copies of either mlc-1+ or mlc-2+ fully rescue all defects of mlc-1(0) mlc-2(0) double mutants including pharyngeal and body-wall muscle defects. Genetic deletion mutants, double mutant analysis, transgenic rescue, in situ hybridization Genetics High 9799259
2019 Cardiac-specific MLC kinase (cMLCK) localises predominantly at the Z-disc where it interacts with α-actinin2 with high affinity (Kd ~13.4 nM) via the cardiac-isoform-specific N-terminus; a cMLCK mutant deficient for α-actinin2 binding fails to promote sarcomere organisation or increase cardiomyocyte size, indicating Z-disc anchoring is required for cMLCK's role in sarcomere organisation upstream of MLC2/MYL9 phosphorylation. Subcellular fractionation/immunofluorescence localisation, kinetic binding assay, N-terminal deletion and point mutants, cardiomyocyte overexpression Scientific reports Medium 31467300
2016 12(S)-HETE activates a 12-HETE receptor → RHO → ROCK → MYPT → MLC2 signalling cascade in lymph endothelial cells to phosphorylate MLC2, driving actomyosin contraction, LEC retraction, and endothelial barrier breaching; MLC2 siRNA abolishes LEC retraction. siRNA knockdown of MLC2, RHO/ROCK inhibitors, Western blot for p-MLC2, CCID barrier breaching assay British journal of cancer Medium 27362730
2025 DDX3X promotes MYL9 expression in pancreatic cancer by destabilising TLE2 mRNA, releasing KLF4-mediated repression of MYL9; elevated MYL9 remodels F-actin and enhances tumour cell traction forces, facilitating metastasis. CRISPR-Cas9 screen, mRNA stability assays, transcription factor ChIP, traction force microscopy, in vivo orthotopic xenograft Science advances Medium 40938994
2012 Transfection of MYL9 into MYL9-negative Jurkat cells significantly increases surface CD3 expression, establishing that MYL9 enables a distinct pathway for CD3 surface expression independent of the PKC-dependent pathway. MYL9 gene transfection into Jurkat cells, flow cytometry for surface CD3, PMA stimulation Journal of smooth muscle research Medium 23538510
2024 DAPK3, released from LUZP1-mediated suppression by COP1-mediated LUZP1 ubiquitination and degradation, phosphorylates and activates MYL9 to promote EMT and colorectal cancer liver metastasis. Patient-derived organoid multi-omics (WES, bulk and scRNA-seq), Co-IP, in vitro and in vivo functional assays Experimental hematology & oncology Medium 41937206
2024 RUNX1 isoforms B and C differentially regulate MYL9 expression in megakaryocytes/platelets: RUNX1B negatively correlates with MYL9 transcripts in platelets while RUNX1C positively regulates target genes; both isoforms bind and regulate the RUNX1 P1 and P2 promoters. ChIP, luciferase promoter assay, RUNX1 isoform overexpression in HEL and HeLa cells, platelet RNAseq from 85 volunteers bioRxiv (preprint)preprint Medium bio_10.1101_2024.06.18.599563

Source papers

Stage 0 corpus · 87 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2009 VE-Cadherin-mediated cell-cell interaction suppresses sprouting via signaling to MLC2 phosphorylation. Current biology : CB 142 19345098
2016 Sodium Butyrate Promotes Reassembly of Tight Junctions in Caco-2 Monolayers Involving Inhibition of MLCK/MLC2 Pathway and Phosphorylation of PKCβ2. International journal of molecular sciences 135 27735862
2014 PKM2 phosphorylates MLC2 and regulates cytokinesis of tumour cells. Nature communications 125 25412762
2021 Atractylodes oil alleviates diarrhea-predominant irritable bowel syndrome by regulating intestinal inflammation and intestinal barrier via SCF/c-kit and MLCK/MLC2 pathways. Journal of ethnopharmacology 72 33592255
2009 MAL/SRF complex is involved in platelet formation and megakaryocyte migration by regulating MYL9 (MLC2) and MMP9. Blood 66 19724058
2006 Decreased platelet expression of myosin regulatory light chain polypeptide (MYL9) and other genes with platelet dysfunction and CBFA2/RUNX1 mutation: insights from platelet expression profiling. Journal of thrombosis and haemostasis : JTH 65 17059412
2016 MicroRNA-223 Attenuates Hypoxia-induced Vascular Remodeling by Targeting RhoB/MLC2 in Pulmonary Arterial Smooth Muscle Cells. Scientific reports 64 27121304
2010 Regulation of platelet myosin light chain (MYL9) by RUNX1: implications for thrombocytopenia and platelet dysfunction in RUNX1 haplodeficiency. Blood 63 20876458
2009 Drastic increase of myosin light chain MLC-2 in senescent skeletal muscle indicates fast-to-slow fibre transition in sarcopenia of old age. European journal of cell biology 60 19616867
2006 Endosomes generate localized Rho-ROCK-MLC2-based contractile signals via Endo180 to promote adhesion disassembly. The Journal of cell biology 60 17043135
2013 PKC-β exacerbates in vitro brain barrier damage in hyperglycemic settings via regulation of RhoA/Rho-kinase/MLC2 pathway. Journal of cerebral blood flow and metabolism : official journal of the International Society of Cerebral Blood Flow and Metabolism 54 23963366
2013 Histone methyltransferase SMYD3 promotes MRTF-A-mediated transactivation of MYL9 and migration of MCF-7 breast cancer cells. Cancer letters 53 24189459
2016 Colon cancer cell-derived 12(S)-HETE induces the retraction of cancer-associated fibroblast via MLC2, RHO/ROCK and Ca2+ signalling. Cellular and molecular life sciences : CMLS 51 28013338
2010 Junb regulates arterial contraction capacity, cellular contractility, and motility via its target Myl9 in mice. The Journal of clinical investigation 47 20551518
2004 Transcriptional regulation of the cardiac-specific MLC2 gene during Xenopus embryonic development. Development (Cambridge, England) 47 14711876
2016 3-Hydroxyflavone inhibits human osteosarcoma U2OS and 143B cells metastasis by affecting EMT and repressing u-PA/MMP-2 via FAK-Src to MEK/ERK and RhoA/MLC2 pathways and reduces 143B tumor growth in vivo. Food and chemical toxicology : an international journal published for the British Industrial Biological Research Association 44 27600294
2008 MR-1 modulates proliferation and migration of human hepatoma HepG2 cells through myosin light chains-2 (MLC2)/focal adhesion kinase (FAK)/Akt signaling pathway. The Journal of biological chemistry 44 18948272
2018 Homozygous deletion in MYL9 expands the molecular basis of megacystis-microcolon-intestinal hypoperistalsis syndrome. European journal of human genetics : EJHG 43 29453416
2023 MYL9 expressed in cancer-associated fibroblasts regulate the immune microenvironment of colorectal cancer and promotes tumor progression in an autocrine manner. Journal of experimental & clinical cancer research : CR 42 37926835
2021 Novel circular RNA circSOBP governs amoeboid migration through the regulation of the miR-141-3p/MYPT1/p-MLC2 axis in prostate cancer. Clinical and translational medicine 42 33784000
2013 Decreased expression of myosin light chain MYL9 in stroma predicts malignant progression and poor biochemical recurrence-free survival in prostate cancer. Medical oncology (Northwood, London, England) 39 24338276
2019 Long non-coding RNA MBNL1-AS1 regulates proliferation, migration, and invasion of cancer stem cells in colon cancer by interacting with MYL9 via sponging microRNA-412-3p. Clinics and research in hepatology and gastroenterology 38 31255531
2002 Rho kinase mediates serum-induced contraction in fibroblast fibers independent of myosin LC20 phosphorylation. American journal of physiology. Cell physiology 35 12388108
2016 Cancer cell-derived 12(S)-HETE signals via 12-HETE receptor, RHO, ROCK and MLC2 to induce lymph endothelial barrier breaching. British journal of cancer 32 27362730
2011 Dynamic regulation of vascular myosin light chain (MYL9) with injury and aging. PloS one 31 22003410
2018 N-terminal acetylation and methylation differentially affect the function of MYL9. The Biochemical journal 28 30242065
2011 Unusual anchor of a motor complex (MyoD-MLC2) to the plasma membrane of Toxoplasma gondii. Traffic (Copenhagen, Denmark) 28 21143563
2011 Regulation of basal LC20 phosphorylation by MYPT1 and CPI-17 in murine gastric antrum, gastric fundus, and proximal colon smooth muscles. Neurogastroenterology and motility 27 21883701
2010 Androgens induce nongenomic stimulation of colonic contractile activity through induction of calcium sensitization and phosphorylation of LC20 and CPI-17. Molecular endocrinology (Baltimore, Md.) 26 20207835
2024 Kaempferol mitigates sepsis-induced acute lung injury by modulating the SphK1/S1P/S1PR1/MLC2 signaling pathway to restore the integrity of the pulmonary endothelial cell barrier. Chemico-biological interactions 24 38823539
1993 Endogenous MLC2 phosphorylation and Ca(2+)-activated force in mechanically skinned skeletal muscle fibres of the rat. Pflugers Archiv : European journal of physiology 24 8351204
2022 Elevated Myl9 reflects the Myl9-containing microthrombi in SARS-CoV-2-induced lung exudative vasculitis and predicts COVID-19 severity. Proceedings of the National Academy of Sciences of the United States of America 21 35895716
2020 SIRT1-mediated deacetylation of NF-κB inhibits the MLCK/MLC2 pathway and the expression of ET-1, thus alleviating the development of coronary artery spasm. American journal of physiology. Heart and circulatory physiology 21 33095054
1998 Functions of the Caenorhabditis elegans regulatory myosin light chain genes mlc-1 and mlc-2. Genetics 21 9799259
2011 Nogo-B regulates migration and contraction of airway smooth muscle cells by decreasing ARPC 2/3 and increasing MYL-9 expression. Respiratory research 20 21251247
1995 Polyamines inhibit myosin phosphatase and increase LC20 phosphorylation and force in smooth muscle. The American journal of physiology 19 7573385
2016 Effects of CC-chemokine receptor 5 on ROCK2 and P-MLC2 expression after focal cerebral ischaemia-reperfusion injury in rats. Brain injury 16 26983670
2018 Aberrant endoplasmic reticulum stress mediates coronary artery spasm through regulating MLCK/MLC2 pathway. Experimental cell research 15 29378169
2006 Smooth muscle myosin isoform expression and LC20 phosphorylation in innate rat airway hyperresponsiveness. American journal of physiology. Lung cellular and molecular physiology 15 16766577
2023 PRPF19 facilitates colorectal cancer liver metastasis through activation of the Src-YAP1 pathway via K63-linked ubiquitination of MYL9. Cell death & disease 14 37031206
2020 MiR-663a, regulated by lncRNA GAS5, contributes to osteosarcoma development through targeting MYL9. Human & experimental toxicology 14 32633150
2015 RhoA-mediated MLC2 regulates actin dynamics for cytokinesis in meiosis. Cell cycle (Georgetown, Tex.) 14 26701676
2014 Phospho-GlcNAc modulation of slow MLC2 during soleus atrophy through a multienzymatic and sarcomeric complex. Pflugers Archiv : European journal of physiology 14 24477671
2013 Myocardin-related transcription factor A is up-regulated by 17β-estradiol and promotes migration of MCF-7 breast cancer cells via transactivation of MYL9 and CYR61. Acta biochimica et biophysica Sinica 14 24084383
2013 Potential regulation of human muscle plasticity by MLC2 post-translational modifications during bed rest and countermeasures. Archives of biochemistry and biophysics 14 24184274
2022 ROCK1/MLC2 inhibition induces decay of viral mRNA in BPXV infected cells. Scientific reports 13 36280692
2024 An Inulin-Type Fructan CP-A from Codonopsis pilosula Alleviated 5-Fluorouracil-Induced Intestinal Mucositis via the ERK/MLCK/MLC2 Pathway and Regulation of Gut Microbiota. Pharmaceuticals (Basel, Switzerland) 11 38543083
2023 Inhibition of the MLCK/MLC2 pathway protects against intestinal heat stroke-induced injury in rats. Journal of thermal biology 11 37506522
2021 CD4+ T cells in inflammatory diseases: pathogenic T-helper cells and the CD69-Myl9 system. International immunology 11 34427648
2012 Gene duplication and conversion events shaped three homologous, differentially expressed myosin regulatory light chain (MLC2) genes. European journal of cell biology 11 22425609
2023 MYL9 promotes squamous cervical cancer migration and invasion by enhancing aerobic glycolysis. The Journal of international medical research 10 37950670
1992 Effects of myosin kinase inhibiting peptide on contractility and LC20 phosphorylation in skinned smooth muscle. The American journal of physiology 10 1535480
2023 Protective effects of low-intensity pulsed ultrasound (LIPUS) against cerebral ischemic stroke in mice by promoting brain vascular remodeling via the inhibition of ROCK1/p-MLC2 signaling pathway. Cerebral cortex (New York, N.Y. : 1991) 9 37771006
2021 The ROCK inhibitor Y-27632 ameliorates blood-spinal cord barrier disruption by reducing tight junction protein degradation via the MYPT1-MLC2 pathway after spinal cord injury in rats. Brain research 9 34634287
2025 Protection of glutamine: The NF-κB/MLCK/MLC2 signaling pathway mediated by tight junction affects oxidative stress, inflammation and apoptosis in snakehead (Channa argus). Fish & shellfish immunology 8 39826630
2024 The interplay between alterations in esophageal microbiota associated with Th17 immune response and impaired LC20 phosphorylation in achalasia. Journal of gastroenterology 8 38472375
2021 Clinicopathological significance of MYL9 expression in pancreatic ductal adenocarcinoma. Cancer reports (Hoboken, N.J.) 8 34821071
2019 Cardiac MLC2 kinase is localized to the Z-disc and interacts with α-actinin2. Scientific reports 8 31467300
2021 Downregulation of PARVA promotes metastasis by modulating integrin-linked kinase activity and regulating MAPK/ERK and MLC2 signaling in prostate cancer. Translational andrology and urology 7 33718092
2023 LAMB3 Promotes Myofibrogenesis and Cytoskeletal Reorganization in Endometrial Stromal Cells via the RhoA/ROCK1/MYL9 Pathway. Cell biochemistry and biophysics 6 37801199
2022 Oxycodone relieves permeability damage and apoptosis of oxygen-glucose deprivation/reoxygenation-induced brain microvascular endothelial cells through ras homolog family member A (RhoA)/ Rho-associated coiled-coil containing kinases (ROCK)/ myosin light chain 2 (MLC2) signal. Bioengineered 6 35170371
2012 Surface CD3 expression proceeds through both myosin regulatory light chain 9 (MYL9)-dependent and MYL9-independent pathways in Jurkat cells. Journal of smooth muscle research = Nihon Heikatsukin Gakkai kikanshi 6 23538510
2022 Novel role of Dipterocarpus tuberculatus as a stimulator of focal cell adhesion through the regulation of MLC2/FAK/Akt signaling pathway. Cell adhesion & migration 5 35615953
2022 MYL9 deficiency is neonatal lethal in mice due to abnormalities in the lung and the muscularis propria of the bladder and intestine. PloS one 5 35802750
2021 Fermentation characteristics and bacterial dynamics during Chinese sauerkraut fermentation by Lactobacillus curvatus LC-20 under varied salt concentrations reveal its potential in low-salt suan cai production. Journal of bioscience and bioengineering 5 33865692
2024 CT45A1-mediated MLC2 (MYL9) phosphorylation promotes natural killer cell resistance and outer cell fate in a cell-in-cell structure, potentiating the progression of microsatellite instability-high colorectal cancer. Molecular oncology 4 39322998
2021 ITPRIP promotes glioma progression by linking MYL9 to DAPK1 inhibition. Cellular signalling 4 34111521
1992 Polylysine activates smooth muscle actin-myosin interaction without LC20 phosphorylation. The American journal of physiology 4 1535481
2025 Activation of S1PR2 on macrophages and the hepatocyte S1PR2/RhoA/ROCK1/MLC2 pathway in vanishing bile duct syndrome. PloS one 3 39854311
2025 RNA binding protein DDX3X drives pancreatic cancer progression via the TLE2-MYL9 axis. Science advances 3 40938994
2024 Phosphorylated CPI-17 and MLC2 as Biomarkers of Coronary Artery Spasm-Induced Sudden Cardiac Death. International journal of molecular sciences 3 38474189
2024 MYL9 binding with MYO19 suppresses epithelial-mesenchymal transition in non-small-cell lung cancer. Physiological genomics 3 39437553
2023 The autocrine glycosylated-GREM1 interacts with TGFB1 to suppress TGFβ/BMP/SMAD-mediated EMT partially by inhibiting MYL9 transactivation in urinary carcinoma. Cellular oncology (Dordrecht, Netherlands) 2 36920729
2025 The Minute Virus of Canines (MVC) Activates the RhoA/ROCK1/MLC2 Signal Transduction Pathway Resulting in the Dissociation of Tight Junctions and Facilitating Occludin-Mediated Viral Infection. Microorganisms 1 40142587
2025 The Network of Exosomes miRNA and p-MLC2 Regulatory Pathway Induced Pathological Cardiac Hypertrophy in Vasn Deficient Mice. Journal of cellular and molecular medicine 1 41235503
2025 DPSCs-Exos promote OPCs differentiation and white matter repair via Myl9-mediated PRMT5 nucleation after ischemic stroke. Journal of cerebral blood flow and metabolism : official journal of the International Society of Cerebral Blood Flow and Metabolism 1 41272428
2024 Activation of VEGFR3 and MLC2 are Critical for GLP-2 Enhancement of Chylomicron Transport. Gastro hep advances 1 40242171
2023 ZC3H4 governs epithelial cell migration through ROCK/p-PYK2/p-MLC2 pathway in silica-induced pulmonary fibrosis. Environmental toxicology and pharmacology 1 37866415
1995 Transactivation of cardiac MLC-2 promoter by MyoD in 10T1/2 fibroblast cells is independent of E-box requirement but depends upon new proteins that recognize MEF-2 site. Cellular & molecular biology research 1 8589760
2026 Integrative GWAS and RNA-seq identify MYL9 as a key regulator of pullorum disease resistance in chickens. Poultry science 0 41671841
2026 Multi-omics analysis of patient-derived organoids reveals that E3 ligase COP1 promotes liver metastasis and oxaliplatin resistance in colorectal cancer through LUZP1 degradation and MYL9 phosphorylation. Experimental hematology & oncology 0 41937206
2026 Integrated bulk RNA-seq and scRNA-seq identification of a novel "PET-SPI1-MYL9" transcriptional axis in lung adenocarcinoma driven by polyethylene terephthalate exposure. Translational cancer research 0 41969450
2025 MLC2: Physiological Functions and Potential Roles in Tumorigenesis. Cell biochemistry and biophysics 0 40089610
2025 Lnc237 regulates duck myoblasts proliferation and differentiation via the miR-22-5p/MYL9 axis. Poultry science 0 41130043
2024 Ghrelin may protect against vascular endothelial injury in Acute traumatic coagulopathy by mediating the RhoA/ROCK/MLC2 pathway. Journal of thrombosis and thrombolysis 0 39179950
2023 Staphylococcal Enterotoxin C2 Mutant-Induced Antitumor Immune Response Is Controlled by CDC42/MLC2-Mediated Tumor Cell Stiffness. International journal of molecular sciences 0 37511553
1997 [The expression of MLC2-chymase fusion gene in cultured neonatal myocardial cells]. Zhongguo yi xue ke xue yuan xue bao. Acta Academiae Medicinae Sinicae 0 10453502

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