Affinage

Showing ARHGEF12 — LARG is a alias.

ARHGEF12

Rho guanine nucleotide exchange factor 12 · UniProt Q9NZN5

Length
1544 aa
Mass
173.2 kDa
Annotated
2026-06-09
86 papers in source corpus 40 papers cited in narrative 39 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ARHGEF12 (LARG) is a RhoA-subfamily-specific guanine nucleotide exchange factor that couples diverse upstream receptors and mechanical inputs to RhoA-dependent cytoskeletal and contractile programs (PMID:11094164, PMID:21454492). Its DH domain is specific for RhoA, RhoB, and RhoC and catalyzes nucleotide exchange with the highest reported activity among Dbl-family GEFs, with an N-terminal regulatory region of the DH domain mediating association with GDP-bound RhoA and the PH domain contributing regulatory protein interactions near the membrane rather than catalysis itself (PMID:19560536, PMID:21454492). LARG operates as an integral effector linking Gα12/13-coupled GPCRs to Rho signaling, and combined genetic loss of LARG with related RGS-RhoGEFs abolishes Gα12/13-to-Rho signaling, causing embryonic lethality (PMID:11094164, PMID:23467409). It is additionally activated by Gαq-coupled receptors, by Plexin-B/semaphorin receptors through a PDZ-domain interaction with the Plexin C-terminus, and by integrin-borne mechanical force via the Src-family kinase Fyn, the last establishing LARG as the principal mediator of force-driven RhoA activation and cellular stiffening in adhesion complexes (PMID:12183458, PMID:12196628, PMID:21572419, PMID:22100544, PMID:29208526). LARG activity is tuned by multiple layers of regulation: autoinhibitory C-terminal oligomerization (PMID:14712228), stimulatory phosphorylation by RSK2 at Ser1288 that drives membrane translocation (PMID:29279389, PMID:41338458), inhibitory Cdk1 phosphorylation at Ser190/Ser1176 during mitosis (PMID:26483157), CYLD-mediated deubiquitination that enhances exchange activity (PMID:23405219), and a Dia1/RhoA-dependent positive feedback loop in which the Dia1 FH2 domain stimulates LARG GEF activity (PMID:17575049). Through these inputs LARG coordinates vascular smooth muscle contraction and myogenic vasoconstriction (PMID:18084302, PMID:31549965), airway smooth muscle hyperresponsiveness (PMID:30385725), smooth muscle differentiation (PMID:20702813), cell migration and Rac-to-Rho crosstalk (PMID:38102112), cytokinesis abscission at the central spindle and midbody (PMID:23885121), MTOC polarity (PMID:17959834), and erythroid differentiation via a p38 pathway (PMID:31467124). In tight-junction-forming endothelial cells LARG instead selectively activates Rap1A to maintain barrier integrity (PMID:35294066).

Mechanistic history

Synthesis pass · year-by-year structured walk · 35 steps
  1. 2000 High

    Established that LARG is a functional RhoA activator and the molecular link between Gα12-family-coupled GPCRs and Rho signaling, defining its core pathway position.

    Evidence In vivo RhoA activation assay and co-immunoprecipitation in mammalian cells

    PMID:11094164

    Open questions at the time
    • Catalytic specificity and kinetics not yet defined
    • Direct versus indirect Gα12 coupling not resolved at this stage
  2. 2002 High

    Identified the PDZ domain as a receptor-docking module that connects LARG to Plexin-B/semaphorin signaling, showing LARG is recruited by specific receptor tails to activate RhoA.

    Evidence Affinity pulldown from brain, PDZ mutagenesis, and RhoA activation/dominant-negative assays, replicated by two labs

    PMID:12183458 PMID:12196628

    Open questions at the time
    • Structural basis of PDZ recognition not defined
    • Whether Plexin binding alters LARG catalytic state unknown
  3. 2002 Medium

    Showed LARG activity is enhanced by tyrosine phosphorylation downstream of GPCRs, introducing the concept of kinase-driven feedback on LARG.

    Evidence Tyrosine phosphorylation assays, kinase inhibitors, and RhoA-GTP pulldown after thrombin stimulation

    PMID:11799111

    Open questions at the time
    • Phosphosites not mapped
    • Single lab; mechanism of activity enhancement undefined
  4. 2004 Medium

    Revealed autoinhibition via C-terminal oligomerization, explaining how LARG basal activity is restrained.

    Evidence Co-IP, in vitro GEF assay, SRE reporter and focus-formation assays with C-terminal truncations

    PMID:14712228

    Open questions at the time
    • Structural detail of oligomerization interface lacking
    • Physiological trigger for de-oligomerization unknown
  5. 2007 High

    Defined a Dia1/RhoA-dependent positive feedback loop in which the RhoA effector Dia1 directly stimulates LARG exchange activity, amplifying RhoA signaling.

    Evidence In vitro GEF reconstitution, Co-IP, autoinhibition-release and siRNA assays

    PMID:17575049

    Open questions at the time
    • Quantitative contribution of feedback in vivo unclear
    • Whether feedback is cell-type restricted not resolved
  6. 2007 Medium

    Extended LARG function to MTOC polarization and microtubule dynamics downstream of G12/13, linking LARG to centrosomal organization through pericentrin association.

    Evidence Gα12/13-deficient MEFs, siRNA, confocal imaging, pericentrin Co-IP

    PMID:17959834

    Open questions at the time
    • Whether centrosomal RhoA activation is required not shown
    • Direct vs indirect pericentrin interaction unresolved
  7. 2007 High

    Demonstrated physiological necessity of G12/13-LARG signaling in vascular smooth muscle for salt-induced hypertension, distinguishing it from basal blood pressure control.

    Evidence Smooth muscle-specific inducible LARG knockout mice with blood pressure telemetry and salt loading

    PMID:18084302

    Open questions at the time
    • Upstream receptor in salt-induced response not pinpointed
    • Compensation by other RhoGEFs not fully excluded
  8. 2007 Medium

    Placed LARG downstream of fibronectin/integrin adhesion, showing it acts with p115 RhoGEF to drive stress fiber and focal adhesion formation.

    Evidence Active-GEF pulldown, siRNA, actin/focal adhesion staining and RhoA-GTP pulldown

    PMID:17971419

    Open questions at the time
    • Direct adhesion-receptor coupling mechanism undefined at this stage
  9. 2009 Medium

    Connected LARG to axon guidance, showing FAK-dependent tyrosine phosphorylation drives RGMa/Unc5B-induced RhoA activation and growth cone collapse.

    Evidence Co-IP, siRNA, tyrosine phosphorylation and growth cone collapse assays

    PMID:19273616

    Open questions at the time
    • Phosphosites mediating activation not mapped
    • Single lab
  10. 2009 Medium

    Showed mDia1 activates the LARG/RhoA/ROCK axis in neutrophil chemotaxis, reinforcing the Dia1-LARG feedback in a physiological migration context.

    Evidence mDia1-KO neutrophils, RhoA/ROCK assays, chemotaxis assays

    PMID:19265163

    Open questions at the time
    • Direct vs indirect mDia1-LARG coupling in neutrophils not isolated
  11. 2009 High

    Dissected the PH domain's role, showing a conserved hydrophobic patch is needed for cellular RhoA activation through regulatory interactions, not catalysis or membrane targeting per se.

    Evidence PH domain mutagenesis, in vitro exchange assay, reporter assay, membrane-targeting fusion rescue

    PMID:19560536

    Open questions at the time
    • Identity of the regulatory proteins bound by the patch unknown
  12. 2011 High

    Quantified LARG catalytic specificity and power, establishing it as RhoA/B/C-specific with the highest reported Dbl-family GEF activity and an N-terminal DH region engaging GDP-bound RhoA.

    Evidence In vitro kinetic nucleotide-exchange assays with substrate-specificity panel and comparative DH analysis

    PMID:21454492

    Open questions at the time
    • Structural basis of the N-terminal regulatory region not solved here
  13. 2011 High

    Identified Fyn as the upstream kinase activating LARG under integrin tensional force, defining a mechanotransduction pathway distinct from GEF-H1/FAK-Ras-ERK.

    Evidence Magnetic twisting cytometry, GEF activation assays, siRNA, Co-IP and atomic force microscopy

    PMID:21572419

    Open questions at the time
    • Fyn phosphosites on LARG not mapped
    • How force is transduced to Fyn-LARG not fully defined
  14. 2011 Medium

    Showed Gαq-coupled receptors activate LARG via its DH-PH tandem independently of PLCβ, broadening LARG's upstream Gα repertoire beyond G12/13.

    Evidence Co-IP, SRF reporter, dominant-active Gαq, siRNA and receptor pharmacology

    PMID:22100544

    Open questions at the time
    • Direct Gαq-LARG binding interface undefined
    • Single lab
  15. 2012 Medium

    Established NIS as a binding partner that sequesters LARG at the plasma membrane to modulate RhoA-dependent motility and invasion.

    Evidence Co-IP, migration/invasion assays, NIS localization manipulation

    PMID:22962269

    Open questions at the time
    • Direct binding interface not mapped
    • Single lab; physiological relevance limited to specific contexts
  16. 2012 Medium

    Linked LARG to angiotensin II/AT1R-driven vascular contraction through the RhoA/MYPT1 axis, supporting its role in vascular tone.

    Evidence siRNA, RhoA activity and MYPT1 phosphorylation Western blots, isometric contraction

    PMID:23123644

    Open questions at the time
    • Direct receptor-LARG coupling step not shown
  17. 2013 High

    Provided definitive genetic proof that LARG, redundantly with PDZ-RhoGEF and p115, is required for Gα12/13-to-Rho signaling and embryonic development.

    Evidence Single/double/triple knockout mice and MEF signaling assays (RhoA-GTP, JNK/p38)

    PMID:23467409

    Open questions at the time
    • Non-redundant LARG-specific functions in development not isolated here
  18. 2013 Medium

    Identified a cytokinesis function, localizing LARG to the central spindle and midbody and showing it is required specifically for abscission.

    Evidence siRNA, live imaging, immunofluorescence colocalization with tubulin/RhoA, rescue, Aurora-B inhibition

    PMID:23885121

    Open questions at the time
    • Recruitment mechanism to the midbody unknown
    • Relationship to Aurora-B abscission checkpoint not fully defined
  19. 2013 Medium

    Established CYLD as a deubiquitinase that activates LARG, adding ubiquitin removal as a regulatory layer controlling exchange activity and cell polarity.

    Evidence Co-IP, ubiquitination/deubiquitination assays, RhoA-GTP pulldown, cytoskeletal phenotyping

    PMID:23405219

    Open questions at the time
    • Ubiquitin sites on LARG not mapped
    • E3 ligase that ubiquitinates LARG unidentified
  20. 2014 Medium

    Extended LARG mechanotransduction to endothelial ICAM-1, showing it drives RhoA-mediated stiffening required for leukocyte crawling and transendothelial migration.

    Evidence Magnetic twisting cytometry on ICAM-1, siRNA, RhoA pulldown, TEM and traction force assays

    PMID:24585879

    Open questions at the time
    • Direct ICAM-1-LARG coupling not shown
  21. 2014 Medium

    Showed TGF-β/ALK5-driven EMT degrades LARG (and GEF-H1) via the proteasome, reducing stiffness and the mechanical stiffening response while promoting invasion.

    Evidence TGF-β treatment, proteasome inhibition, siRNA, magnetic twisting cytometry, invasion assays

    PMID:25143398

    Open questions at the time
    • E3 ligase mediating LARG degradation unidentified
  22. 2014 Medium

    Demonstrated inflammatory JNK signaling upregulates LARG expression to sustain RhoA/Rho-kinase-driven smooth muscle hypercontraction.

    Evidence siRNA, JNK inhibitor, Western blot, Rho kinase activity and contraction assays

    PMID:24740538

    Open questions at the time
    • Transcriptional mechanism of LARG induction not defined
  23. 2015 High

    Established Cdk1 as a mitotic kinase that phosphorylates LARG at Ser190/Ser1176 to suppress its GEF activity, providing cell-cycle control of RhoA activation.

    Evidence In vitro Cdk1 kinase assay, phosphospecific antibodies, phosphonull/phosphomimetic mutants, RhoA activity assay, immunofluorescence

    PMID:26483157

    Open questions at the time
    • Structural basis of phospho-inhibition unknown
    • Coordination with abscission role not reconciled
  24. 2015 Medium

    Confirmed in vivo that LARG is required for C-RGMa/Rho/ROCK-mediated axon layer targeting in the developing tectum.

    Evidence Dominant-negative LARG in chick tectum via in ovo electroporation and axon projection analysis

    PMID:26292756

    Open questions at the time
    • Dominant-negative approach; endogenous LARG loss-of-function not tested
  25. 2017 High

    Identified RSK2 as a kinase that binds and phosphorylates LARG at Ser1288 to activate RhoA/RhoB-dependent migration and invasion.

    Evidence Co-IP, in vitro kinase assay, RhoA-GTP pulldown, site-directed mutagenesis, migration/invasion assays

    PMID:29279389

    Open questions at the time
    • Membrane translocation mechanism not yet defined (addressed later)
  26. 2017 Medium

    Placed LARG in the DC-SIGN signalosome facilitating HIV-1 internalization in dendritic cells, with cocaine enhancing complex assembly.

    Evidence Co-IP, HIV-1 internalization and trafficking imaging, siRNA

    PMID:28094782

    Open questions at the time
    • Direct DC-SIGN-LARG interaction not isolated
    • Single lab
  27. 2017 Medium

    Established LARG as the primary mediator of mechanical strain-induced RhoA activation in mesenchymal stem cell lineage commitment, restraining adipogenesis.

    Evidence siRNA, RhoA-GTP pulldown, mechanical strain, adipogenesis/osteogenesis assays

    PMID:29208526

    Open questions at the time
    • Upstream strain sensor coupling to LARG not defined
  28. 2018 High

    Demonstrated LARG is required for IL17A-induced airway smooth muscle contractility and airway hyperresponsiveness in vivo, independent of inflammation.

    Evidence Arhgef12-KO mice, tracheal ring contraction, RhoA activation, house dust mite allergen model

    PMID:30385725

    Open questions at the time
    • IL17A receptor-to-LARG coupling mechanism undefined
  29. 2019 High

    Confirmed G12/13-LARG-RhoA signaling is required for pressure-induced myogenic vasoconstriction and systemic vascular resistance.

    Evidence Smooth muscle-specific conditional LARG knockout mice, pressure myography, RhoA activity and Ca2+ measurements

    PMID:31549965

    Open questions at the time
    • Molecular pressure sensor upstream of LARG not identified
  30. 2019 Medium

    Extended LARG function to erythropoiesis, showing it regulates erythroid differentiation through a RhoA/p38 pathway.

    Evidence Zebrafish morpholino/CRISPR KO, p38 assay, RhoA/p38 rescue, K562 and 32D differentiation assays

    PMID:31467124

    Open questions at the time
    • Link between RhoA and p38 in this context not mechanistically detailed
  31. 2022 Medium

    Revealed a context-specific switch in which endothelial LARG selectively activates Rap1A (not Rho GTPases) to maintain tight-junction barrier integrity under TNF challenge.

    Evidence Cell-free GEF assay with immunoprecipitated ArhGEF12, active-GEF pulldown, TEER and tight junction staining

    PMID:35294066

    Open questions at the time
    • Structural determinant of the RhoA-vs-Rap1A switch unknown
    • Single lab; unexpected substrate specificity
  32. 2023 High

    Positioned LARG as the molecular link coupling Rac activity to RhoA activation at cell protrusions/retractions during migration.

    Evidence FRET Rho GTPase biosensors, optogenetic Rac activation, siRNA, live migration analysis

    PMID:38102112

    Open questions at the time
    • Mechanism of Rac-dependent membrane recruitment of LARG not defined
  33. 2025 Medium

    Refined the RSK2-LARG axis by showing Ser1288 phosphorylation drives LARG membrane translocation and LARG-RhoA complex assembly downstream of EGF in glioblastoma.

    Evidence Phosphospecific antibodies, membrane fractionation, Co-IP, RhoA-GTP pulldown, EGF stimulation in patient-derived GBM cells

    PMID:41338458

    Open questions at the time
    • Membrane anchor recognizing phospho-LARG not identified
    • Single lab
  34. 2025 Medium

    Identified Solo as a partner maintaining LARG activity at cell-substrate adhesions for substrate-stiffness-dependent actin remodeling.

    Evidence BioID, Co-IP, siRNA, live imaging, actin polymerization and traction-force assays

    PMID:41739636

    Open questions at the time
    • Direct vs proximity interaction not fully resolved
    • Single lab
  35. 2025 Medium

    Linked an ARHGEF12 E620K mutation to oncogenic Rap1 activation, ITGA6 upregulation, and pro-metastatic exosome-driven fibroblast education in ovarian cancer.

    Evidence Ectopic E620K expression, in vitro migration/invasion/colony assays, xenograft, Rap1 assay, exosome characterization, fibroblast co-culture

    PMID:40860157

    Open questions at the time
    • How E620K shifts substrate preference toward Rap1 not structurally defined
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • The structural basis governing LARG's substrate switch between RhoA-subfamily GTPases and Rap1A, and how the converging regulatory inputs (oligomerization, phosphorylation, ubiquitination, partner binding) are integrated on a single molecule, remain unresolved.
  • No structure of full-length regulated LARG
  • Determinants of RhoA-vs-Rap1A specificity unknown
  • Hierarchy among competing regulatory inputs undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060089 molecular transducer activity 4 GO:0098772 molecular function regulator activity 4 GO:0140096 catalytic activity, acting on a protein 2
Localization
GO:0005856 cytoskeleton 3 GO:0005886 plasma membrane 3 GO:0005815 microtubule organizing center 2 GO:0005829 cytosol 1
Pathway
R-HSA-162582 Signal Transduction 4 R-HSA-1266738 Developmental Biology 3 R-HSA-397014 Muscle contraction 3 R-HSA-1640170 Cell Cycle 2
Partners
CYLDDIAPH1FYNGNA13PCNTPLXNB1RHOARSK2
Complex memberships
DC-SIGN signalosome

Evidence

Reading pass · 39 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2000 LARG activates RhoA in vivo and functions as an integral component linking G protein-coupled receptors and heterotrimeric G proteins of the Gα12 family to Rho-dependent signaling pathways. In vivo RhoA activation assay, co-immunoprecipitation, overexpression in mammalian cells FEBS letters High 11094164
2002 LARG directly interacts with the cytoplasmic tail of Plexin-B1 and Plexin-B2 via its PDZ domain binding to a PDZ-binding motif in the Plexin-B C-terminus; this interaction is required for semaphorin-induced RhoA activation and cytoskeletal reorganization. Affinity pulldown from mouse brain, co-immunoprecipitation, PDZ domain mutagenesis, RhoA activation assay, dominant-negative experiments The Journal of biological chemistry; Proceedings of the National Academy of Sciences High 12183458 12196628
2002 FAK tyrosine-phosphorylates LARG in response to thrombin receptor activation, enhancing RhoA activation; this constitutes a positive feedback loop for sustained Rho activation downstream of GPCRs. Tyrosine phosphorylation assays, kinase inhibitor experiments, co-immunoprecipitation, RhoA-GTP pulldown The Journal of biological chemistry Medium 11799111
2004 LARG forms homo- and hetero-oligomers with PDZ-RhoGEF via their C-terminal regions; deletion of the C-terminal tail increases in vivo RhoA-GEF activity and transforming potential, identifying C-terminal oligomerization as an autoinhibitory mechanism. Co-immunoprecipitation, in vitro GEF assay, serum response element reporter assay, Rho-GTP pulldown, focus formation assay Oncogene Medium 14712228
2007 LARG is required for G12/G13-mediated MTOC polarization and microtubule dynamics; LARG associates with pericentrin and localizes to the MTOC and along microtubule tracks. Gα12/13-deficient MEFs, siRNA knockdown, confocal microscopy, co-immunoprecipitation with pericentrin Molecular biology of the cell Medium 17959834
2007 The RhoA effector Dia1 binds to LARG in a RhoA-dependent manner (releasing Dia1 autoinhibition), and the Dia1 FH2 domain stimulates LARG guanine nucleotide exchange activity in vitro, creating a positive feedback loop for RhoA activation. In vitro GEF assay, co-immunoprecipitation, autoinhibition release assay, siRNA knockdown with RhoA/ROCK signaling readout Genes & development High 17575049
2007 G12-G13-LARG signaling in vascular smooth muscle is required for salt-induced hypertension but not for basal blood pressure regulation, established using smooth muscle-specific inducible Cre/loxP deletion of LARG in mice. Smooth muscle-specific inducible LARG knockout mouse, blood pressure telemetry, salt-loading model Nature medicine High 18084302
2007 LARG is activated downstream of fibronectin adhesion and, together with Lsc/p115 RhoGEF, mediates RhoA activation and formation of stress fibers and focal adhesions in response to fibronectin. Affinity pulldown for active GEFs, siRNA knockdown, actin/focal adhesion staining, RhoA-GTP pulldown Journal of cell science Medium 17971419
2009 LARG associates with Unc5B coreceptor downstream of neogenin/RGMa; FAK-dependent tyrosine phosphorylation of LARG mediates RGMa-induced RhoA activation and growth cone collapse. Co-immunoprecipitation, siRNA knockdown, tyrosine phosphorylation assay, RhoA activation assay, growth cone collapse assay The Journal of cell biology Medium 19273616
2009 mDia1 activates the LARG/RhoA/ROCK signaling axis in neutrophil chemotaxis; mDia1-deficient neutrophils show impaired LARG and RhoA activation, polarization, and directional migration. mDia1 knockout mouse neutrophils, RhoA-GTP pulldown, ROCK activity assay, chemotaxis assay Journal of immunology Medium 19265163
2009 The LARG PH domain contains a conserved hydrophobic patch that is required for RhoA activation and stress fiber induction in cells but not for nucleotide exchange activity in vitro; this patch mediates regulatory protein interactions near the membrane rather than membrane targeting per se. PH domain mutagenesis, in vitro nucleotide exchange assay, transcriptional reporter assay, confocal microscopy, membrane-targeting domain fusion rescue Cellular signalling High 19560536
2010 S1P receptor 2 (S1PR2) activates RhoA in smooth muscle cells specifically through LARG (not PDZ-RhoGEF or p115), and this LARG-RhoA pathway promotes smooth muscle differentiation marker gene expression. siRNA knockdown, RhoA-GTP pulldown, smooth muscle promoter reporter assay, myocardin-related transcription factor nuclear localization, scratch wound/transwell migration assay Arteriosclerosis, thrombosis, and vascular biology Medium 20702813
2011 Application of tensional force to integrins activates LARG through the Src family kinase Fyn, recruiting LARG to adhesion complexes and increasing cellular stiffness (reinforcement); this is mechanistically distinct from GEF-H1 activation via FAK-Ras-ERK. Magnetic twisting cytometry, biochemical GEF activation assays, siRNA knockdown, co-immunoprecipitation, atomic force microscopy Nature cell biology High 21572419
2011 The DH domain of LARG is specific for RhoA, RhoB, and RhoC (inactive toward Rac1, Cdc42, TC10) and exhibits the highest catalytic GEF activity reported for a Dbl protein (~10^7-fold acceleration of nucleotide exchange); the PH domain does not contribute to DH activity in LARG, and a novel N-terminal regulatory region of the DH domain mediates association with GDP-bound RhoA. In vitro kinetic nucleotide exchange assays (fluorescent RhoA reporter), comparative DH domain analysis, substrate specificity panel The Journal of biological chemistry High 21454492
2011 LARG is activated by Gαq-coupled receptors (histamine H1 receptor) through a mechanism requiring only the catalytic DH-PH tandem of LARG, independent of PLCβ activation and distinct from Gαq-mediated p63RhoGEF activation; activated Gαq physically interacts with LARG. Co-immunoprecipitation, SRF-reporter assay, dominant-active Gαq expression, siRNA knockdown, receptor-specific pharmacology Cellular signalling Medium 22100544
2012 NIS (sodium/iodide symporter) binds LARG directly; this interaction sequesters LARG at the plasma membrane, modulating RhoA activity and cell motility/invasiveness. Co-immunoprecipitation, cell migration/invasion assays, NIS subcellular localization manipulation Cancer research Medium 22962269
2012 Angiotensin II activates the LARG/RhoA/MYPT1 axis in vascular smooth muscle cells via the AT1 receptor; siRNA knockdown of LARG reduces Ang II-induced RhoA activity, MYPT1 phosphorylation, and aortic ring contraction. siRNA knockdown, Western blot for RhoA activity and MYPT1 phosphorylation, isometric contraction measurement Acta pharmacologica Sinica Medium 23123644
2013 Combined genetic deletion of PDZ-RhoGEF and LARG in mice causes early embryonic lethality and developmental defects; triple deletion of PDZ-RhoGEF, LARG, and p115 abolishes Gα12/13-to-Rho signaling and thrombin-induced proliferation, directional migration, and JNK/p38 activation in MEFs. Knockout mouse generation, embryonic lethality analysis, MEF signaling assays, RhoA-GTP pulldown, JNK/p38 activation The Journal of biological chemistry High 23467409
2013 LARG localizes to the central spindle and midbody during cytokinesis and is required for abscission; LARG depletion causes persistent intercellular bridges and apoptosis without affecting furrow regression or early mitotic timing. Aurora-B inhibition rescues the delayed abscission kinetics. siRNA knockdown, live cell imaging, immunofluorescence colocalization with α-tubulin and RhoA, rescue expression, Aurora-B inhibition Molecular biology of the cell Medium 23885121
2013 CYLD deubiquitinase interacts with LARG and deubiquitinates it; CYLD-mediated deubiquitination of LARG enhances its ability to stimulate GDP/GTP exchange on RhoA, thereby regulating cytoskeletal rearrangement and cell polarity. Co-immunoprecipitation, ubiquitination assay, deubiquitination assay, RhoA-GTP pulldown, cytoskeletal phenotype analysis PloS one Medium 23405219
2014 LARG acts downstream of mechanically stimulated ICAM-1 clusters in endothelial cells to increase RhoA activity and cellular stiffening; LARG depletion decreases leukocyte crawling and inhibits transendothelial migration. Magnetic twisting cytometry on ICAM-1 clusters, siRNA knockdown, RhoA-GTP pulldown, leukocyte TEM assay, traction force microscopy Journal of immunology Medium 24585879
2014 TGF-β-induced EMT causes proteasomal degradation of LARG and GEF-H1 via ALK5, leading to decreased cell stiffness, loss of stiffening response to integrin force, and increased invasion capacity. TGF-β treatment, proteasome inhibition, siRNA knockdown, magnetic twisting cytometry, Boyden chamber invasion assay, Western blot Molecular biology of the cell Medium 25143398
2014 Jun kinase activity drives overexpression of LARG in response to inflammatory cytokines (IL-1β, TNF-α) in longitudinal smooth muscle cells, resulting in enhanced LARG/RhoA/Rho kinase signaling and sustained hypercontraction. siRNA, JNK inhibitor (SP600125), Western blot, Rho kinase activity assay, smooth muscle contraction assay American journal of physiology. Cell physiology Medium 24740538
2015 Cdk1 directly phosphorylates LARG at Ser190 and Ser1176 during mitosis; phosphonull LARG is more active than phosphomimetic LARG in RhoA activation assays, indicating Cdk1 phosphorylation suppresses LARG GEF activity during mitosis. Phosphorylated LARG localizes to mitotic organizing centers and flanking the midbody. In vitro Cdk1 kinase assay, phosphospecific antibodies, phosphonull/phosphomimetic mutants, RhoA activity assay, immunofluorescence Cellular signalling High 26483157
2015 C-RGMa peptide uses a LARG/Rho/ROCK pathway to inhibit axonal growth; dominant-negative LARG (LARG-PDZ) overexpression in the developing optic tectum causes overshoots in superficial tectal layers, establishing LARG as required for C-RGMa-mediated layer targeting. Dominant-negative LARG overexpression in chick embryo tectum, in ovo electroporation, axonal projection analysis Cell death and differentiation Medium 26292756
2017 RSK2 directly interacts with LARG and phosphorylates LARG at Ser1288, activating RhoA/RhoB-dependent cell migration and invasion; RSK2 Thr577 phosphorylation is required for LARG-RhoA pathway activation. Co-immunoprecipitation, in vitro kinase assay, RhoA-GTP pulldown, site-directed mutagenesis, transwell migration/invasion assay Proceedings of the National Academy of Sciences High 29279389
2017 LARG is a component of the DC-SIGN signalosome and participates in DC-SIGN-mediated HIV-1 internalization in dendritic cells; cocaine enhances DC-SIGN/LARG/LSP1 complex formation, activating LARG-RhoA and focal adhesion signaling to facilitate HIV-1 transfer. Co-immunoprecipitation, HIV-1 internalization assay, intracellular trafficking imaging, siRNA knockdown Scientific reports Medium 28094782
2017 LARG knockdown in mesenchymal stem cells accelerates adipogenesis, reduces basal RhoA activity, and almost entirely inhibits mechanical strain-induced RhoA activation, establishing LARG as the primary mediator of force-driven RhoA activation in MSC lineage commitment. siRNA knockdown, RhoA-GTP pulldown, mechanical strain application, adipogenesis/osteogenesis assays (Oil-Red-O, ALP staining, qPCR) Bone Medium 29208526
2018 Arhgef12 is required for IL17A-induced airway smooth muscle contractility and airway hyperresponsiveness; Arhgef12-KO tracheal rings show decreased contractility and RhoA activation in response to IL17A, and Arhgef12-KO mice have reduced airway hyperresponsiveness in a house dust mite model without effects on inflammation. Arhgef12-KO mouse, tracheal ring contraction assay, RhoA activation assay, in vivo allergen sensitization model, airway hyperresponsiveness measurement JCI insight High 30385725
2019 G12/G13- and LARG-mediated RhoA signaling in smooth muscle cells is required for myogenic vasoconstriction; smooth muscle-specific loss of G12/G13 or ARHGEF12 abolishes pressure-induced RhoA activation and myogenic tone in small arteries, reducing systemic vascular resistance. Smooth muscle-specific conditional LARG knockout mouse, pressure myography, RhoA activity assay, intracellular Ca2+ measurement eLife High 31549965
2019 ARHGEF12-RhoA signaling regulates erythroid differentiation via the p38 kinase pathway; arhgef12 knockdown/knockout in zebrafish impairs erythropoiesis, and active RhoA or p38 rescues this defect. Morpholino knockdown and CRISPR/Cas9 KO in zebrafish, p38 kinase assay, rescue with active RhoA/p38, human K562 and mouse 32D cell differentiation assays Haematologica Medium 31467124
2022 In tight junction-forming human dermal microvascular endothelial cells, ArhGEF12 selectively activates Rap1A (not RhoA, RhoB, RhoC, or Rap1B) in cell-free assays; ARHGEF12 depletion exacerbates TNF-induced barrier disruption by reducing GTP-bound Rap1A. siRNA knockdown, trans-endothelial electrical resistance, cell-free GEF assay with immunoprecipitated ArhGEF12, active GEF pulldown, tight junction staining FASEB journal Medium 35294066
2023 Arhgef12 (and Arhgef11) are enriched at transient cell protrusions and retractions, recruited to the plasma membrane by active Rac; their depletion reduces Rac-to-Rho activity crosstalk, cell protrusion-retraction dynamics, and migration distance, placing Arhgef12 as the molecular link coupling Rac activity to RhoA activation during cell migration. FRET-based Rho GTPase activity biosensors, optogenetic Rac activation, siRNA knockdown, live cell migration analysis Nature communications High 38102112
2025 RSK2 phosphorylation of LARG at S1288 promotes membrane translocation of LARG and markedly enhances assembly of the LARG-RhoA complex and GTP loading of RhoA in response to EGF; this pathway is active in glioblastoma cells. Phosphospecific antibodies, membrane fractionation, co-immunoprecipitation, RhoA-GTP pulldown, EGF stimulation, patient-derived GBM cell lines The Journal of biological chemistry Medium 41338458
2025 ARHGEF12 E620K mutation activates Rap1 signaling, upregulates ITGA6 expression, and promotes formation of ITGA6-high exosomes that educate ovarian fibroblasts toward cancer-associated fibroblast phenotypes, facilitating pre-metastatic niche formation. Ectopic expression of E620K mutant, in vitro migration/invasion/colony assays, xenograft OM model, Rap1 signaling assay, exosome characterization, fibroblast co-culture Theranostics Medium 40860157
2025 Solo interacts with LARG and is required to maintain LARG activity at cell-substrate adhesions; LARG is required for Solo-induced actin polymerization and for actin cytoskeletal remodeling in response to substrate stiffness. BioID proximity labeling, co-immunoprecipitation, siRNA knockdown, live cell imaging, actin polymerization assay, traction force on substrates of varying stiffness Molecular biology of the cell Medium 41739636
2006 A Tyr1306Cys variant of LARG has reduced RhoA GEF activity in transfection assays in NIH3T3 cells compared to the Tyr1306 form, demonstrating that this residue influences LARG catalytic function. Transient transfection, RhoA activation assay (reporter/activity) Diabetes Low 16644711
2014 LARG interacts with TELO2 and pericentrin (PCNT) and a subset co-localizes with PCNT at the centrosome; LARG-deficient cells exhibit replication stress signaling defects including supernumerary centrosomes, reduced γH2AX and RPA foci, and reduced Chk1 activation. Yeast 2-hybrid, co-immunoprecipitation, immunofluorescence co-localization, siRNA knockdown, γH2AX/RPA foci assay, Chk1 activation assay Cell cycle Low 25485589
2008 The LARG PDZ domain has degenerate binding specificity for carboxyterminal peptides, binding five diverse biological partners; phage display identifies a broad sequence preference rather than a strict binding motif. Phage display peptide library, fluorescence polarization binding assay, synthetic peptide binding assays Acta biochimica Polonica Low 18542831

Source papers

Stage 0 corpus · 86 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 G12-G13-LARG-mediated signaling in vascular smooth muscle is required for salt-induced hypertension. Nature medicine 581 18084302
2011 The Rho GEFs LARG and GEF-H1 regulate the mechanical response to force on integrins. Nature cell biology 305 21572419
2000 Leukemia-associated Rho guanine nucleotide exchange factor (LARG) links heterotrimeric G proteins of the G(12) family to Rho. FEBS letters 199 11094164
2002 Plexin B regulates Rho through the guanine nucleotide exchange factors leukemia-associated Rho GEF (LARG) and PDZ-RhoGEF. The Journal of biological chemistry 189 12183458
2002 The semaphorin receptor plexin-B1 signals through a direct interaction with the Rho-specific nucleotide exchange factor, LARG. Proceedings of the National Academy of Sciences of the United States of America 151 12196628
2007 Positive feedback between Dia1, LARG, and RhoA regulates cell morphology and invasion. Genes & development 141 17575049
2002 Regulation of G protein-linked guanine nucleotide exchange factors for Rho, PDZ-RhoGEF, and LARG by tyrosine phosphorylation: evidence of a role for focal adhesion kinase. The Journal of biological chemistry 135 11799111
2004 Homo- and hetero-oligomerization of PDZ-RhoGEF, LARG and p115RhoGEF by their C-terminal region regulates their in vivo Rho GEF activity and transforming potential. Oncogene 94 14712228
2007 A novel role for Lsc/p115 RhoGEF and LARG in regulating RhoA activity downstream of adhesion to fibronectin. Journal of cell science 92 17971419
2009 The mDial formin is required for neutrophil polarization, migration, and activation of the LARG/RhoA/ROCK signaling axis during chemotaxis. Journal of immunology (Baltimore, Md. : 1950) 82 19265163
2015 ARHGEF12 influences the risk of glaucoma by increasing intraocular pressure. Human molecular genetics 79 25637523
2009 Unc5B associates with LARG to mediate the action of repulsive guidance molecule. The Journal of cell biology 72 19273616
2011 Mechanistic insights into specificity, activity, and regulatory elements of the regulator of G-protein signaling (RGS)-containing Rho-specific guanine nucleotide exchange factors (GEFs) p115, PDZ-RhoGEF (PRG), and leukemia-associated RhoGEF (LARG). The Journal of biological chemistry 59 21454492
2000 Effects of NO donors and synthase agonists on endothelial cell uptake of L-Arg and superoxide production. American journal of physiology. Cell physiology 58 10644521
2013 PDZ-RhoGEF and LARG are essential for embryonic development and provide a link between thrombin and LPA receptors and Rho activation. The Journal of biological chemistry 56 23467409
2014 The RhoA guanine nucleotide exchange factor, LARG, mediates ICAM-1-dependent mechanotransduction in endothelial cells to stimulate transendothelial migration. Journal of immunology (Baltimore, Md. : 1950) 50 24585879
2014 TGF-β regulates LARG and GEF-H1 during EMT to affect stiffening response to force and cell invasion. Molecular biology of the cell 45 25143398
2010 Sphingosine 1-phosphate receptor 2 signals through leukemia-associated RhoGEF (LARG), to promote smooth muscle cell differentiation. Arteriosclerosis, thrombosis, and vascular biology 45 20702813
2007 LARG and mDia1 link Galpha12/13 to cell polarity and microtubule dynamics. Molecular biology of the cell 45 17959834
2002 Characterization of the expression of PDZ-RhoGEF, LARG and G(alpha)12/G(alpha)13 proteins in the murine nervous system. The European journal of neuroscience 45 12492428
2012 Iodide transporter NIS regulates cancer cell motility and invasiveness by interacting with the Rho guanine nucleotide exchange factor LARG. Cancer research 42 22962269
2017 RSK2 drives cell motility by serine phosphorylation of LARG and activation of Rho GTPases. Proceedings of the National Academy of Sciences of the United States of America 40 29279389
2009 High-throughput screening for small-molecule inhibitors of LARG-stimulated RhoA nucleotide binding via a novel fluorescence polarization assay. Journal of biomolecular screening 40 19196702
2009 LARG at chromosome 11q23 has functional characteristics of a tumor suppressor in human breast and colorectal cancer. Oncogene 32 19734946
2023 Rho GTPase activity crosstalk mediated by Arhgef11 and Arhgef12 coordinates cell protrusion-retraction cycles. Nature communications 29 38102112
2017 LARG GEF and ARHGAP18 orchestrate RhoA activity to control mesenchymal stem cell lineage. Bone 27 29208526
2003 Anti-thrombotic activity of PDR, a newly synthesized L-Arg derivative, on three thrombosis models in rats. Thrombosis research 24 12893027
2014 Automated NMR fragment based screening identified a novel interface blocker to the LARG/RhoA complex. PloS one 22 24505392
2013 Leukemia-associated RhoGEF (LARG) is a novel RhoGEF in cytokinesis and required for the proper completion of abscission. Molecular biology of the cell 22 23885121
2009 A conserved hydrophobic surface of the LARG pleckstrin homology domain is critical for RhoA activation in cells. Cellular signalling 22 19560536
2019 MicroRNA-214-3p Regulates Hypoxia-Mediated Pulmonary Artery Smooth Muscle Cell Proliferation and Migration by Targeting ARHGEF12. Medical science monitor : international medical journal of experimental and clinical research 21 31373336
2019 ARHGEF12 regulates erythropoiesis and is involved in erythroid regeneration after chemotherapy in acute lymphoblastic leukemia patients. Haematologica 21 31467124
2019 Myogenic vasoconstriction requires G12/G13 and LARG to maintain local and systemic vascular resistance. eLife 19 31549965
2013 Modulation of NO and ROS production by AdiNOS transduced vascular cells through supplementation with L-Arg and BH4: implications for gene therapy of restenosis. Atherosclerosis 18 23958248
2011 LARG links histamine-H1-receptor-activated Gq to Rho-GTPase-dependent signaling pathways. Cellular signalling 17 22100544
2010 Photocytotoxicity and DNA cleavage activity of L-arg and L-lys Schiff base oxovanadium(IV) complexes having phenanthroline bases. Dalton transactions (Cambridge, England : 2003) 17 20563340
2015 ϒ-secretase and LARG mediate distinct RGMa activities to control appropriate layer targeting within the optic tectum. Cell death and differentiation 16 26292756
2012 Angiotensin II regulates the LARG/RhoA/MYPT1 axis in rat vascular smooth muscle in vitro. Acta pharmacologica Sinica 16 23123644
2018 Arhgef12 drives IL17A-induced airway contractility and airway hyperresponsiveness in mice. JCI insight 15 30385725
2013 CYLD regulates RhoA activity by modulating LARG ubiquitination. PloS one 14 23405219
2017 Cocaine Enhances DC to T-cell HIV-1 Transmission by Activating DC-SIGN/LARG/LSP1 Complex and Facilitating Infectious Synapse Formation. Scientific reports 13 28094782
2007 Effects of endogenous nitric oxide induced by 5-fluorouracil and L-Arg on liver carcinoma in nude mice. World journal of gastroenterology 13 18069768
2012 Supplement of L-Arg improves protective immunity during early-stage Plasmodium yoelii 17XL infection. Parasite immunology 12 22709481
2006 A functional Tyr1306Cys variant in LARG is associated with increased insulin action in vivo. Diabetes 12 16644711
2003 Characterization of leukemia-associated Rho guanine nucleotide exchange factor (LARG) expression during murine development. Cell and tissue research 12 14513355
2022 Identification of Arhgef12 and Prkci as genetic modifiers of retinal dysplasia in the Crb1rd8 mouse model. PLoS genetics 11 35675330
2021 Therapy-induced Deletion in 11q23 Leading to Fusion of KMT2A With ARHGEF12 and Development of B Lineage Acute Lymphoplastic Leukemia in a Child Treated for Acute Myeloid Leukemia Caused by t(9;11)(p21;q23)/KMT2A-MLLT3. Cancer genomics & proteomics 11 33419897
2021 Regulation of Leukaemia Associated Rho GEF (LARG/ARHGEF12). Small GTPases 11 34304710
2015 Mitotic-dependent phosphorylation of leukemia-associated RhoGEF (LARG) by Cdk1. Cellular signalling 10 26483157
2021 Enzymatic Cβ-H Functionalization of l-Arg and l-Leu in Nonribosomally Derived Peptidyl Natural Products: A Tale of Two Oxidoreductases. Journal of the American Chemical Society 9 34767710
2014 Jun kinase-induced overexpression of leukemia-associated Rho GEF (LARG) mediates sustained hypercontraction of longitudinal smooth muscle in inflammation. American journal of physiology. Cell physiology 9 24740538
2022 Targeting ARHGEF12 promotes neuroblastoma differentiation, MYCN degradation, and reduces tumorigenicity. Cellular oncology (Dordrecht, Netherlands) 8 36520365
2021 MiR-361-5p/abca1 and MiR-196-5p/arhgef12 Axis Involved in γ-Sitosterol Inducing Dual Anti-Proliferative Effects on Bronchial Epithelial Cells of Chronic Obstructive Pulmonary Disease. International journal of chronic obstructive pulmonary disease 8 34675500
2018 Inhibition of aggregation of physically modified rice proteins by isoconcentration of l-Arg and l-Glu. International journal of biological macromolecules 8 30244131
2015 Role of GABAB receptor and L-Arg in GABA-induced vasorelaxation in non-diabetic and streptozotocin-induced diabetic rat vessels. Iranian biomedical journal 8 25864813
2015 CAT-1 as a novel CAM stabilizes endothelial integrity and mediates the protective actions of L-Arg via a NO-independent mechanism. Journal of molecular and cellular cardiology 8 26283571
2022 ArhGEF12 activates Rap1A and not RhoA in human dermal microvascular endothelial cells to reduce tumor necrosis factor-induced leak. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 7 35294066
2021 Identification of KMT2A-ARHGEF12 fusion in a child with a high-grade B-cell lymphoma. Cancer genetics 7 34237703
2019 Reduction of blood pressure elevation by losartan in spontaneously hypertensive rats through suppression of LARG expression in vascular smooth muscle cells. Journal of the Formosan Medical Association = Taiwan yi zhi 7 30962047
2008 Degenerate specificity of PDZ domains from RhoA-specific nucleotide exchange factors PDZRhoGEF and LARG. Acta biochimica Polonica 6 18542831
1998 Effects of L-NAME and L-Arg on arterial blood pressure in normotensive and hypertensive streptozotocin diabetic rats. Acta physiologica, pharmacologica et therapeutica latinoamericana : organo de la Asociacion Latinoamericana de Ciencias Fisiologicas y [de] la Asociacion Latinoamericana de Farmacologia 4 9695876
2024 What is the Origin of the Regioselective C3-Hydroxylation of L-Arg by the Nonheme Iron Enzyme Capreomycin C? Chemistry (Weinheim an der Bergstrasse, Germany) 3 39190221
2014 The leukemia-associated Rho guanine nucleotide exchange factor LARG is required for efficient replication stress signaling. Cell cycle (Georgetown, Tex.) 3 25485589
2025 Comprehensive exome profiling identifies ARHGEF12 mutation as a driver in gastric cancer with ovarian metastasis. Theranostics 2 40860157
2020 Conserved Dynamic Mechanism of Allosteric Response to L-arg in Divergent Bacterial Arginine Repressors. Molecules (Basel, Switzerland) 2 32397647
2018 Prophylactic treatment of L-Arg improves malaria outcomes by regulating host immune responses during Plasmodium yoelii 17XL infection. Experimental parasitology 2 30266573
2012 Interactions of L-Arg with calf thymus DNA using neutral red dye as a fluorescence probe. Spectrochimica acta. Part A, Molecular and biomolecular spectroscopy 2 22842133
1980 Biological studies of Chlorella pyrenoidosa (strain LARG-1) cultures grown under space flight conditions. Life sciences and space research 2 11971285
2025 METTL3 Promotes Cutaneous T-Cell Lymphoma Progression by Regulating ARHGEF12 Expression. International journal of molecular sciences 1 40332203
2025 The effect of L-Arg supplementation on L-Arg/NO metabolic and AMPK/ACC-1 signalling pathways in adipose cells (3T3 L1). Amino acids 1 40764680
2025 Impact of KCl and L-Arg substitution on low salt beef patties: oxidative, textural, flavor, and microbial insights. Food chemistry 1 41092626
2025 Integrative multi-omics and single-cell transcriptomics reveal ARHGEF12 driving chemoresistance in bladder cancer. Hereditas 1 41310897
2025 Correction: Gan et al. METTL3 Promotes Cutaneous T-Cell Lymphoma Progression by Regulating ARHGEF12 Expression. Int. J. Mol. Sci. 2025, 26, 3640. International journal of molecular sciences 1 41373919
2010 [Effect of L-Arg on inflammatory reaction and nuclear factor-kappa B signal pathway in the acute lung injury in rats induced by lipopolysaccaride]. Zhongguo ying yong sheng li xue za zhi = Zhongguo yingyong shenglixue zazhi = Chinese journal of applied physiology 1 20476576
2007 Genetic variants in the leukemia-associated Rho guanine nucleotide exchange factor (ARHGEF12) gene are not associated with T2DM and related parameters in Caucasians (KORA study). European journal of endocrinology 1 17766704
1998 [Changes of nitric oxide synthase in hypoxic pulmonary hypertension and the effect of L-arg and L-NAME on pulmonary circulation]. Zhonghua bing li xue za zhi = Chinese journal of pathology 1 11245006
2026 Solo regulates the localization and activity of LARG for actin cytoskeletal remodeling in cell-substrate adhesions. Molecular biology of the cell 0 41739636
2026 Variant-to-gene mapping identifies ARHGEF12 as a primary open-angle glaucoma effector gene operating within retinal ganglion cells. bioRxiv : the preprint server for biology 0 41929112
2026 Engineering L-Arg@ZIF-8 nanoparticles to modulate the immune-osteogenic-angiogenic microenvironment for accelerated bone fracture healing. Discover nano 0 42068476
2026 Investigating therapeutic response to netarsudil in glaucoma subjects with the ARHGEF12 risk variant. Frontiers in pharmacology 0 42181883
2025 Polymorphic variants of ABCA1, PMM2, and ARHGEF12 genes and the risk of glaucoma in an Iranian population. International journal of ophthalmology 0 40385107
2025 Phosphorylation at S1288 of leukemia associated RhoGEF (LARG/ARHGEF12) induces plasma membrane localization and promotes binding and activation of RhoA. The Journal of biological chemistry 0 41338458
2023 Probing mechanistic questions in the PLP- and O2-dependent l-Arg oxidases. Methods in enzymology 0 37245913
2013 [Effects of L-Arg on expression of PI3K and PKB in liver among low-birth-weight newborn rats]. Zhongguo dang dai er ke za zhi = Chinese journal of contemporary pediatrics 0 23965885
1997 [Effect of L-arg and SNP on pulmonary arterial pressure and vascular structural changes of chronically hypoxic rats]. Zhongguo yi xue ke xue yuan xue bao. Acta Academiae Medicinae Sinicae 0 10453488
1997 [Activation of L-Arg: no pathway in canine brain by the damage from complete cerebral ischemia-reperfusion]. Hunan yi ke da xue xue bao = Hunan yike daxue xuebao = Bulletin of Hunan Medical University 0 9868109

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